SPECIES: Artemisia cana

 INTRODUCTORY

SPECIES: Artemisia cana
photos courtesy of ©Jody Javersak

AUTHORSHIP AND CITATION:
Howard, Janet L. 2002. Artemisia cana. 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/ [].

FEIS ABBREVIATION:
ARTCAN
ARTCANB
ARTCANC
ATCANV

SYNONYMS:
Seriphidium canum (Pursh) Weber [190]
     = Artemisia cana [75,92,191]
A. c. var. viscidula Osterh. [191]
     = A. c. ssp. viscidula [12,92,117]


NRCS PLANT CODE [180]:
ARCA13
ARCAB3
ARCAC5
ARCAV2

COMMON NAMES:
Species names:
    silver sagebrush
    hoary sagebrush
    dwarf sagebrush
Subspecies names:
    Bolander silver sagebrush
    plains silver sagebrush
    mountain silver sagebrush

TAXONOMY:
The scientific name of silver sagebrush is Artemisia cana Pursh (Asteraceae) [57,75,92,191]. Based upon differences in morphology and ploidy level [12,117], some systematists recognize 3 subspecies:

Artemisia cana spp. bolanderi (Gray) G.H. Ward [12,75,92,117]    Bolander silver sagebrush
Artemisia cana ssp. cana   plains silver sagebrush
Artemisia cana ssp. viscidula (Osterh.) Beetle [12,92,117]   mountain silver sagebrush

Within this summary, "silver sagebrush" refers to the species as a whole. Infrataxa are referred to by the 3 common names listed above. Not all authorities recognize infrataxa within silver sagebrush [38,190].

Hybrids: The sagebrush genus is evolving rapidly, and hybridization is an important factor in its evolution [96,97,113,116,188]. Silver sagebrush readily hybridizes with other woody sagebrush species (Artemisia subgenus Tridentatae) including threetip sagebrush (A. tripartita), big sagebrush (A. tridentata) [117,188,190,191], and low sagebrush (A. arbuscula) [12,117,190]. Silver sagebrush mountain big sagebrush (A. t. ssp. vaseyana) and silver sagebrush Lahontan sagebrush (A. a. ssp. longicaulis) hybrids form stable, self-reproducing populations that some authorities classify as distinct taxonomic entities (snowfield sagebrush (A. spiciformis Osterh.) and coaltown sagebrush (A. argilosa Beetle), respectively) [12,114,116,118,163]. Silver sagebrush infrataxa also hybridize with each other, although Bolander silver sagebrush is geographically isolated from the other 2 subspecies and does not do so naturally [12,38,114]. Sagebrush hybrids are important ecologically as well as evolutionarily, often tolerating a broader range of ecological conditions than either parent [116]. Fire effects to silver sagebrush hybrids are discussed in that section of this review.

LIFE FORM:
Shrub

FEDERAL LEGAL STATUS:
No special status

OTHER STATUS:
Silver sagebrush/basin wildrye (Leymus cinereus), silver sagebrush/mountain big sagebrush/Cusick's bluegrass (Poa cusickii), silver sagebrush/Fendler bluegrass (P. fendleriana), and silver sagebrush/Sandberg bluegrass (P. secunda) communities of Nevada are state ranked as SP: having potential for being threatened or vulnerable but needing further reporting or documentation [128].

The Nature Conservancy ranks silver sagebrush communities of the Great Plains as regionally endangered [156].


DISTRIBUTION AND OCCURRENCE

SPECIES: Artemisia cana
GENERAL DISTRIBUTION:
Species: Silver sagebrush is widely distributed across western North America. It occurs from southern British Columbia east to southwestern Manitoba and south to Minnesota, Nebraska, northern New Mexico and Arizona, and southern California [71,92]. Distributed over 53,221 mi2 (137,800 km2) of the western United States, it is 2nd only to big sagebrush in total area occupied by a sagebrush species [12,13]. Silver sagebrush is most common in the northern Great Plains, Rocky Mountain, and Intermountain regions [198]. It is relatively uncommon in the Great Basin [126]. It is rare in Utah, British Columbia, and Manitoba, and absent from Washington [38,92].  Plants database provides maps showing the overall distribution of silver sagebrush and distributions of the subspecies.

Subspecies: Bolander silver sagebrush occurs from north-central Oregon, where it occurs in montane meadows of the Ochoco and Blue mountains and on the eastern slope of the Cascade Range [37], south through mountainous regions and the eastern edge of the Great Basin to Inyo and Tulare counties, California, and east to Humboldt and Washoe counties, Nevada. Bolander silver sagebrush populations are disjunct from the other 2 subspecies [30,71,92,93,112,126].

Plains silver sagebrush occurs from southern British Columbia east to southwestern Manitoba and south to western Nebraska, eastern Colorado, central Wyoming, and south-central Montana [71,92,190]. Most common east of the Continental Divide, it is widely distributed in the northern Great Plains. Common in Saskatchewan, it becomes increasingly sparse to the south except along watercourses and bottomlands, where it may be locally abundant  [90,112]. Distributions of plains and mountain silver sagebrush overlap in eastern Idaho, western Montana, across Wyoming, and in north-central Colorado. Outlying populations occur in west-central Colorado and the eastern Dakotas [71].

Mountain silver sagebrush is most common west of the Continental Divide. It is distributed from central Idaho and western Montana east to central Colorado, south to northeastern New Mexico and north-central Arizona, and west to central Nevada and eastern Oregon [71,92,112,126,190].

ECOSYSTEMS [55]:
FRES17 Elm-ash-cottonwood
FRES20 Douglas-fir
FRES21 Ponderosa pine
FRES29 Sagebrush
FRES30 Desert shrub
FRES34 Chaparral-mountain shrub
FRES36 Mountain grasslands
FRES37 Mountain meadows
FRES38 Plains grasslands

STATES:
AZ CA CO ID MN MT NE
NV NM ND OR SD UT WY

AB BC MB SK

BLM PHYSIOGRAPHIC REGIONS [16]:
2 Cascade Mountains
3 Southern Pacific Border
4 Sierra Mountains
5 Columbia Plateau
6 Upper Basin and Range
7 Lower Basin and Range
8 Northern Rocky Mountains
9 Middle Rocky Mountains
10 Wyoming Basin
11 Southern Rocky Mountains
12 Colorado Plateau
13 Rocky Mountain Piedmont
14 Great Plains
15 Black Hills Uplift
16 Upper Missouri Basin and Broken Lands

KUCHLER [101] PLANT ASSOCIATIONS:
K011 Western ponderosa forest
K012 Douglas-fir forest
K016 Eastern ponderosa forest
K037 Mountain-mahogany-oak scrub
K038 Great Basin sagebrush
K040 Saltbush-greasewood
K050 Fescue-wheatgrass
K051 Wheatgrass-bluegrass
K055 Sagebrush steppe
K056 Wheatgrass-needlegrass shrubsteppe
K063 Foothills prairie
K064 Grama-needlegrass-wheatgrass
K065 Grama-buffalo grass
K066 Wheatgrass-needlegrass
K067 Wheatgrass-bluestem-needlegrass
K068 Wheatgrass-grama-buffalo grass
K069 Bluestem-grama prairie

SAF COVER TYPES [50]:
16 Aspen
63 Cottonwood
217 Aspen
220 Rocky Mountain juniper
222 Black cottonwood-willow
229 Pacific Douglas-fir
235 Cottonwood-willow
237 Interior ponderosa pine
238 Western juniper

SRM (RANGELAND) COVER TYPES [162]:
101 Bluebunch wheatgrass
102 Idaho fescue
103 Green fescue
107 Western juniper/big sagebrush/bluebunch wheatgrass
216 Montane meadows
301 Bluebunch wheatgrass-blue grama
302 Bluebunch wheatgrass-Sandberg bluegrass
303 Bluebunch wheatgrass-western wheatgrass
304 Idaho fescue-bluebunch wheatgrass
305 Idaho fescue-Richardson needlegrass
306 Idaho fescue-slender wheatgrass
307 Idaho fescue-threadleaf sedge
308 Idaho fescue-tufted hairgrass
309 Idaho fescue-western wheatgrass
310 Needle-and-thread-blue grama
311 Rough fescue-bluebunch wheatgrass
312 Rough fescue-Idaho fescue
313 Tufted hairgrass-sedge
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
323 Shrubby cinquefoil-rough fescue
324 Threetip sagebrush-Idaho fescue
401 Basin big sagebrush
402 Mountain big sagebrush
403 Wyoming big sagebrush
404 Threetip sagebrush
405 Black sagebrush
406 Low sagebrush
407 Stiff sagebrush
408 Other sagebrush types
409 Tall forb
411 Aspen woodland
413 Gambel oak
422 Riparian
501 Saltbush-greasewood
604 Bluestem-grama prairie
605 Sandsage prairie
606 Wheatgrass-bluestem-needlegrass
607 Wheatgrass-needlegrass
608 Wheatgrass-grama-needlegrass
609 Wheatgrass-grama
610 Wheatgrass
611 Blue grama-buffalo grass
612 Sagebrush-grass
613 Fescue grassland
614 Crested wheatgrass
615 Wheatgrass-saltgrass-grama
704 Blue grama-western wheatgrass

HABITAT TYPES AND PLANT COMMUNITIES:
Composition of silver sagebrush communities is variable across the species' broad geographical range. The communities are often compositionally and structurally simple, but are sometimes diverse, especially in riparian zones [67,78,127,177,198]. Brief descriptions of common codominants and associates are presented below by subspecies. More detailed descriptions of silver sagebrush-dominated plant communities are available in the publications listed at the end of this section.

Bolander silver sagebrush: There are few descriptions of Bolander silver sagebrush communities in the literature. Bolander silver sagebrush types on the Toiyabe National Forest of west-central Nevada and  east-central California are codominated by Douglas' sedge (Carex douglasii) and/or Baltic rush (Juncus balticus) [110]. Descriptions of Bolander silver sagebrush communities in California are particularly sparse, although a community codominated by mat muhly (Muhlenbergia richardsonis) has been documented in Lassen County [202]. In nearby Plumas County, Bolander silver sagebrush co-occurs with mountain big sagebrush, Nebraska sedge (Carex nebrascensis), and tufted hairgrass (Deschampsia cespitosa) [32]. In central Oregon, Bolander silver sagebrush/mat muhly communities occur on seasonal ponds, expanding and contracting as climate drives water levels. Big sagebrush, Sierra lodgepole pine (Pinus contorta var. murrayana), and quaking aspen (Populus tremuloides) border and may invade the community [40]. In meadows of the Ochoco and Blue mountains, Bolander silver sagebrush often codominates with big sagebrush. Cusick's bluegrass usually dominates the understory; western yarrow (Achillea millefolium) is a constant associate [98]. Bolander silver sagebrush is classified as a major indicator shrub for riparian zones in central Oregon [100].

Plains silver sagebrush: Species diversity tends to be low in plains silver sagebrush communities [78,127,177]. There are few other shrub species; the communities are mostly composed of silver sagebrush, bunchgrasses, and other graminoid associates. Forbs are typically infrequent [78,177] but occasionally important. Nelson [127] reported 20% forb cover in a plains silver sagebrush community in the Badlands of North Dakota. In Saskatchewan, plains silver sagebrush dominates sandhill prairies, and is common in the blue grama-green needlegrass (Bouteloua gracilis-Nassella viridula) association [88]. Plains silver sagebrush dominates mesic mountain steppes in the northwestern portion of its range, often mixed with Wyoming big sagebrush (Artemisia tridentata var. wyomingensis). It merges with and associates in western wheatgrass-Idaho fescue (Pascopyrum smithii-Festuca idahoensis) and bluebunch wheatgrass (Pseudoroegneria spicata)-Idaho fescue mountain grasslands [71] and in black greasewood (Sarcobatus vermiculatus) communities [3]. Plains silver sagebrush-dominated communities are often upslope from black cottonwood (Populus balsamifera ssp. trichocarpa) floodplains throughout the subspecies' distribution [80], and may codominate with willow (Salix) species on meandering drainages [158]. Plains silver sagebrush sometimes dominates understories of open narrowleaf cottonwood (Populus angustifolia) stands [31]. In the shortgrass prairie, plains silver sagebrush communities occur on or upslope from floodplains. Western wheatgrass is the most common codominant; other frequently codominating or associated grasses include green needlegrass, prairie sandreed (Calamovilfa longifolia), and blue grama [21,31,64,127,139,177]. Nelson [127] found blue grama codominated upland sites in the Badlands of North Dakota, while a plains silver sagebrush-western snowberry (Symphoricarpos occidentalis)/western wheatgrass formation occurred on bottomlands. Japanese brome (Bromus japonicus) and cheatgrass (B. tectorum) are fairly constant invaders in plains silver sagebrush communities [177]. Thatcher [176] reported that plains silver sagebrush in eastern Wyoming was the only woody sagebrush species growing in association with basin wildrye.

Mountain silver sagebrush typically occurs in steppe vegetation. Species diversity and community production can be great in mountain silver sagebrush communities, which often border riparian zones and moist mountain meadows. The most common understory dominants in mountain silver sagebrush communities of Wyoming and Montana are tufted hairgrass and Idaho fescue [198]. In the Gros Ventre area of Wyoming, high-elevation (> 7,000 ft. (2,000 m)) mountain silver sagebrush/Idaho fescue communities occur on moraines and landslides [20]. Lower-elevation mountain silver sagebrush communities of Wyoming may be dominated by bluegrasses (Poa spp.) and bromes (Bromus spp.) [143]. Shrubby cinquefoil (Dasiphora floribunda) is a common associate in mountain silver sagebrush communities, and usually the only other shrub present [20,143]. Shrubby cinquefoil is the only constant shrub associate in mountain silver sagebrush/Thurber fescue (F. thurberi) steppes of Colorado [176,178] and in mountain silver sagebrush/Idaho fescue steppes of central and eastern Idaho [155]. Mountain silver sagebrush is locally dominant on mountain grassland clay soils of central Idaho, where it forms the "Camas Prairie" association with common camas (Camassia quamash) [149]. Mountain silver sagebrush steppes in northern Utah and southern Idaho are codominated by tufted hairgrass, sheep fescue (F. ovina), and Kentucky bluegrass (P. pratensis). Mountain silver sagebrush/Kentucky bluegrass is probably a sere induced by heavy grazing [77,131]. Mountain silver sagebrush communities of Utah merge into open quaking aspen-Rocky Mountain Douglas-fir (Pseudotsuga menziesii var. glauca) parklands, where mountain silver sagebrush becomes an understory component [166]. In eastern Oregon, Douglas' sedge and Cusick's bluegrass dominate mountain silver sagebrush understories. The mountain silver sagebrush community merges into upland mountain big sagebrush and streamside Kentucky bluegrass or sedge (Carex spp.) communities [132]. In Nevada, mountain silver sagebrush steppes are codominated by slender wheatgrass (Elymus trachycaulus), Idaho fescue, beardless wildrye (Leymus triticoides), sedges, and Baltic rush [109,110].

Vegetation classifications describing silver sagebrush-dominated communities are listed below:

Silver sagebrush (covering both plains and mountain subspecies):
ID: [61]
MT: [66,150]

Bolander silver sagebrush:
CA: [110,202]
NV: [110]
OR: [98,99,170]

Plains silver sagebrush:
AB: [35]
MT: [62,125]
ND: [62,64,69,78,127]
SD: [64]
WY: [31,65,177]

Mountain silver sagebrush:
CO: [54,175,178]
ID: [77,155,203]
MT: [66]
NV: [109,110]
OR: [132]
UT: [131,142,203]
WY: [20,24,31,143,203]

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Artemisia cana
GENERAL BOTANICAL CHARACTERISTICS:
Silver sagebrush is a native shrub with semiwoody to woody stems [92]. Mature plants generally range from 1.3 to 3.3 feet (0.4-1 m) in height and are densely branched [38,168]. Plants on upland sites tend to be smaller than plants on bottomlands and other moist sites [176]. The lanceolate leaves are thin and narrow, ranging from 0.8 to 3.5 inches (2-9 cm) in length [57,157,168]. Plants with lobed leaves are probably showing introgression with big sagebrush [12,188]. Vegetative shoots are evergreen and perennial; floral shoots are annual but somewhat persistent after drying [72]. The inflorescence is a panicle of perfect disc flowers [57,75]. The fruit is a small (~ 2.5 1 mm), sticky-walled cypsela bearing a single, tiny seed [38,111,168,171,191]. Harvey [71] reported mean mass, length, and width of plains silver sagebrush seed from Montana at 680 g, 2.1 mm, and 0.90 mm, respectively. The root system consists of a taproot with lateral roots [71]. Plants in Saskatchewan had deep taproots (7-10+ ft. (2-4+ m)) without much lateral branching in upper soil layers [36]. Silver sagebrush is rhizomatous; rhizomes are generally located within a few inches of the soil surface [71]. Rhizome length of plains silver sagebrush in eastern Montana averaged 3.4 feet (1.1 m): 3.3 times more than mean plant height of 13 inches (32 cm). Individual rhizomes bore 1 to 52 sprouts [187].

Plant descriptions provided in this section present information relevant to fire ecology and are not meant as identification keys. Several florae [38,57,75,111,168,191] provide keys for identifying silver sagebrush.

Subspecies: Infrataxa are morphologically distinguished by plant height and relative leaf size, color, and hairiness of herbage [75,191]. Distinctions are not clear-cut, and there is considerable overlap in the subspecies' characteristics [38]. Beetle [12] and Winward [198] provide keys for distinguishing silver sagebrush infrataxa. Gross differences in the subspecies' morphology follow.

Bolander silver sagebrush is a relatively low, thickly branched, round shrub [91]. It is less than 3 feet (0.9 m) in height. Stems are woody at the base and bear relatively small (1- to 2-inch-long (3-5 cm)), narrow leaves. Fruits are less than 1.2 mm in length, and resinous [75]. Bolander silver sagebrush is easily distinguished because of its restricted range [12].

Plains silver sagebrush is the tallest of the subspecies. It has an erect, freely branching growth form, typically reaching 3 to 4 feet (0.9-1 m) in height at maturity but sometimes as much as 4.9 to 6.6 feet (1.5-2 m) [57,90,168]. It has relatively larger, wider leaves (2-4 inches 0.4 inch (4-9 1 cm)) with a denser, whiter covering of tomentum [38,89] compared to mountain silver sagebrush. 

Mountain silver sagebrush is erect and freely branching in form [90]. It grows from 0.3 to 1 foot (0.1-0.3 m) in height. It is differentiated from plains silver sagebrush by its relatively shorter stature, smaller (0.8-5.3 cm long), narrower, more strongly lobed leaves, and less silvery-white (often dark green) herbage [12,85,191].

Species' stand/age class structure: Silver sagebrush communities are usually structurally simple, consisting of silver sagebrush and an understory of graminoids. Silver sagebrush is often the only shrub [65,143,155,203], and forb cover is usually sparse [64,132]. Spacing varies from widely spaced silver sagebrush plants or clustered colonies to a close arrangement of shrubs [65,69,78,132,186]. Silver sagebrush in late succession can form nearly closed cover [69,198]. Hirsch [78] found a range of 674 to 4,519 silver sagebrush per hectare in a plains silver sagebrush-western wheatgrass community in southwestern North Dakota. Another western North Dakota study showed the few forbs in the community were most abundant near plains silver sagebrush, while bunchgrasses were more plentiful in shrub interspaces [73]. Bare ground and biological soil crust cover tend to be low in undisturbed silver sagebrush communities [15,132]. On a mountain silver sagebrush community in eastern Oregon, mean bare ground and litter cover on undisturbed soils were < 3% and 80%, respectively. Maximum bare ground cover on disturbed soils was 17% [132].

Woody sagebrush species are long lived, with some species exceeding 100 years of age [193]. As a cloning species, however, silver sagebrush probably does not fit this general pattern. Maximum life span and typical age class structure of silver sagebrush need further investigation. Wambolt and others [187] found age class structure of plains silver sagebrush in eastern Montana was as follows:

Plant part Mean age (yrs) Standard deviation Number of samples
stems 3.4a,1 2.0 204
taproots 6.9b 3.1 28
parent rhizome2 8.8c 3.7 68
rhizome system3 6.0b 2.5 128
1Significant (p<0.05) differences followed by different letters
2Rhizome originating from a parent (or dead) stump to which sprout was directly connected.
3Rhizomes sections other than in 2 above.

Physiology:
Flooding - Silver sagebrush has superior flooding tolerance compared to other woody sagebrush species [157].

Drought - Drought tolerance of silver sagebrush is uncertain. The species is markedly sensitive to water stress in the seedling stage [186,187], and has been noted as either drought intolerant [112] or tolerant [47,60] at maturity. Within the species, plains, mountain, and Bolander silver sagebrush have been ranked least to most drought tolerant, respectively [163]. Anecdotal evidence from northern Great Plains studies during and after the "dust bowl" era (1937-1943) suggests that silver sagebrush is well adapted to survive severe, prolonged drought. On study sites across the northern Great Plains, adult postdrought survivorship of silver sagebrush was comparable to big sagebrush, and silver sagebrush showed better seedling establishment than big sagebrush after the drought had ended [47,144].

RAUNKIAER [141] LIFE FORM:
Phanerophyte
Geophyte

REGENERATION PROCESSES:
Silver sagebrush reproduces from seed and by sprouting and layering [71,186].

Breeding system: Silver sagebrush's perfect flowers are uniformly fertile [111]. Most breeding is accomplished by outcrossing, although some selfing occurs [112]. DNA studies of plains silver sagebrush showed evidence of interpopulation outcrossing, but differences in genetic diversity among populations were not great. McArthur and others [115] state that large population sizes and wind-effected pollination tend to minimize between-population differences in silver sagebrush.

Pollination: Pollen is mostly spread by wind. Since bees, beetles, and flies regularly visit silver sagebrush flowers, some insect-mediated pollination probably occurs [112,115,182].

Seed production: Sagebrush species tend to have high fruit set, seed set, and germination rates. High seed production and germination rates help offset the genus' generally low rate of seedling establishment (see Seedling establishment) [182]. Although a good fruit and seed producer, silver sagebrush has a lower rate of seed production than other species in its subgenus, with fruit set 18% lower than big sagebrush [71,179]. Silver sagebrush 1st starts producing seed at about 4 years of age [147].

Seed dispersal: At time of dispersal, the involucral bracts spread and release the ripe fruits [168]. The single seed remains contained within the fruit, which falls beneath or slightly downwind of the parent plant. Some long-distance dispersal probably occurs when the sticky fruits adhere to passing animals [71,159]. Wind and water also disperse seed away from the parent plant [12,49,187]. Fruits that fall on crusted snow may be moved long distances by wind. Sagebrush seeds float, and water currents may also carry seeds considerable distances [159].

Seed banking: Studies on potential seed banking of silver sagebrush are sparse, but a few studies suggest that silver sagebrush may build up a short-term seed bank.  Silver sagebrush seed retains its ability to geminate when stored under cool, dry conditions, but looses germinative capacity rapidly when exposed to unfavorable conditions. Eddleman [45] obtained better (73%) and faster germination from stratified, year-old seed than from unstratified 2-month-old seed (26% germination) in the laboratory. However, Romo and Young [148] found plains silver sagebrush seed rapidly lost its ability to germinate when exposed to field environments in Saskatchewan. Spring-sown seed showed significantly better emergence than fall-sown seed, which was more likely to be killed by fungi or drastic fluctuations in temperature. Some seeds that failed to germinate within the 1st year were still viable, and the authors suggest that silver sagebrush may have a small soil seed bank reserve on safe sites. Further studies are needed to determine ability of silver sagebrush to establish from soil-stored seed.

Germination: Silver sagebrush seed is immediately germinable, although best germination occurs when seeds are retained on the parent plant long enough to experience cold temperatures [45,186]. Seed germinates with or without light [184,186] over a wide range of temperatures [186]. Plains silver sagebrush seed collected in Montana, stored at room temperature for 2 years, and sown in the greenhouse showed 51% and 64% germination on filter paper (without and with fungicide treatment, respectively). Optimum germination appears to occur with persistently cool soil temperatures. When sown in the greenhouse, seeds showed 93% germination at 68 oFahrenheit (20 oC) and 97% germination at 59 oFahrenheit (15 oC) [71]. A Saskatchewan study found best germination of plains silver sagebrush occurred with stratification (> 28 days' duration) around 50 oFahrenheit (10 oC). Temperature range for germination was 41-77 oFahrenheit (5-25 oC) [146]. In the Montana study, exposure to relatively high temperatures (86 oFahrenheit (30 oC) for 14 days) prior to planting resulted in loss of seed viability. Germination rates were best for seed planted 2.5 mm below the soil surface. No germination occurred in seed more than 7.5 mm below the soil surface. Seeds sown on the soil surface germinated readily but were highly susceptible to desiccation [71]. 

Seedling establishment/growth: Although silver sagebrush seed production is high, few plants survive the germination and seedling stages [185]. Artificial plains silver sagebrush regeneration, established in southern Montana by seeding-in, showed  mean 1st-year survivorship of 3.4% [71]. Seedlings require open ground that is free from competition [71,147], and they are rare in undisturbed to lightly disturbed, mature stands [71,73]. On undisturbed steppes in the Badlands of North Dakota, plains silver sagebrush comprised 100% of total shrub cover, only 4% of which was seedlings [186]. Good establishment occurs on soils disturbed by animal burrowing or tilling [147]; the incidence of postfire seedling establishment needs further study. Silver sagebrush seedling mortality is greatest in the 1st year, tending to level out after that [71,186]. Investigators in Saskatchewan [83] found that up to a critical mass, seedling establishment and growth rates of plains silver sagebrush seedlings increased with increasing weight of their originating seeds; however, plants from seeds weighting > 0.57 mg showed poor germination, and growth of seedlings from the heaviest seeds was slow. In a Saskatchewan study, Romo and Grilz [147] found that only 5-6% of sown silver sagebrush seed produced seedlings, but 85% of seedlings that survived their 1st year also survived their 2nd. Drought is the primary cause of seedling death. Seedlings require moist soil for establishment and growth [186,187], and pulses of seedling establishment have been noted in years of above-average precipitation [187].

Acclimated seedlings are cold-hardy. Plains silver sagebrush seedlings from Saskatchewan survived exposure to -38 oFahrenheit (-39 oC) temperatures after hardening-off, while nonacclimated seedlings did not survive exposure to 7 oFahrenheit (-14 oC) temperatures. The researchers suggested that seedlings germinating early in spring may show better winter survivorship than later-germinating seedlings, as older seedlings that have gradually acclimated to cooling temperatures have had more time to develop protection from freezing by lignifying cell walls and expanding twig buds [82].

Asexual regeneration: Cloning is silver sagebrush's most common method of reproduction [148,187]. With 3 methods of sprouting and the ability to layer [12,71,72,90,189], silver sagebrush is the strongest sprouter in its subgenus [71,186,200]. It sprouts from the roots [12,14,17,38,71,72,114,126], rhizomes [12,17,114,126,157,171,189], and the root crown [47,152,189] after disturbance damages stem tissue [38,72,126,157] and removes apical dominance. A strong sprouting response from buried organs increases silver sagebrush's ability to survive flood, ice scour, drought, and even severe fire damage [47,186,187]. Enlargement of colonies via sprouting and layering is nearly the sole method of silver sagebrush regeneration in the absence of disturbance; both seedlings and sprouts may occur on disturbed sites. Origin of excavated plains silver sagebrush plants in Montana was approximately 37% from seed (no connecting rhizomes) and 63% from sprouts (plants connected by rhizomes). An excavated colony consisted of 15 individuals, all connected by a rhizome running within 2 inches (5 cm) of the soil surface. Each plant had its own root system. Soil texture may affect ability to clone: the only sites where plants established from seed outnumbered those originating from rhizomes were those with very gravelly or very clayey soils [71]. Similarly, another Montana study of plains silver sagebrush regeneration found plants started from seed outnumbered cloned plants only on sites with clay pans underlaid with gravel. Typical rates of establishment were 1/3rd from seed and 2/3rds from cloning [187]. 

Harvey [71] found layering was an important means of regeneration for silver sagebrush in eroded gullies. Stems that had been buried in silt by periodic floods were producing adventitious roots and sprouting; rhizomes were also producing sprouts.

Sprouting from the root crown is less frequently reported in the literature than sprouting from roots or rhizomes, but root crown sprouting may be an important adaptive response to drought and browsing. Approximately 3/5ths of the plains silver sagebrush plants on an eastern Montana site gradually died back to the root crown during the severe drought of the early1930s, sprouting back from the root crown in the spring of 1935. The remaining 2/5ths did not recover [47]. In a study of the response of native shrubs to pruning (a surrogate for browsing), silver sagebrush responded to annual pruning to a height of 0.5 foot (0.2 m ) by "vigorous" sprouting from the root crown. The sprouts grew quickly: after 3 years of treatment, there was no difference in mean height between pruned and unpruned silver sagebrush plants (3 feet (0.9 m)), although crown area was significantly less in pruned vs. unpruned plants (03.2 vs. 42.4 ft2 (2.81 vs. 3.91 m2), respectively; p = 0.05). [152].

Hybrids: Silver sagebrush hybrids may inherit ability to sprout and layer [71,115]. For example, a natural mountain silver sagebrush mountain big sagebrush hybrid from Gallatin County, Montana, produced root sprouts in the field and layered when moved to the greenhouse [71].

SITE CHARACTERISTICS:
Silver sagebrush requires more moisture than most sagebrush species, growing in areas that receive > 10 inches (250 mm) mean annual precipitation and have a water table within 3 feet (1 m) of the soil surface [157,176]. For example, a Bolander silver sagebrush/Sandberg bluegrass community in Plumas County, California, occurred on streamside soils with a mean July moisture content of 36% and a mean water table depth of 47.6 inches (121 cm) [173]. Because it requires moist soils, silver sagebrush typically grows on the edges of streambanks and drainages and on floodplains, bottomlands, and moist meadows [57,157,190,190].

Soils: Silver sagebrush occupies moister, colder soils than any other woody sagebrush species in North America [193]. Soil drainage is often slow. Silver sagebrush communities are also common on transitional wet-to-dryland sites [63,109] where soils dry by late summer. A mountain silver sagebrush/tufted hairgrass community in Yellowstone National Park was described as "the driest wetland community type" within the Park [24]. Phosphorus, potassium, nitrogen, organic matter, and cation exchange capacity are often lower in silver sagebrush soils than in soils of surrounding communities. Silver sagebrush cannot tolerate strongly saline or calcareous soils [73,157,185]. Soil pH ranges from slightly acidic to strongly alkaline (6.0-8.5) [24,63,66,176]. Parent materials of soils supporting silver sagebrush include sandstones, shales, and granites [130]; soil textures include clay, silt, loam, sand, and gravel [71,85,187]. Silver sagebrush in Wyoming occurred on shallow to deep soil profiles, being most common (9/14 sites) in deep soils (depths of > 3 feet (1 m)) [176]. Best growth occurs on well-drained, coarse-textured, alluvial soils that are moist in the upper 6 inches (20 cm) of the soil profile [63,185].

Elevation: Silver sagebrush's overall elevational range is from 5,000 to nearly 11,000 feet (2,000-3,000 m) elevation [17,157,198].

Subspecies:
Bolander silver sagebrush is most common on pluvial lakebeds, internally drained basins with alkaline soils, and in snow catchments with granitic soils [71,91,113,163,203]. It also grows on meadows, streambanks, and moist, gravelly soils [17,75,98]. It is the only sagebrush in North America that can tolerate temporary inundation [182]. Bolander silver sagebrush occurs from 4,400 to 11,000 feet (1,300-3,400 m) elevation [17,75,98]. In the Ochoco Mountains, it occurs on the margins of mid-elevation meadows (4,400-5,600 ft. (1,300-1,700 m)) and on lower-elevation, inactive floodplains [98]. It occurs from 5,000 to 9,600 feet (1,500-2,900 m) elevation in Nevada [93].

Plains silver sagebrush occurs on low hillsides, riparian zones, and in valleys. Ranging from 4,000 to 7,000 feet (1,000-2,000 m) elevation, it is most common below 5,000 feet (1,550 m) on moist, sandy soil bordering prairie creeks, ephemeral water courses, and the bottoms of eroded hills [14,71,90,117,168]. Topography is usually flat to gently sloping [127]. Scattered plants may occur on upland sites [14].

Mountain silver sagebrush is most common on mountain meadows, stream terraces, basins, and areas of heavy winter snowpack throughout its distribution. It is occasionally found on moist upland slopes [12,85,113,191].  In the Southwest it is most common in the mountains but also occurs on plains and in valleys [111]. Mountain silver sagebrush is common on limestone-derived soils [163]. It occurs from 6,000 to 10,000 feet (2,100-3,050 m) elevation across its range [17,71,191]. Ranges for mountain silver sagebrush by state are:

State Elevation
AZ, NM 5,000-9,000 feet (1,500-2,700 m) [111]
ID 6,000-8,000 feet (1,800-2,400 m) [155]
MT 1,860-5,994 feet (567-1,827 m) [65,66]
NV 5,000-9,600 feet (1,500-2,900 m) [93]
WY 1,000-10,000 feet (1,800-3,000 m)  [20,85,130]

SUCCESSIONAL STATUS:
Silver sagebrush occurs in early to late stages of succession [12,94,131,148]. Reed [143] reported mountain silver sagebrush invading abandoned hayfields in the Jackson Hole Wildlife Park of Wyoming. These silver sagebrush steppes were sometimes stable, and sometimes succeeded to quaking aspen. Silver sagebrush prefers open sites, and is intolerant of all but light shade [189]. It grows on newly exposed soils and can survive fire, browsing, and short periods of flooding disturbance [63,99]. Researchers in Montana have documented a riparian sere on the Yellowstone River that begins on bare, disturbed sandbars, progresses through development of plains cottonwood/sandbar willow (Salix exigua) gallery to a Wood's rose-western snowberry (Rosa woodsii-Symphoricarpos occidentalis) shrubland stage as trees age and die out, and climaxes in a plains silver sagebrush-western wheatgrass-prairie sandreed steppe [18,19,67]. Hansen and others [67] suggest this riverside plains silver sagebrush steppe may be a zootic disclimax induced by cattle overbrowsing the riparian zone. Nelson [127] noted a different successional trajectory in the Badlands of North Dakota. There was no evidence of shrubland and climax steppe vegetation phases after tree death; instead, plains silver sagebrush invaded open plains cottonwood (Populus deltoides ssp. monilifera) galleries, sometimes dominating the understory.

Silver sagebrush may invade overgrazed mountain meadows [142]. Silver sagebrush steppes of Utah occur on mountain rangelands disturbed by heavy grazing; the steppes are usually codominated by herbaceous invaders. In the absence of disturbance, the silver sagebrush steppes progress to stable, late-successional seres codominated by bunchgrasses [131].

Jakubos and Romme [88] noted invasion of Rocky Mountain lodgepole pine (Pinus contorta var. latifolia) into big sagebrush-silver sagebrush/sedge meadows in Yellowstone National Park. The authors attributed the tree invasion to a trend towards warmer climate since the late 1880s.

SEASONAL DEVELOPMENT:
Although classified as evergreen, silver sagebrush sheds many of its leaves in winter [75], retaining a few until spring regrowth [168]. Leaves may drop early in response to drought [47]. Seedlings emerge and new growth of older plants starts from March to June, depending upon latitude and elevation [12,71,82,147]. In southern Montana, established plants initiate growth in late March, with seedlings emerging from mid-April to mid-June [71]. Flowering occurs in July and August in California and Oregon [72,75], from July through September in New Mexico [111], and in August and September in the Great Basin, northern Great Plains, and the interior Pacific Northwest [38,45,57,93,119,126]. The tomentum tends to thin around flowering time [38]. Seed dispersal is prolonged from early fall through the following spring [148]. Fruit and seed disperse as a unit, with most seed shattering from September through November [12,45,57,119,148,186]. Some fruits are retained on the parent plant until winter or spring [148].

Phenology of plains silver sagebrush on 6 sites across southern Montana was as follows [71]:

Event Timing
dormancy ends early to late March
apical bud enlargement late March  to mid-April
twig elongation mid-April to mid-May
lateral branch bud enlargement mid-May to mid-June
lateral branch elongation late May to mid-Aug.
floral branch elongation mid-June to mid-Aug.
floral bud enlargement July
anther development mid-July to mid-Aug.
flowering mid-Aug. to early Sept.
fruit ripens early Sept. to mid-Oct.
seed dissemination mid-Sept. to mid-Oct.
onset of winter dormancy mid-Oct. to mid-Nov.

FIRE ECOLOGY

SPECIES: Artemisia cana
FIRE ECOLOGY OR ADAPTATIONS:
Fire adaptations: Silver sagebrush has a strong sprouting response after top-kill by fire [25,38,199,200]. Because it possesses several organs capable of regeneration, including roots and rhizomes that are protected by soil, it is not as susceptible to fire mortality as most woody sagebrush species [13,119,186,187].

Although sprouting is silver sagebrush's primary method of postfire regeneration, some seedling establishment can occur if early postfire years are favorably wet during the growing season [187]. Time required for postfire sprouts to begin seed production is undocumented, but given silver sagebrush's rapid rate of sprout growth [152], sprouting stems probably fruit and seed within 2 or 3 postfire years. Seeds produced from postfire sprouts are the most likely sources of seedling establishment, but wind, water, and animal transport of seed onto burns may also contribute to postfire seedling establishment [12,49,71,159,187].

Fuels: Occupying bottomlands and other areas that are wetter than surrounding vegetation, silver sagebrush communities often have heavier litter and vertical fuel loads than surrounding plant communities [132,177,186]. Fires tend to carry into and spread in silver sagebrush unless the community has been overgrazed [17,186]. Heavy litter build-up tends to decrease rhizomatous sprouting in silver sagebrush colonies, while removal of the litter by fire or other means encourages silver sagebrush sprouting and colonial expansion [185,187].

Fire regimes: Silver sagebrush steppes experience stand-replacement fires [14,134,195]. Fire frequencies are uncertain: fire histories for silver sagebrush communities are sparse to altogether lacking. Since plant productivity and community structure vary across the species' wide geographical distribution [65,69,78,132,186], historic fire intervals were probably similarly varied.

Bolander silver sagebrush: As of this writing (2002), there are no published fire history studies on Bolander silver sagebrush communities. Bolander silver sagebrush grows on sites with widely varying characteristics, including fuel loads. Fire was probably rare in Bolander silver sagebrush communities on alkali sinks in the rain shadow of the Sierra Nevada, and frequent on Bolander silver sagebrush meadows of the Ochoco and Blue mountains. Since even plant community composition, structure, and production are poorly described in the literature, a great deal of research is needed to understand fire effects and historic fire regimes for Bolander silver sagebrush communities. Fire and rangeland managers with experience in this type are encouraged to share their findings with Fire Effects Information System staff.

Plains silver sagebrush: Fire history studies of plains silver sagebrush-grama-needlegrass steppes of the northern Great Plains are based on historic records and tree fire scars obtained from adjacent, coniferous communities and a few lone, on-site conifers. These studies suggest that stand-replacement fires occurred frequently in plains silver sagebrush steppes. Wright and Bailey [199] estimated fire frequencies of 5 to 10 years on rolling-to-level topography, where plains silver sagebrush mostly occurs as widely scattered plants in a landscape dominated by mixed grasses [71]. On topography dissected with breaks and rivers, where silver sagebrush is most common, they estimated fire return intervals of 20 to 30 years. Other researchers report similar fire return intervals for plains grassland steppes [27,52,192]. For example, fire return intervals range from 16 to 47+ years in relict western wheatgrass-green needlegrass-plains silver sagebrush communities of western North Dakota [137].

Most plains grassland fires historically occurred in summer or fall, with lightning and human ignition sources. Lightning-caused ignitions were most common in July and August. Sieg [165] suggests that for the northern Great Plains, the natural fire regime of low draws that were vegetated with sprouting species such as silver sagebrush was mostly late-growing-season fire, ignited after fine fuels had cured and dried. Late-season fires were probably "intense," but because natural fires also occurred in other seasons, the overall fire regime was highly variable.  Native Americans traditionally set fires throughout the year, although April, September, and October were their peak ignition times [76].

Mountain silver sagebrush: The cool, moist montane sites supporting mountain silver sagebrush can be highly productive, with graminoids providing ample flashy fuels to carry fire [197]. Fire ecologists estimate frequent stand-replacement fires in this type, with mean fire return intervals ranging from 3 to 45+ years [5,6,74]. Houston [84] estimated on Snake River Plains of Idaho, fires probably cycled about every 25 years in the wetter areas favored by mountain silver sagebrush.

The following table provides some fire regime intervals for plant communities and ecosystems where silver sagebrush is a dominant or important component of the vegetation. For further information on these fire regimes, see the FEIS species summaries on the community and ecosystem dominants listed below.

Community or Ecosystem Dominant Species Fire Return Interval Range (years)
silver sagebrush Artemisia cana 5-45 [74,137,199]
sagebrush steppe A. tridentata/Pseudoroegneria spicata 20-70 [133]
basin big sagebrush A. t. var. tridentata 12-43 [154]
mountain big sagebrush A. t. var. vaseyana 15-40 [7,29,122]
Wyoming big sagebrush A. t. var. wyomingensis 10-70 (40**) [183,201]
saltbush-greasewood Atriplex confertifolia-Sarcobatus vermiculatus < 35 to < 100
plains grasslands Bouteloua spp. < 35
blue grama-needle-and-thread grass-western wheatgrass B. gracilis-Hesperostipa comata-Pascopyrum smithii < 35
blue grama-buffalo grass B. g.-Buchloe dactyloides < 35
cheatgrass Bromus tectorum < 10
western juniper Juniperus occidentalis 20-70
Rocky Mountain juniper J. scopulorum < 35
wheatgrass plains grasslands P. smithii < 35 [133]
interior ponderosa pine* Pinus ponderosa var. scopulorum 2-30 [6,10,104]
eastern cottonwood Populus deltoides < 35 to 200 [133]
quaking aspen (west of the Great Plains) P. tremuloides 7-120 [6,58,121]
mountain grasslands Pseudoroegneria spicata 3-40 (10**) [5,6]
Rocky Mountain Douglas-fir* Pseudotsuga menziesii var. glauca 25-100 [6,7,8]
*fire return interval varies widely; trends in variation are noted in the species summary
**mean

POSTFIRE REGENERATION STRATEGY [169]:
Small shrub, adventitious bud/root crown
Rhizomatous shrub, rhizome in soil
Geophyte, growing points deep in soil
Initial off-site colonizer (off-site, initial community)
Secondary colonizer (on-site or off-site seed sources)

FIRE EFFECTS

SPECIES: Artemisia cana
IMMEDIATE FIRE EFFECT ON PLANT:
Surface fire top-kills silver sagebrush [14,134,195]. Because perennating buds on rhizomes and roots are protected by soil, silver sagebrush ordinarily survives even severe surface fire [98]. Fuel loads in most silver sagebrush communities are usually sufficient to ensure a high rate of top-kill. The foliage is moderately flammable, and silver sagebrush litter is highly flammable when dry [124]. Fires usually spread in silver sagebrush communities during dry seasons [25,26,124,200].

DISCUSSION AND QUALIFICATION OF FIRE EFFECT:
Fall burning under dry conditions may result in greater mortality compared to spring burning due to internal water stress of silver sagebrush plants [119,195]. Fire Case Studies summarizes a study on seasonal differences in fire effects on silver sagebrush.

Ground fires are rarely reported in silver sagebrush steppes; however, in Saskatchewan, Rowe [151] noted a September, lightning-ignited ground fire in a silver sagebrush/green needlegrass-blue grama-western wheatgrass association. In some spots, the fire burned into the taproots of silver sagebrush plants and killed them.

PLANT RESPONSE TO FIRE:
Silver sagebrush has the strongest sprouting response of any woody sagebrush species in North America [25,199]. It sprouts from the root crown [134], rhizomes [12,14,17,114,187], and roots [12,14,17,114] after top-kill by fire [25,198,200]. Because of its strong postfire sprouting response, fire on a 5 to 20-year rotation promotes silver sagebrush [17,187]. It can return to or exceed canopy coverage in 4 to 6 postfire years [198]. For example, fall prescribed burning of a Bolander silver sagebrush/Baltic rush wet meadow in Inyo National Forest, California, produced no significant change in postfire silver sagebrush cover (measured for 4 postfire years) compared to prefire cover [70]. Dry weather and fuel conditions before fire can reduce the number of sprouts silver sagebrush produces after fire [195]. Very frequent fire tends to favor associated bunchgrasses over silver sagebrush [98].

Silver sagebrush requires an open, disturbed seedbed [71,147], and fire may prepare a favorable site for seedling establishment. In eastern Montana, plains silver sagebrush seedling establishment has been documented on sites disturbed by fire or ice scouring. Rhizome sprouting was more common, however (approximately 1 seedling per 3 sprouts) [187]. In a rehabilitation study using artificial regeneration, mixed sagebrush steppe in Wyoming was treated to control invading Utah juniper (Juniperus osteosperma). Control methods included herbicide (picloram) application, bulldozing, tree harvest, and a fortuitous wildfire. After the treatments and the wildfire, native species including silver, big, and black (Artemisia nova) sagebrush were seeded in, with time of seeding (spring vs. fall) and seeding method (furrow vs. broadcast seeding) evaluated. All sagebrush species established best on burned plots compared to unburned plots; fall planting and broadcasting also favored the sagebrushes compared to spring seeding and furrowing [53].

Hybrids: Silver sagebrush hybrids may inherit the ability to sprout after fire and other top-killing events. Silver sagebrush mountain big sagebrush and silver sagebrush threetip sagebrush hybrids both sprout after fire [97,112]. Inadequately studied, postfire sprouting response of silver threetip sagebrush hybrids is probably strong because each parent has genetic ability to sprout. Silver and threetip sagebrushes were once assumed to be closely related due to common sprouting ability [12,164]; however, DNA-based phylogenetic studies show a close genetic relationship among black sagebrush, low sagebrush, and silver sagebrush infrataxa (the silver sagebrush lineage)  [96,97,120], and a more distant genetic relationship between the silver sagebrush lineage and the big sagebrush lineage (which includes threetip sagebrush and big sagebrush infrataxa) [96,97]. Studies on the sprouting ability of silver, black, and low sagebrush combinations are lacking, but such studies might provide insight into the genetic basis for sprouting in woody sagebrushes and ultimately, produce woody sagebrush plants that can survive on sites where exotic annual grasses have altered fire regimes. Sprouting, drought-tolerant sagebrush hybrids may be useful in rehabilitating sagebrush rangelands that have converted to annual grasslands dominated by cheatgrass or medusahead (Taeniatherum caput-medusae) after repeated, short-interval fires. Big and threetip sagebrush seedlings are more drought tolerant than silver sagebrush seedlings. Crosses among silver, big, and threetip sagebrush may produce sprouting plants that can survive on dry sites prone to reburns [112]. McArthur and others [115] have produced a plains silver sagebrush Wyoming big sagebrush hybrid with apparently fertile seed.

DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:
A late summer wildfire promoted plains silver sagebrush on an Alberta western wheatgrass-blue grama prairie. The grassland, used as a cattle range, was in excellent condition prior to the wildfire. Three successive years of drought followed the wildfire, and cattle were allowed free access to burned and unburned portions of the range. Stocking rate was 0.64 AUM/ha. Plains silver sagebrush was the most common shrub in the postfire plant community, and its biomass was greater on burned than on unburned plots on both upland and bottomland sites [48].

Fire also favored silver sagebrush on the Grand Mesa National Forest, Colorado. A Gambel oak/common snowberry-Saskatoon serviceberry community (Quercus gambelii/Symphoricarpos albus-Amelanchier alnifolia) was prescribed burned, sprayed, and/or chained to reduce density and height of big game and cattle browse. Production of silver sagebrush (kg/ha) before and after treatments is given below [102].

1 yr before 2 yrs after 5 yrs after 10 yrs after
burning 10.67 11.80 20.27 17.33
spraying data not given ---- ---- ----
chaining 8.40 2.93 4.40 2.20
burning, spraying, and chaining 1.20 0.53 0.60 2.00


FIRE MANAGEMENT CONSIDERATIONS:
Beetle and Johnson [14] recommend prescribed fire to increase silver sagebrush density on rangelands. Blaisdell and others [17] provide general guidelines for burning in sagebrush-grasslands.

A variety of fire treatments may be needed to mimic historic fire regimes. Because the fire regime of the northern Great Plains was highly variable, Sieg [165] recommends varying fire intensities and seasons, including use of mid-summer fire, to restore natural disturbance patterns. Howe [87] stated "the strategy should avoid a uniformity of timing of burns or in intervals between burns that artificially simplifies what was probably a more complex system."


FIRE CASE STUDIES

SPECIES: Artemisia cana
FIRE CASE STUDY CITATION:
Howard, Janet L. 2002. Seasonal effects of prescribed burning in eastern Montana on silver sagebrush. In: Artemisia cana. 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/ [].

REFERENCE:
White, Richard S.; Currie, Pat O. 1983. The effects of prescribed burning on silver sagebrush. Journal of Range Management. 36(5): 611-613. [195].

SEASON/SEVERITY CLASSIFICATION:
Spring (mid-April)/Low severity
Fall (late Oct.)/Moderate severity

STUDY LOCATION:
The study sites were on the Livestock and Range Research Station near Miles City, Montana.

PREFIRE VEGETATIVE COMMUNITY:
The community is a plains silver sagebrush/western wheatgrass-blue grama (Artemisia cana spp. cana/Pascopyrum smithii-Bouteloua gracilis) steppe. Threadleaf sedge (Carex filifolia), needle-and-thread grass (Hesperostipa comata), green needlegrass (Nassella viridula), and Japanese brome (Bromus japonicus) are common associates.

TARGET SPECIES PHENOLOGICAL STATE:
Silver sagebrush was breaking dormancy mid-April. In mid-October it was dispersing fruits and seed.

SITE DESCRIPTION:
Not provided.

FIRE DESCRIPTION:
Spring burning was conducted in mid-April of 1977, shortly after spring run-off. Soil moisture was near field capacity. A headfire was used.

Fall burning was conducted in early October of 1979, after summer drought "substantially reduced soil moisture throughout the soil profile." A backfire was used.

Fuel loadings and weather conditions were similar on the spring- and fall-burned sites. Fuel loads ranged from 300 kg/ha pm plains silver sagebrush/blue grama sites to more than 1,000 kg/ha on plains silver sagebrush/western wheatgrass sites. Temperature averaged 21 oC and wind speed was less than 8 km/hr.

FIRE EFFECTS ON TARGET SPECIES:
Plant measurements were taken the July after burning (n=210 plants). Fire effects were classified in 3 progressively severe groups: foliage consumed; foliage and twigs consumed with main stems remaining; and entire plant burned to the root crown (top-kill). Silver sagebrush survivorship, number of sprouts, height of sprouts at plant center, and an index of canopy cover* were measured. Mean annual precipitation was below normal throughout the study period.
*Model based upon linear intercept of foliage as measured on major and minor plant axes of the plant's elliptical area

Immediate fire effect: The fires accomplished partial to complete top-kill as described in the severity groups above. Silver sagebrush mortality was dependent upon fire severity and season. The spring fire killed approximately 1/3rd of the plants that were completely burned. Plants that were not completely top-killed sustained mortalities of less than 10%. The fall fire killed about 3/4ths of the completely burned plants, with 40% mortality of incompletely consumed plants. Mortality differences were highly significant (p<0.01) between seasons.

Plant response to fire: Prefire plant size had no significant effect (p < 0.05) on either fire survival or postfire growth. Severe fire reduced height growth of postfire sprouts on top-killed plants compared to fires that only partially consumed plants. Plants that had only foliage removed tended to sprout more quickly than those also losing branches. Spring burning resulted in plants with relatively fewer sprouts, superior height growth, and greater canopy indices  compared to fall burning. For spring-burned silver sagebrush, height growth by the next July was > 20 cm for top-killed plants, and > 30 cm for partially consumed plants. Canopy indices* (see above) were > 350 mm. For the fall burns, incompletely burned plants usually had > 10 sprouts per plant, and the sprouts were < 20 cm in height by the next July. Top-killed plants averaged 4 sprouts per plant, with sprouts < 4 cm in height by the next July. Mortality and postfire growth data are given below.

Mean growth in relation to fire severity. Measurements were made in July 1977 or 1980, after plants had passed through 1 spring growing season.

Fire severity

Growth characteristics

  Number of sprouts Height (mm) Canopy index (mm)
Spring fire
   foliage consumed 4.5a1 322a 394a
   twigs & foliage consumed 3.8a 305a 424a
   top-killed 3.3b 216b 358a
Fall fire
   foliage consumed 10.3a 188a 341a
   twigs & foliage consumed 13.3a 188a 279a
   top-killed   4.2b   56b   96b
1Differences between values are significant (p < 0.05) when followed by a different letter.

FIRE MANAGEMENT IMPLICATIONS:
Prescribed burning can be used to alter silver sagebrush communities. Burning under favorable spring moisture conditions, before silver sagebrush stem elongation occurs, results in relatively low silver sagebrush mortality and vigorous postfire sprouting. In this study, silver sagebrush cover was nearly back to prefire levels the 1st growing season after the spring fire. Burning in fall, just before onset of dormancy, results in relatively high mortality, less vigorous sprouting, and reduced canopy cover.

MANAGEMENT CONSIDERATIONS

SPECIES: Artemisia cana
IMPORTANCE TO LIVESTOCK AND WILDLIFE:
Silver sagebrush provides valuable habitat and forage for wildlife. Deer, elk, pronghorn, bighorn sheep, and sage-grouse browse the foliage [51,198]. Voles (Microtus spp.) feed on silver sagebrush roots and rhizomes [186]. On the Great Plains, browsing ungulates generally make little use of silver sagebrush foliage in summer [14], but use plains silver sagebrush habitats all year due to their relatively mesic conditions. Bison and mule deer, for example, make heavy year-round use of use of plains silver sagebrush/western wheatgrass steppes of Theodore Roosevelt National Park, North Dakota [65]. Mule deer may browse silver sagebrush heavily when other forage is dormant [103,108]. Silver sagebrush is also important on fall and winter ranges [89,171,186]. Bolander silver sagebrush/bottlebrush squirreltail (Elymus elymoides) communities were rated as highest-use areas for mule deer on the Silver Lake-Fort Rock winter range of Lake County, Oregon, from November through May [106]. On the Missouri River Breaks of Montana, Rocky Mountain mule deer made greatest use of silver sagebrush in winter; it was a minor component of their spring and fall diets. Silver sagebrush on the Missouri River Breaks composed 33% total volume of  elk rumens (n=12) from fall samples; however, it was only present in trace amounts in samples from a preceding year. It was not found in rumen samples from cattle [108].

In regions with dry summers, livestock and wildlife may browse silver sagebrush when upland vegetation has already dried. In Modoc County of northeastern California, Bolander silver sagebrush communities were preferred summer-use areas for pronghorn, feral horses, and cattle [153]. Although a common staple for wildlife, livestock use of silver sagebrush is variable depending upon availability of palatable herbs [186]. Domestic sheep generally browse silver sagebrush more heavily than cattle [14]. Cattle may prefer grazing in silver sagebrush steppes, however, because their relatively mesic soils promotes heavier grass cover than in adjacent communities, especially in summer [95]. Livestock may actually make greater use of silver sagebrush when there is ample grass to go with it [198].

Silver sagebrush steppes are important habitat for wildlife. For example, they are the 2nd-most used habitat for sharp-tailed grouse in eastern Montana; the birds used only upland western wheatgrass-blue grama grassland habitats more than plains silver sagebrush steppes [174]. Silver sagebrush communities also provide important corridors for wildlife traveling from riparian zones to forested or grassland communities [129,132]. Three wildlife species, pronghorn, sage-grouse, and a native grasshopper, sometimes rely on silver sagebrush steppes as their primary habitat, and on silver sagebrush browse as their primary forage.

Silver sagebrush is used as an indicator species in pronghorn habitat suitability index models [3]. It provides winter maintenance and emergency forage [59,172]. Pronghorn also use silver sagebrush in the fall, particularly when vegetation on less mesic sites has dried [4,11,43]. For example, stomach samples of pronghorn taken during hunting season (Sept.-Oct.) in southeastern Montana were primarily sagebrush (plains silver and big sagebrush); several samples contained no other forage [34]. Pronghorn in Saskatchewan were concentrated on fall rangelands where plains silver sagebrush and creeping juniper (Juniperus horizontalis) were available [43]. Heavy livestock grazing can degrade silver sagebrush habitat conditions for pronghorn [66]. Fires that top-kill large patches of silver sagebrush can adversely affect pronghorn habitat [9] until sprouts have attained a few years' growth.

When plentiful, silver sagebrush is important in the diet of sage-grouse [105,140]. Where it is the primary sagebrush cover, silver sagebrush communities are critical sage-grouse habitat [1]. Crop analysis of adult sage-grouse in White Pine County, Nevada, showed the birds were consuming almost nothing but mountain silver sagebrush [59]. Sage-grouse populations at the northern and eastern boundaries of the bird's distribution are most likely to be ecologically dependant on silver sagebrush [23]. Sage-grouse in southeastern Alberta -- where other woody sagebrush taxa are uncommon -- nest under plains silver sagebrush almost exclusively, and silver sagebrush is their primary diet item [1].

Silver or sand sagebrush are the primary diet items of the sagebrush grasshopper (Melanoplus bowditchi). A North Dakota study showed crop contents of sagebrush grasshoppers were 97% silver sagebrush. High densities of sagebrush grasshoppers have never been recorded, suggesting few economic impacts and little damage to silver sagebrush populations are caused by the insect. However, little is known of the ecology and life history of sagebrush grasshopper, and further studies are needed to assess the ecological and economic impacts of the grasshopper [135].

Palatability: Silver sagebrush is 1 of the more palatable and nutritious sagebrush species [181,198]. With a mean of 11%, winter (Jan.-March) crude protein content of Bolander silver sagebrush on the Fort Rock-Silver Lake winter mule deer range in central Oregon exceeded that of either low or big sagebrush [181]. Mule deer, fed 7 sagebrush taxa collected from the Silver Lake Range in fall and winter, showed highest relative preference* for Bolander silver, low, and mountain big sagebrush in both fall and winter feeding trials. Domestic sheep showed moderate preference (browsed but did not prefer) for Bolander silver and mountain big sagebrush in fall, and showed highest preference for Bolander silver, low, and mountain big sagebrush in winter. Area of collection did not affect the animals' choices, while taxa consistently did [161].
*relative preference index=% of diet/% composition available

Walton [186] has ranked Bolander, mountain, and plains silver sagebrush as most to least palatable, respectively.

Nutritional value: Nutritional content of silver sagebrush is greatest in the spring and declines slowly over winter. Silver sagebrush is rated "fair" in energy value and "fair to good" in protein value [44]. Mean chemical composition of plains silver sagebrush on the Little Missouri National Grasslands, North Dakota, was as follows [51].

  Ca* Mg* K* Cu* Fe (ppm) Mn (ppm) Zn (ppm)
Jan. 0.36 0.20 0.82 7.7 210 36 13
Feb. 0.46 0.61 1.87 4.7 290 33 12
March 0.46 0.17 0.81 4.7 223 39 14
April 0.56 0.29 1.78 5.9 233 72 21
May 0.84 0.57 2.90 6.3 437 119 18
June 0.79 0.47 1.33 5.4 212 46 19
July 1.02 0.45 1.25 0.8 193 39 13
Aug. 0.49 0.33 1.27 4.6 227 35 16
Sept. 0.65 0.30 0.97 3.9 207 43 12
Oct. 0.67 0.33 0.88 3.2 293 48 12
Nov. 0.58 0.31 0.86 3.7 250 47 11
Dec. 0.64 0.30 0.81 3.7 257 32 14
*Dry-weight pecentages.

Based upon nutritional content (protein, carbohydrate, fat, vitamin, and mineral ), palatability, and dependability of supply on rangelands, silver sagebrush is rated low to fair forage for cattle, fair to good for domestic sheep, and good for pronghorn, elk, and deer [63,145].

Cover value: Silver sagebrush provides thermal and hiding cover for small mammals, small nongame birds, and game birds [44,66]. At the lower end of its elevational range, it provides important sage-grouse nesting cover [198].

VALUE FOR REHABILITATION OF DISTURBED SITES:
Silver sagebrush has potential as a soil stabilizer and for use in rangeland, wildlife, and riparian restoration projects [63,112,136,148,159,198]. It is established from seed, containerized stock, and nursery-grown cuttings [136,147]. Due to the species' strong tendency to layer, stem cuttings root easily [71]. Transplants have shown good growth on favorably moist sites. Plains silver sagebrush cuttings, used as part of an upland game bird habitat improvement project on the Snake River Plain of southern Idaho, showed 95% survival and good growth 5 years after transplanting [160]. Silver sagebrush also regenerates when seeded on moist sites, and is recommended for inclusion in range seeding mixtures. Seeding-in is not recommended on dry sites [41,136,198]; however, silver sagebrush seed can germinate on soils with low water potentials due to dissolved salts [45]. Spring sowing is recommended [147,148]. The seed is commercially available [138].

Rehabilitation projects show variability in silver sagebrush's ability to establish on coal mine spoils. In southern Montana, silver sagebrush sown on coal mine spoils showed poor survivorship and seedling growth compared to germinants on unmined plots [71]. However, silver sagebrush cuttings transplanted on coal mine sites in Wyoming and Colorado showed good to excellent survivorship in their 1st year. Transplants on 2 Wyoming sites treated with topsoil showed 100% survivorship. In Colorado, transplants showed 50% survivorship on sites treated with topsoil and 100% survivorship on untreated coal mine spoils. Neither the Wyoming nor the Colorado coal mine sites were irrigated after transplanting [86]. Poor results on some sites may be due to toxic soil contaminants and/or dry soils. Further information is needed on ability of silver sagebrush to establish on contaminated soils.

Information on seed collection, processing, storage, and planting is available for silver sagebrush [45,147,159,189].

OTHER USES:
Native Americans of the northern Rocky Mountains used a decoction of silver sagebrush leaves as a general tonic, and chewed the leaves to quench thirst. Blackfoot and Lakota used silver sagebrush as winter horse browse, and considered it the best sagebrush browse available for big game species. Tribes of the Great Basin used silver sagebrush branches as a fuelbed for roasting pinyon (Cembra) pinecones [123]. Many tribes use the branches in ceremonial rites [56].

Silver sagebrush is listed as a heat and drought tolerant plant suitable for xeriscaping [60]. Since young plants are not drought tolerant [186,187], generous dry-season irrigation is probably warranted for their 1st few years until plants establish a deep root system.

OTHER MANAGEMENT CONSIDERATIONS:
Rangelands: Occupying mesic to moist sites with good graminoid coverage, silver sagebrush communities are often highly productive. A plains silver sagebrush/western wheatgrass community in Theodore Roosevelt National Park, North Dakota, averaged 85% grass and 9% forb coverage [65]. Dry mass production of herbs may reach 2,000 lbs/acre [77,197]. Silver sagebrush tends to increase at the expense of grasses on overgrazed rangelands [89,98,107,142,144,171]. Due to its palatability, however, it too can be overused [89,127,171]. In eastern Montana, a plains silver sagebrush riparian community with a long-term history of year-round cattle grazing was changed to a rest-rotation grazing system. Seventeen years later, silver sagebrush cover had significantly increased (p>0.01) from 10 to 85%. The authors attributed the increase to a combination of increased precipitation and decreased browsing pressure [22]. Smoliak and others [167] recommend that no more than 40% of  silver sagebrush's annual production be browsed in order to maintain silver sagebrush health and productivity. Winward [198] gives the following recommendations for managing silver and other sagebrush rangelands:

Cunningham [39] found that productivity of silver sagebrush in northwestern Colorado was negatively correlated (r=-0.52, p<0.01) with productivity of associated graminoids; forbs were not correlated with silver sagebrush growth. Presence of extractable magnesium in the 12- to 24-inch (30-60 cm) soil layer increased silver sagebrush productivity (r=0.47, p<0.05), as did rock and gravel in the soil profile (r=0.25, p<0.05). A high percentage of sand in the 12- to 24-inch soil layer negatively affected growth (r=-0.15, p<0.01). Harvey [71] provides a model for predicting silver sagebrush productivity based upon leaf and woody tissue measurements.

Silver sagebrush may allelopathically inhibit germination of some herbaceous species, as well as its own seed. Aqueous extracts of silver sagebrush leaves reduced germination of 6 mixed-grass species, including silver sagebrush. Laboratory results using extracts do not necessarily indicate allelopathy under field conditions; however, a field study of plant species distributional patterns in western North Dakota suggests that some forb species are more likely to associate beneath or nearby silver sagebrush canopies. Other species, especially bunchgrasses, tend to segregate away from silver sagebrush [73]. Possible allelopathy of silver sagebrush, and its effects on plant community composition and structure, need further investigation.

Control: Since silver sagebrush provides important wildlife forage and cover, controlling it is not generally recommended [134,198]. Silver sagebrush can obtain nearly closed cover on some sites [69], however, and control may be desired if silver sagebrush is so dense that it is excluding perennial grasses.

Chemical: Silver sagebrush can be controlled with 2,4-D or picloram [17,81,134,194]; 2-,4-D is most effective [17,196]. Follow-up spraying may be necessary to control sprouts [33]. Mechanical controls such as plowing, disking, and chaining are not recommended for mature plants because they are likely to promote sprouting [134]. They may be effective for young plants. White and Currie [194] obtained up to 67% control by plowing silver sagebrush less than 1 m3 in size.

Fire: A single application of prescribed fire is not recommended for control, as it is likely to promote sprouting [134]. Used in sequence, prescribed fire and herbicides may reduce silver sagebrush to desired levels. Using various combinations and rates of herbicides, Whitson and others [196] obtained a range of 8-91% control with fall prescribed fire followed by July application of herbicides to sprouts. Frequent reburning may also reduce silver sagebrush cover, and further studies are needed on the possibility of reduced silver sagebrush using repeated fires. Blaisdell and others [17] provide general guidelines for managing sagebrush-grass rangelands, including assessment of rangeland condition and recommendations for burning. See Fire Management Considerations for further information on managing silver sagebrush with fire.

Artemisia cana: References


1. Aldridge, Cameron L.; Brigham, R. Mark. 2002. Sage-grouse nesting and brood habitat use in southern Canada. Journal of Wildlife Management. 66(2): 433-444. [41664]

2. Allen, Arthur W.; Cook, John G.; Armbruster, Michael J. 1984. Habitat suitability index models: pronghorn. FWS/OBS-82/10.65. Washington, DC: U.S. Department of the Interior, Fish and Wildlife Service. 22 p. [11709]

3. Allen, Eugene O. 1968. Range use, foods, condition, and productivity of white-tailed deer in Montana. Journal of Wildlife Management. 32(1): 130-141. [16331]

4. Amstrup, Steven C. 1978. Activities and habitat use patterns of pronghorns on Montana-Wyoming coal lands. In: Proceedings, 8th biennial pronghorn antelope workshop; 1978 May 2-4; Jasper, AB. Edmonton, AB: Alberta Recreation, Parks, and Wildlife, Fish and Wildlife Division: 270-304. [3300]

5. Arno, Stephen F. 1980. Forest fire history in the Northern Rockies. Journal of Forestry. 78(8): 460-465. [11990]

6. Arno, Stephen F. 2000. Fire in western forest ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 97-120. [36984]

7. Arno, Stephen F.; Gruell, George E. 1983. Fire history at the forest-grassland ecotone in southwestern Montana. Journal of Range Management. 36(3): 332-336. [342]

8. Arno, Stephen F.; Scott, Joe H.; Hartwell, Michael G. 1995. Age-class structure of old growth ponderosa pine/Douglas-fir stands and its relationship to fire history. Res. Pap. INT-RP-481. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 25 p. [25928]

9. Bailey, Arthur W. 1978. Prescribed burning as an important tool for Canadian rangelands. In: McAvoy, S. D. A. M.; Gordon, R. C., co-chairs. Fire and range management: [Seminar proceedings]; [1978, April]; Regina, SK. [Regina, SK: Saskatchewan Department of Agriculture, Lands Branch]: 15-27. On file with: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. [18390]

10. Baisan, Christopher H.; Swetnam, Thomas W. 1990. Fire history on a desert mountain range: Rincon Mountain Wilderness, Arizona, U.S.A. Canadian Journal of Forest Research. 20: 1559-1569. [14986]

11. Bayless, Steve. 1971. Relationships between big game and sagebrush. Paper presented at: Annual meeting of the Northwest Section of the Wildlife Society; 1971 March 25-26; Bozeman, MT. 14 p. On file with: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. [17098]

12. Beetle, A. A. 1960. A study of sagebrush: The section Tridentatae of Artemisia. Bulletin 368. Laramie, WY: University of Wyoming, Agricultural Experiment Station. 83 p. [416]

13. Beetle, Alan A. 1977. Recognition of Artemisia subspecies--a necessity. In: Johnson, Kendall L., ed. Wyoming shrublands: Proceedings, 6th Wyoming shrub ecology workshop; 1977 May 24-25; Buffalo, WY. Laramie, WY: Shrub Ecology Workshop: 35-42. [419]

14. Beetle, Alan A.; Johnson, Kendall L. 1982. Sagebrush in Wyoming. Bull. 779. Laramie, WY: University of Wyoming, Agricultural Experiment Station. 68 p. [421]

15. Belnap, Jayne; Kaltenecker, Julie Hilty; Rosentreter, Roger; [and others]. 2001. Biological soil crusts: ecology and management. Technical Reference 1730-2. Denver, CO: U.S. Department of the Interior, Bureau of Land Management, National Science and Technology Center, Information and Communications Group. 110 p. [40277]

16. 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]

17. Blaisdell, James P.; Murray, Robert B.; McArthur, E. Durant. 1982. Managing Intermountain rangelands--sagebrush-grass ranges. Gen. Tech. Rep. INT-134. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 41 p. [467]

18. Boggs, Keith Webster. 1984. Succession in riparian communities of the lower Yellowstone River, Montana. Bozeman, MT: Montana State University. 107 p. Thesis. [7245]

19. Boggs, Keith; Weaver, T. 1992. Response of riparian shrubs to declining water availability. In: Clary, Warren P.; McArthur, E. Durant; Bedunah, Don; Wambolt, Carl L., compilers. Proceedings--symposium on ecology and management of riparian shrub communities; 1991 May 29-31; Sun Valley, ID. Gen. Tech. Rep. INT-289. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 48-51. [19094]

20. Bramble-Brodahl, Mary K. 1978. Classification of Artemisia vegetation in the Gros Ventre area, Wyoming. Moscow, ID: University of Idaho. 126 p. Thesis. [506]

21. Brand, Michael D. 1980. Secondary succession in the mixed grass prairie of southwestern North Dakota. Fargo, ND: North Dakota State University. 77 p. Dissertation. [14147]

22. Branson, Farrel A.; Miller, Reuben F. 1981. Effects of increased precipitation and grazing management on northeastern Montana rangelands. Journal of Range Management. 34(1): 3-10. [7507]

23. Braun, Clait E. 1998. Sage grouse declines in western North America: what are the problems? In: Proceedings, Western Association of Fish and Wildlife Agencies; 1998 June 26-July 2; Jackson, WY. Cheyenne, WY: Western Association of Fish and Wildlife Agencies: 139-156. [35365]

24. Brichta, Paul Harold. 1986. Environmental relationships among wetland community types of the northern range, Yellowstone National Park. Missoula, MT: University of Montana. 74 p. Thesis. [6727]

25. Britton, Carlton M. 1979. Fire on the range. Western Wildlands. 5(4): 32-33. [514]

26. Britton, Carlton M.; Ralphs, Michael H. 1979. Use of fire as a management tool in sagebrush ecosystems. In: The sagebrush ecosystem: a symposium: Proceedings; 1978 April; Logan, UT. Logan, UT: Utah State University, College of Natural Resources. 101-109. [518]

27. Brown, Peter M.; Sieg, Carolyn Hull. 1996. Fire history in interior ponderosa pine communities of the Black Hills, South Dakota, USA. International Journal of Wildland Fire. 6(3): 97-105. [29220]

28. Bunting, Stephen C.; Kilgore, Bruce M.; Bushey, Charles L. 1987. Guidelines for prescribed burning sagebrush-grass rangelands in the northern Great Basin. Gen. Tech. Rep. INT-231. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 33 p. [5281]

29. Burkhardt, Wayne J.; Tisdale, E. W. 1976. Causes of juniper invasion in southwestern Idaho. Ecology. 57: 472-484. [565]

30. CalFlora. 2000. Information on California plants for education, research and conservation [Online]. Berkeley, CA: The CalFlora Database (Producer). Available: http://www.calflora.org/ [2002, September 19]. [42048]

31. Collins, Ellen I. 1984. Preliminary classification of Wyoming plant communities. Cheyenne, WY: Wyoming Natural Heritage Program/The Nature Conservancy. 42 p. [661]

32. Conroy, Scott D.; Svejcar, Tony J. 1991. Willow planting success as influenced by site factors and cattle grazing in northeastern California. Journal of Range Management. 44(1): 59-63. [14929]

33. Cornelius, Donald R.; Graham, Charles A. 1958. Sagebrush control with 2,4-D. Journal of Range Management. 11: 122-125. [690]

34. Couey, Faye M. 1946. Antelope foods in southeastern Montana. Journal of Wildlife Management. 10: 367. [4236]

35. Coupland, Robert T. 1950. Ecology of mixed prairie in Canada. Ecological Monographs. 20(4): 271-315. [700]

36. Coupland, Robert T.; Johnson, R. E. 1965. Rooting characteristics of native grassland species of Saskatchewan. Journal of Ecology. 53: 475-507. [702]

37. Crawford, Rex C.; Kagan, Jimmy. 2001. Wildlife habitat definitions: No. 16. Shrub-steppe. In: Northwest Habitat Institute, IBIS (Interactive Biodiversity Information System) [Online]. [Adapted from Johnson, David H.; O'Neil, Thomas A., eds. Wildlife habitat relationships in Oregon and Washington. Corvallis, OR: Oregon State University Press]. Available: http://www.nwhi.org/ibis/wildhabs/WHDR_H16.asp [2002, September 19]. [42046]

38. Cronquist, Arthur; Holmgren, Arthur H.; Holmgren, Noel H.; [and others]. 1994. Intermountain flora: Vascular plants of the Intermountain West, U.S.A. Vol. 5. Asterales. New York: The New York Botanical Garden. 496 p. [28653]

39. Cunningham, Hugh. 1971. Soil-vegetation relationships of a bitterbrush-sagebrush association in northwestern Colorado. Fort Collins, CO: Colorado State University. 94 p. Thesis. [724]

40. Dealy, J. Edward. 1971. Habitat characteristics of the Silver Lake mule deer range. Res. Pap. PNW-125. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 99 p. [782]

41. DePuit, Edward J.; Coenenberg, Joe G.; Skilbred, Chester L. 1980. Establishment of diverse native plant communities on coal surface-mined lands in Montana as influenced by seeding method, mixture and rate. Research Report 163. Bozeman, MT: Montana State University, Agricultural Experiment Station. 64 p. [221]

42. Dirschl, Herman J. 1962. Sieve mesh size related to analysis of antelope rumen contents. Journal of Wildlife Management. 26(3): 327-328. [3997]

43. Dirschl, Herman J. 1963. Food habits of the pronghorn in Saskatchewan. Journal of Wildlife Management. 27(1): 81-93. [5939]

44. 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]

45. Eddleman, Lee E. 1977. Indigenous plants of southeastern Montana. I. Viability and suitability for reclamation in the Fort Union Basin. Special Publication 4. Missoula, MT: University of Montana, School of Forestry, Montana Forest and Conservation Experiment Station. 122 p. [42440]

46. Eddleman, Lee E.; Doescher, Paul S. 1978. Selection of native plants for spoils revegetation based on regeneration characteristics and successional status. In: ANL/LRP-2. Annual report July 1976-October 1977. Argonne, IL: Argonne National Laboratory, Energy & Environmental Systems Division, Land Reclamation Program: 132-138. [5729]

47. Ellison, Lincoln; Woolfolk, E. J. 1937. Effects of drought on vegetation near Miles City, Montana. Ecology. 18(3): 329-336. [6264]

48. Erichsen-Arychuk, Catherine; Bork, Edward W.; Bailey, Arthur W. 2002. Northern dry mixed prairie responses to summer wildfire and drought. Journal of Range Management. 55(2): 164-170. [40694]

49. Evans, Raymond A.; Young, James A. 1982. Microhabitat variation in relation to weed seed germination and seedling emergence. In: Hatfield, Jerry L.; Thomason, Ivan J., eds. Biometerology in integrated pest management. New York: Academic Press: 421-448. [42759]

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

51. Fairaizl, Steven D. 1978. Bighorn sheep in North Dakota: population estimates, food habits and their biogeochemistry. Grand Forks, ND: University of North Dakota. 83 p. Thesis. [Project No.: W-67-R-17. Bismark, ND: North Dakota State Game and Fish Department. 51 p. + appendices]. [25841]

52. Fisher, R. F.; Jenkins, M. J.; Fisher, William F. 1986. Fire and the prairie-forest mosaic of Devils Tower National Monument. The American Midland Naturalist. 117(2): 250-257. [15638]

53. Fisser, Herbert G. 1981. Shrub establishment, dominance, and ecology on the juniper and sagebrush-grass types in Wyoming. In: Shrub establishment on disturbed arid and semi-arid lands: Proceedings of the symposium; 1980 December 2-3; Laramie, WY. Laramie, WY: Wyoming Game and Fish Department: 23-28. [926]

54. Francis, Richard E. 1983. Sagebrush-steppe habitat types in northern Colorado: a first approximation. In: Moir, W. H.; Hendzel, Leonard, tech. coords. Proceedings of the workshop on Southwestern habitat types; 1983 April 6-8; Albuquerque, NM. Abluquerque, NM: U.S. Department of Agriculture, Forest Service, Southwestern Region: 67-71. [955]

55. 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]

56. Gilmore, Melvin Randolph. 1919. Uses of plants by the Indians of the Missouri River region. In: 33rd annual report of the Bureau of American Ethnology. Washington, DC: Bureau of American Ethnology: 44-154. [6928]

57. Great Plains Flora Association. 1986. Flora of the Great Plains. Lawrence, KS: University Press of Kansas. 1392 p. [1603]

58. Gruell, G. E.; Loope, L. L. 1974. Relationships among aspen, fire, and ungulate browsing in Jackson Hole, Wyoming. Lakewood, CO: U.S. Department of the Interior, National Park Service, Rocky Mountain Region. 33 p. In cooperation with: U.S. Department of Agriculture, Forest Service, Intermountain Region. [3862]

59. Gullion, Gordon W. 1964. Contributions toward a flora of Nevada. No. 49: Wildlife uses of Nevada plants. CR-24-64. Beltsville, MD: U.S. Department of Agriculture, Agricultural Research Service, National Arboretum Crops Research Division. 170 p. [6729]

60. Gutknecht, Kurt W. 1989. Xeriscaping: an alternative to thirsty landscapes. Utah Science. 50(4): 142-146. [10166]

61. Hall, James B.; Hansen, Paul L. 1997. A preliminary riparian habitat type classification system for the Bureau of Land Management districts in southern and eastern Idaho. Tech. Bull. No. 97-11. Boise, ID: U.S. Department of the Interior, Bureau of Land Management; Missoula, MT: University of Montana, School of Forestry, Riparian and Wetland Research Program. 381 p. [28173]

62. Hansen, Paul L.; Boggs, Keith; Pfister, Robert D.; [and others]. 1994. Classification and management of riparian and wetland sites in Montana. In: Hamre, R. H., ed. Workshop on western wetlands and riparian areas: public/private efforts in recovery, management, and education: Proceedings; 1993 September 9-11; Snowbird, UT. Boulder, CO: Thorne Ecological Institute: 1-17. [27800]

63. Hansen, Paul L.; Chadde, Steve W.; Pfister, Robert D. 1988. Riparian dominance types of Montana. Misc. Publ. No. 49. Missoula, MT: University of Montana, School of Forestry, Montana Forest and Conservation Experiment Station. 411 p. [5660]

64. Hansen, Paul L.; Hoffman, George R. 1988. The vegetation of the Grand River/Cedar River, Sioux, and Ashland Districts of the Custer National Forest: a habitat type classification. Gen. Tech. Rep. RM-157. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 68 p. [771]

65. Hansen, Paul L.; Hoffman, George R.; Bjugstad, Ardell J. 1984. The vegetation of Theodore Roosevelt National Park, North Dakota: a habitat type classification. Gen. Tech. Rep. RM-113. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 35 p. [1077]

66. Hansen, Paul L.; Pfister, Robert D.; Boggs, Keith; [and others]. 1995. Classification and management of Montana's riparian and wetland sites. Miscellaneous Publication No. 54. Missoula, MT: The University of Montana, School of Forestry, Montana Forest and Conservation Experiment Station. 646 p. [24768]

67. Hansen, Paul; Boggs, Keith; Pfister, Robert; Joy, John. 1990. Classification and management of riparian and wetland sites in central and eastern Montana. Missoula, MT: University of Montana, School of Forestry, Montana Forest and Conservation Experiment Station, Montana Riparian Association. 279 p. [12477]

68. Hansen, Paul; Pfister, Robert; Joy, John; [and others]. 1989. Classification and management of riparian sites in southwestern Montana. Missoula, MT: University of Montana, School of Forestry, Montana Riparian Association. 292 p. Draft Version 2. [8900]

69. Hanson, Herbert C.; Whitman, Warren. 1938. Characteristics of major grassland types in western North Dakota. Ecological Monographs. 8(2): 57-114. [15]

70. Hargis, Christina; McCarthy, Clinton. 1986. Vegetation changes following a prescribed burn on a Great Basin meadow. Transactions, Western Section of the Wildlife Society. 22: 47-51. [15955]

71. Harvey, Stephen John. 1981. Life history and reproductive strategies in Artemisia. Bozeman, MT: Montana State University. 132 p. M.S. thesis. [1102]

72. Hayes, Doris W.; Garrison, George A. 1960. Key to important woody plants of eastern Oregon and Washington. Agric. Handb. 148. Washington, DC: U.S. Department of Agriculture, Forest Service. 227 p. [1109]

73. Hazlett, Donald L.; Hoffman, George R. 1975. Plant species distributional patterns in Artemisia tridentata- and Artemisia cana-dominated vegetation in western North Dakota. Botanical Gazette. 136(1): 72-77. [1111]

74. Heyerdahl, Emily K.; Berry, Dawn; Agee, James K. 1994. Fire history database of the western United States. Final report. Interagency agreement: U.S. Environmental Protection Agency DW12934530; U.S. Department of Agriculture, Forest Service PNW-93-0300; University of Washington 61-2239. Seattle, WA: U.S. Department of Agriculture, Pacific Northwest Research Station; University of Washington, College of Forest Resources. 28 p. [+ Appendices]. Unpublished report on file with: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. [27979]

75. Hickman, James C., ed. 1993. The Jepson manual: Higher plants of California. Berkeley, CA: University of California Press. 1400 p. [21992]

76. Higgins, Kenneth F. 1984. Lightning fires in North Dakota grasslands and in pine-savanna lands of South Dakota and Montana. Journal of Range Management. 37(2): 100-103. [1148]

77. Hironaka, M.; Fosberg, M. A.; Winward, A. H. 1983. Sagebrush-grass habitat types of southern Idaho. Bulletin Number 35. Moscow, ID: University of Idaho, Forest, Wildlife and Range Experiment Station. 44 p. [1152]

78. Hirsch, Kathie Jean. 1985. Habitat classification of grasslands and shrublands of southwestern North Dakota. Fargo, ND: North Dakota State University. 281 p. Dissertation. [40326]

79. Hoffman, G. R.; Hazlett, D. L. 1977. Effects of aqueous Artemisia extracts and volatile substances on germination of selected species. Journal of Range Management. 30(2): 134-137. [23850]

80. Hopkins, Rick B. 1984. Avian species associated with prairie woodland types. In: Noble, Daniel L.; Winokur, Robert P., eds. Wooded draws: characteristics and values for the northern Great Plains: Proceedings of a symposium; 1984 June 12-13; Rapid City, SD. Great Plains Agricultural Council Publication No. 111. Rapid City, SD: South Dakota School of Mines and Technology, Biology Department: 27-35. [1192]

81. Hormay, August L.; Alberico, Fred J.; Lord, P. B. 1962. Experiences with 2,4-D spraying on the Lassen National Forest. Journal of Range Management. 15(6): 325-328. [38665]

82. Hou, Junqiang; Romo, J. T. 1998. Cold-hardiness of silver sagebrush seedlings. Journal of Range Management. 51(6): 704-708. [41531]

83. Hou, Junqiang; Romo, J. T. 1998. Seed weight and germination time affect growth of 2 shrubs. Journal of Range Management. 51(6): 699-703. [41532]

84. Houston, Douglas B. 1973. Wildfires in northern Yellowstone National Park. Ecology. 54(5): 1111-1117. [5781]

85. Houston, Kent E.; Hartung, Walter J.; Hartung, Carol J. 2001. A field guide for forest indicator plants, sensitive plants, and noxious weeds of the Shoshone National Forest, Wyoming. Gen. Tech. Rep. RMRS-GTR-84. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. 184 p. [40585]

86. Howard, Gene S.; Rauzi, Frank; Schuman, Gerald E. 1979. Woody plant trials at six mine reclamation sites in Wyoming and Colorado. Production Research Report PRR 177/1/79. Washington, DC: U.S. Department of Agriculture. 14 p. [42428]

87. Howe, Henry F. 1994. Managing species diversity in tallgrass prairie: assumptions and implications. Conservation Biology. 8(3): 691-704. [26692]

88. Jakubos, Bonnie; Romme, William H. 1993. Invasion of subalpine meadows by lodgepole pine in Yellowstone National Park, Wyoming, U.S.A. Arctic and Alpine Research. 25(4): 382-390. [22582]

89. Johnson, James R.; Nichols, James T. 1970. Plants of South Dakota grasslands: A photographic study. Bull. 566. Brookings, SD: South Dakota State University, Agricultural Experiment Station. 163 p. [18500]

90. Johnson, Kendall L. 1979. Basic synecological relationships of the sagebrush types on the high plains of Montana, Wyoming and the Dakotas. In: The sagebrush ecosystem: a symposium: Proceedings; 1978 April; Logan, UT. Logan, UT: Utah State University, College of Natural Resources: 42-49. [1281]

91. Johnson, Kendall L. 1987. Sagebrush types as ecological indicators to integrated pest management (IPM) in the sagebrush ecosystem of western North America. In: Onsager, Jerome A., ed. Integrated pest management on rangeland: State-of-the-art in the sagebrush ecosystem. ARS-50. Washington, DC: U.S. Department of Agriculture, Agricultural Research Service: 1-10. [2841]

92. Kartesz, John T.; Meacham, Christopher A. 1999. Synthesis of the North American flora (Windows Version 1.0), [CD-ROM]. Available: North Carolina Botanical Garden. In cooperation with the Nature Conservancy, Natural Resources Conservation Service, and U.S. Fish and Wildlife Service [2001, January 16]. [36715]

93. Kartesz, John Thomas. 1988. A flora of Nevada. Reno, NV: University of Nevada. 1729 p. [In 3 volumes]. Dissertation. [42426]

94. Knapp, Paul A. 1991. The response of semi-arid vegetation assemblages following the abandonment of mining towns in southwestern Montana. Journal of Arid Environments. 20: 205-222. [14894]

95. Knowles, Craig J. 1986. Some relationships of black-tailed prairie dogs to livestock grazing. The Great Basin Naturalist. 46(2): 198-203. [7530]

96. Kornkven, Amy B.; Watson, Linda E.; Estes, James R. 1998. Phylogenetic analysis of Artemisia section Tridentatae (Asteraceae) based on sequences from the internal transcribed spacers (ITS) of nuclear ribosomal DNA. American Journal of Botany. 85(2): 1787-1795. [42433]

97. Kornkven, Amy B.; Watson, Linda E.; Estes, James R. 1999. Molecular phylogeny of Artemisia section Tridentatae (Asteraceae) based on chloroplast DNA restriction site variation. Systematic Botany. 24(1): 69-84. [42432]

98. Kovalchik, Bernard L. 1987. Riparian zone associations: Deschutes, Ochoco, Fremont, and Winema National Forests. R6 ECOL TP-279-87. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 171 p. [9632]

99. Kovalchik, Bernard L.; Chitwood, Lawrence A. 1990. Use of geomorphology in the classification of riparian plant associations in mountainous landscapes of central Oregon, U.S.A. Forest Ecology and Management. 33/34: 405-418. [6830]

100. Kovalchik, Bernard L.; Hopkins, William E.; Brunsfeld, Steven J. 1988. Major indicator shrubs and herbs in riparian zones on national forests of central Oregon. R6-ECOL-TP-005-88. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 159 p. [8995]

101. Kuchler, A. W. 1964. United States [Potential natural vegetation of the conterminous United States]. Special Publication No. 36. New York: American Geographical Society. 1:3,168,000; colored. [3455]

102. Kufeld, Roland C. 1983. Responses of elk, mule deer, cattle, and vegetation to burning, spraying and chaining of Gambel oak rangeland. Tech. Publ. 34. Fort Collins, CO: Colorado Division of Wildlife. 47 p. [253]

103. Kufeld, Roland C.; Wallmo, O. C.; Feddema, Charles. 1973. Foods of the Rocky Mountain mule deer. Res. Pap. RM-111. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 31 p. [1387]

104. Laven, R. D.; Omi, P. N.; Wyant, J. G.; Pinkerton, A. S. 1980. Interpretation of fire scar data from a ponderosa pine ecosystem in the central Rocky Mountains, Colorado. In: Stokes, Marvin A.; Dieterich, John H., technical coordinators. Proceedings of the fire history workshop; 1980 October 20-24; Tucson, AZ. Gen. Tech. Rep. RM-81. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 46-49. [7183]

105. Leach, Howard R.; Hensley, Arthur L. 1954. The sage grouse in California with special reference to food habits. California Fish and Game. 40: 385-394. [42430]

106. Leckenby, Donavin A. 1978. Mule deer occupancy of plant communities on a south-central Oregon winter range. Job Final Report. Research Project Number: W-70-R. July 1, 1965 to June 30, 1976. Portland, OR: Oregon Department of Fish and Wildlife. 81 p. [12496]

107. Lewis, James K.; Van Dyne, George M.; Albee, Leslie R.; Whetzal, Frank W. 1956. Intensity of grazing: Its effect on livestock and forage production. Bulletin 459. Brookings, SD: South Dakota State College, Agricultural Experiment Station. 44 p. [11737]

108. Mackie, Richard J. 1970. Range ecology and relations of mule deer, elk, and cattle in the Missouri River Breaks, Montana. Wildlife Monographs No. 20. 79 p. [5897]

109. Manning, Mary E.; Padgett, Wayne G. 1989. Preliminary riparian community type classification for Nevada. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Region. 135 p. Preliminary draft. [11531]

110. Manning, Mary E.; Padgett, Wayne G. 1995. Riparian community type classification for Humboldt and Toiyabe National Forests, Nevada and eastern California. R4-Ecol-95-01. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Region. 306 p. [42196]

111. Martin, William C.; Hutchins, Charles R. 1981. A flora of New Mexico. Volume 2. Germany: J. Cramer. 2589 p. [37176]

112. McArthur, E. Durant. 1994. Ecology, distribution, and values of sagebrush within the Intermountain Region. In: Monsen, Stephen B.; Kitchen, Stanley G., compilers. Proceedings--ecology and management of annual rangelands; 1992 May 18-22; Boise, ID. Gen. Tech. Rep. INT-GTR-313. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 347-351. [24308]

113. McArthur, E. Durant. 2000. Sagebrush systematics and distribution. In: Entwistle, P. G.; DeBolt, A. M.; Kaltenecker, J. H.; Steenhof, K., compilers. In: Sagebrush steppe ecosystems symposium: Proceedings; 1999 June 21-23; Boise, ID. Publ. No. BLM/ID/PT-001001+1150. Boise, ID: U.S. Department of the Interior, Bureau of Land Management, Boise State Office: 9-14. [41811]

114. McArthur, E. Durant; Blauer, A. Clyde; Plummer, A. Perry; Stevens, Richard. 1979. Characteristics and hybridization of important Intermountain shrubs. III. Sunflower family. Res. Pap. INT-220. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 82 p. [1571]

115. McArthur, E. Durant; Mudge, Joann; Van Buren, Renee; [and others]. 1998. Randomly amplified polymorphic DNA analysis (RAPD) of Artemisia subgenus Tridentatae species and hybrids. The Great Basin Naturalist. 58(1): 12-27. [28609]

116. McArthur, E. Durant; Pope, C. Lorenzo; Freeman, D. Carl. 1981. Chromosomal studies of subgenus Tridentatae of Artemisia: evidence for autopolyploidy. American Journal of Botany. 68(5): 589-605. [42431]

117. McArthur, E. Durant; Sanderson, Stewart C. 1999. Cytogeography and chromosome evolution of subgenus Tridentatae of Artemisia (Asteraceae). American Journal of Botany. 86(12): 1754-1775. [34931]

118. McArthur, E. Durant; Sanderson, Stewart C. 1999. Ecotones: introduction, scale, and big sagebrush example. In: McArthur, E. Durant; Ostler, W. Kent; Wambolt, Carl L., compilers. Proceedings: shrub ecotones; 1998 August 12-14; Ephraim, UT. Proceedings RMRS-P-11. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 3-8. [36053]

119. McArthur, E. Durant; Stevens, Richard. 1986. Composite shrubs. Unpublished manuscript on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. 155 p. [7342]

120. McArthur, E. Durant; Van Buren, Renee; Sanderson, Stewart C.; Harper, Kimball T. 1998. Taxonomy of Sphaeromoeria, Artemisia, and Tanacetum (Compositae, Anthemideae) based on randomly amplified polymorphic DNA (RAPD). The Great Basin Naturalist. 58(1): 1-11. [28610]

121. Meinecke, E. P. 1929. Quaking aspen: A study in applied forest pathology. Tech. Bull. No. 155. Washington, DC: U.S. Department of Agriculture. 34 p. [26669]

122. Miller, Richard F.; Rose, Jeffery A. 1995. Historic expansion of Juniperus occidentalis (western juniper) in southeastern Oregon. The Great Basin Naturalist. 55(1): 37-45. [26637]

123. Moerman, Dan. 1999. Native American ethnobotany database: Foods, drugs, dyes, and fibers of native North American peoples, [Online]. Available: http://www.umd.umich.edu/cgi-bin/herb/ [2002, December 5]. [37492]

124. Monsen, Stephen B. 1994. Selection of plants for fire suppression on semiarid sites. In: Monsen, Stephen B.; Kitchen, Stanley G., compilers. Proceedings--ecology and management of annual rangelands; 1992 May 18-22; Boise, ID. Gen. Tech. Rep. INT-GTR-313. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 363-373. [24310]

125. Montana State University, Montana Agricultural Experiment Station. 1973. Vegetative rangeland types in Montana. Bulletin 671. Bozeman, MT: Montana State University, Montana Agricultural Experiment Station. 15 p. [29827]

126. Mozingo, Hugh N. 1987. Shrubs of the Great Basin: A natural history. Reno, NV: University of Nevada Press. 342 p. [1702]

127. Nelson, Jack Raymond. 1961. Composition and structure of the principal woody vegetation types in the North Dakota Badlands. Fargo, ND: North Dakota State University. 195 p. Thesis. [161]

128. Nevada Department of Conservation and Natural Resources, Natural Heritage Program. 2001. Focal communities for Nevada [Online]. Available: http://www.state.nv.us/nvnhp/ecology/focus.htm [2002, September 13]. [42008]

129. Norland, Jack E.; Marlow, Clayton B. 1984. Use of wooded draws by free-roaming bison. In: Noble, Daniel L.; Winokur, Robert P., eds. Wooded draws: Characteristics and values for the Northern Great Plains: Symposium proceedings; 1984 June 12-13; Rapid City, SD. Great Plains Agricultural Council Publication No. 111. Rapid City, SD: South Dakota School of Mines and Technology, Biology Department: 40-44. [1777]

130. Olson, R. A.; Gerhart, W. A. 1982. A physical and biological characterization of riparian habitat and its importance to wildlife in Wyoming. Cheyenne, WY: Wyoming Game and Fish Department. 188 p. [6755]

131. Padgett, Wayne G.; Youngblood, Andrew P.; Winward, Alma H. 1989. Riparian community type classification of Utah and southeastern Idaho. R4-Ecol-89-01. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Region. 191 p. [11360]

132. Padgett, Wayne George. 1981. Ecology of riparian plant communities in southern Malheur National Forest. Corvallis, OR: Oregon State University. 143 p. Thesis. [14933]

133. Paysen, Timothy E.; Ansley, R. James; Brown, James K.; [and others]. 2000. Fire in western shrubland, woodland, and grassland ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-volume 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 121-159. [36978]

134. Pechanec, Joseph F.; Plummer, A. Perry; Robertson, Joseph H.; Hull, A. C., Jr. 1965. Sagebrush control on rangelands. Agriculture Handbook No. 277. Washington, DC: U.S. Department of Agriculture. 40 p. [1858]

135. Pfadt, R. E. 1994. Species fact sheet--Sagebrush grasshopper (Melanoplus bowditchi Scudder). In: Field guide to common western grasshoppers. Bulletin 912. Laramie, WY: University of Wyoming, Agricultural Experiment Station. Available: http://www.sdvc.uwyo.edu/grasshopper/fieldgde.htm [2002, October 1]. [42042]

136. Platts, William S.; Armour, Carl; Booth, Gordon D.; [and others]. 1987. Methods for evaluating riparian habitats with applications to management. Gen. Tech. Rep. INT-221. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 177 p. [6171]

137. Quinnild, Clayton L.; Cosby, Hugh E. 1958. Relicts of climax vegetation on two mesas in western North Dakota. Ecology. 39(1): 29-32. [1925]

138. Rainier Seeds, Inc. 2002. [Homepage of Rainer Seeds, Inc], [Online]. Available: http://www.rainierseeds.com [2002, November 6]. [42441]

139. Ralston, Robert Dean. 1960. The structure and ecology of the north slope juniper stands of the Little Missouri Badlands. Salt Lake City, UT: University of Utah. 85 p. Thesis. [192]

140. Rasmussen, D. I.; Griner, Lynn A. 1938. Life history and management studies of the sage grouse in Utah, with special reference to nesting and feeding habits. Transactions, North American Wildlife Conference. 3: 852-864. [42197]

141. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843]

142. Ream, Robert Ray. 1964. The vegetation of the Wasatch Mountains, Utah and Idaho. Madison, WI: University of Wisconsin. 178 p. Dissertation. [5506]

143. Reed, John F. 1952. The vegetation of the Jackson Hole Wildlife Park, Wyoming. The American Midland Naturalist. 48(3): 700-729. [1949]

144. Reed, Merton J.; Peterson, Roald A. 1961. Vegetation, soil, and cattle responses to grazing on northern Great Plains range. Tech. Bull. 1252. Washington, DC: U.S. Department of Agriculture, Forest Service. 79 p. [4286]

145. Ricketts, Matt. 1994. Cutting ranching costs: optimizing forage protein value. Rangelands. 16(6): 260-264. [29726]

146. Romo, J. T.; Eddleman, L. E. 1995. Use of degree-days in multiple-temperature experiments. Journal of Range Management. 48(5): 410-416. [41530]

147. Romo, J. T.; Grilz, R. W. 2002. Establishment of silver sagebrush in the Northern Mixed Prairie. Journal of Range Management. 55(3): 217-221. [41755]

148. Romo, James T.; Young, James A. 2002. Temperature profiles and the effects of field experiments on germination of silver sagebrush. Native Plant Journal. 3: 5-13. [42758]

149. Rosentreter, Roger. 1992. Camas prairie and possible evolutionary links with old world Artemisia species: a presymposium tour. In: Clary, Warren P.; McArthur, E. Durant; Bedunah, Don; Wambolt, Carl L., compilers. Proceedings--symposium on ecology and management of riparian shrub communities; 1991 May 29-31; Sun Valley, ID. Gen. Tech. Rep. INT-289. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 223-227. [19126]

150. Ross, Robert L.; Hunter, Harold E. 1976. Climax vegetation of Montana: Based on soils and climate. Bozeman, MT: U.S. Department of Agriculture, Soil Conservation Service. 64 p. [2028]

151. Rowe, J. S. 1969. Lightning fires in Saskatchewan grassland. Canadian Field-Naturalist. 83: 317-324. [6266]

152. Rupp, Larry; Kjelgren, Roger; Ernstsen, Jerriann; Varga, William. 1997. Shearing and growth of five Intermountain native shrub species. Journal of Environmental Horticulture. 15(3): 123-125. [29195]

153. Salwasser, Hal; Shimamoto, Karen. 1984. Pronghorn, cattle, and feral horse use of wetland and upland habitats. In: Warner, Richard E.; Hendrix, Kathleen M., eds. California riparian systems: Ecology, conservation, and productive management: Proceedings of a conference; 1981 September 17-19; Davis, CA. Berkeley, CA: University of California Press: 210-213. [5836]

154. Sapsis, David B. 1990. Ecological effects of spring and fall prescribed burning on basin big sagebrush/Idaho fescue--bluebunch wheatgrass communities. Corvallis, OR: Oregon State University. 105 p. Thesis. [16579]

155. Schlatterer, Edward F. 1972. A preliminary description of plant communities found on the Sawtooth, White Cloud, Boulder and Pioneer Mountains. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Region. Unpublished paper on file with: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. 111 p. [2076]

156. Schneider, R. E.; Faber-Langendoen, D.; Crawford, R. C.; Weakley, A. S. 1997. Shrubland (scrub): Microphyllous evergreen shrubland. In: Great Plains vegetation classification. Supplemental Document 1. In: Ostlie, W. R.; Schneider, R. E.; Aldrich, J. M.; Faust, T. M.; McKim, R. L. B.; Chaplin, S. J., eds. The status of biodiversity in the Great Plains, [Online]. Arlington, VA: The Nature Conservancy (Producer). Available: http://www.greatplains.org/resource/biodiver/gpclass/iiia4.htm [2002, September 17]. [42009]

157. Schultz, Brad; McAdoo, Kent. 2002. Common sagebrush in Nevada. Special Publication SP-02-02. Reno, NV: University of Nevada, Cooperative Extension. 9 p. Available: http://www.unce.unr.edu/publications/Spec%20Pubs/SP-02-02.doc [2002, October 1]. [42043]

158. Severson, Kieth E.; Boldt, Charles E. 1977. Problems associated with management of native woody plants in the western Dakotas. In: Johnson, Kendall L., editor. Wyoming shrublands: Proceedings, 6th Wyoming shrub ecology workshop; 1977 May 24-25; Buffalo, WY. Laramie, WY: Shrub Ecology Workshop: 51-57. [2759]

159. Shaw, Nancy L.; Monsen, Stephen B. 1990. Use of sagebrush for improvement of wildlife habitat. In: Fisser, Herbert G., ed. Wyoming shrublands: Aspen, sagebrush and wildlife management: Proceedings, 17th Wyoming shrub ecology workshop; 1988 June 21-22; Jackson, WY. Laramie, WY: University of Wyoming, Department of Range Management; Shrub Ecology Workshop 19-35. [22929]

160. Shaw, Nancy; Sands, Alan; Turnipseed, Dale. 1984. Potential use of fourwing saltbush and other dryland shrubs for upland game bird cover in southern Idaho. 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: 206-214. [2125]

161. Sheehy, Dennis P.; Winward, A. H. 1981. Relative palatability of seven Artemisia taxa to mule deer and sheep. Journal of Range Management. 34(5): 397-399. [2128]

162. Shiflet, Thomas N., ed. 1994. Rangeland cover types of the United States. Denver, CO: Society for Range Management. 152 p. [23362]

163. Shultz, Leila M. 1986. Comparative leaf anatomy of sagebrush: ecological considerations. In: McArthur, E. Durant; Welch, Bruce L., compilers. Proceedings--symposium on the biology of Artemisia and Chrysothamnus; 1984 July 9-13; Provo, UT. Gen. Tech. Rep. INT-200. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 253-264. [2140]

164. Shultz, Leila M. 1986. Taxonomic and geographic limits of Artemisia subgenus Tridentatae (Beetle). In: McArthur, E. Durant; Welch, Bruce L., compilers. Proceedings--symposium on the biology of Artemisia and Chrysothamnus; 1984 July 9-13; Provo, UT. Gen. Tech. Rep. INT-200. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 20-28. [2141]

165. Sieg, Carolyn Hull. 1997. The role of fire in managing for biological diversity on native rangelands of the Northern Great Plains. In: Uresk, Daniel W.; Schenbeck, Greg L.; O'Rourke, James T., tech. coords. Conserving biodiversity on native rangelands: symposium proceedings; 1995 August 17; Fort Robinson State Park, NE. Gen. Tech. Rep. RM-GTR-298. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 31-38. [28054]

166. Smith, Justin G. 1952. Food habits of mule deer in Utah. Journal of Wildlife Management. 16(2): 148-154. [2174]

167. Smoliak, S.; Ditterline, R. L.; Scheetz, J. D.; Holzworth, L. K.; Sims, J. R.; Wiesner, L. E.; Baldridge, D. E.; Tibke, G. L. 2001. Montana Interagency Plant Materials Handbook. EB69 [Online]. Bozeman, MT: Montana State University, Animal and Range Sciences, Extension Service (Producer). Available: http://animalrangeextension.montana.edu/Articles/Forage/Species/Shrubs/Silversagebrush.htm [2002, September 20]. [42052]

168. Stephens, H. A. 1973. Woody plants of the North Central Plains. Lawrence, KS: The University Press of Kansas. 530 p. [3804]

169. Stickney, Peter F. 1989. Seral origin of species originating in northern Rocky Mountain forests. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. 10 p. [20090]

170. Stoms, David M.; Davis, Frank W.; Driese, Kenneth L.; [and others]. 1998. Gap analysis of the vegetation of the Intermountain semi-desert ecoregion. The Great Basin Naturalist. 58(3): 199-216. [30151]

171. Stubbendieck, James; Hatch, Stephan L.; Butterfield, Charles H. 1992. North American range plants. 4th ed. Lincoln, NE: University of Nebraska Press. 493 p. [25162]

172. Sundstrom, Charles; Hepworth, William G.; Diem, Kenneth L. 1973. Abundance, distribution and food habits of the pronghorn: A partial characterization of the optimum pronghorn habitat. Bulletin No. 12. Cheyenne, WY: Wyoming Game and Fish Commission. 59 p. [5906]

173. Svejcar, Tony J.; Riegel, Gregg M.; Conroy, Scott D.; Trent, James D. 1992. Establishment and growth potential of riparian shrubs in the northern Sierra Nevada. In: Clary, Warren P.; McArthur, E. Durant; Bedunah, Don; Wambolt, Carl L., compilers. Proceedings--symposium on ecology and management of riparian shrub communities; 1991 May 29-31; Sun Valley, ID. Gen. Tech. Rep. INT-289. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 151-154. [19110]

174. Swenson, Jon E. 1985. Seasonal habitat use by sharp-tailed grouse, Tympanuchus phasianellus, on mixed-grass prairie in Montana. Canadian Field-Naturalist. 99(1): 40-46. [23501]

175. Terwilliger, Charles, Jr.; Tiedeman, James A. 1978. Habitat types of the mule deer critical winter range and adjacent steppe region of Middle Park, Colorado. Final Report Cooperative Agreement No. 16-739-CA. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 108 p. [5611]

176. Thatcher, Albert P. 1959. Distribution of sagebrush as related to site differences in Albany County, Wyoming. Journal of Range Management. 12(2): 55-61. [2314]

177. Thilenius, John F.; Brown, Gary R.; Medina, Alvin L. 1995. Vegetation on semi-arid rangelands, Cheyenne River Basin, Wyoming. Gen. Tech. Rep. RM-GTR-263. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 60 p. [26478]

178. Tiedeman, James A.; Francis, Richard E.; Terwilliger, Charles, Jr.; Carpenter, Len H. 1987. Shrub-steppe habitat types of Middle Park, Colorado. Res. Pap. RM-273. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 20 p. [2329]

179. Tisdale, E. W.; Hironaka, M. 1981. The sagebrush-grass region: a review of the ecological literature. Bull. 33. Moscow, ID: University of Idaho, Forest, Wildlife and Range Experiment Station. 31 p. [2344]

180. U.S. Department of Agriculture, National Resource Conservation Service. 2003. PLANTS database (2003), [Online]. Available: http://plants.usda.gov/. [34262]

181. Urness, Philip Joel. 1966. Influence of range improvement practices on composition, production, and utilization of Artemisia deer winter range in central Oregon. Corvallis, OR: Oregon State University. 182 p. Dissertation. [3060]

182. Valles, Joan; McArthur, E. Durant. 2001. Artemisia systematics and phylogeny: cytogenetic and molecular insights. In: McArthur, E. Durant; Fairbanks, Daniel J., compilers. Shrubland ecosystem genetics and biodiversity: proceedings; 2000 June 13-15; Provo, UT. Proc. RMRS-P-21. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 67-74. [41956]

183. Vincent, Dwain W. 1992. The sagebrush/grasslands of the upper Rio Puerco area, New Mexico. Rangelands. 14(5): 268-271. [19698]

184. Vories, Kimery C. 1981. Growing Colorado plants from seed: a state of the art. Volume I: Shrubs. Gen. Tech. Rep. INT-103. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 80 p. [3426]

185. Walton, Todd P.; White, Richard S.; Wambolt, Carl L. 1986. Artemisia reproductive strategies: a review with emphasis on plains silver sagebrush. In: McArthur, E. Durant; Welch, Bruce L., compilers. Proceedings--symposium on the biology of Artemisia and Chrysothamnus; 1984 July 9-13; Provo, UT. Gen. Tech. Rep. INT-200. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 67-74. [2447]

186. Walton, Todd Patrick. 1984. Reproductive mechanisms of plains silver sagebrush Artemisia cana cana in southeastern Montana. Bozeman, MT: Montana State University. 161 p. Thesis. [100]

187. Wambolt, Carl L.; Walton, Todd; White, Richard S. 1989. Seed dispersal characteristics of plains silver sagebrush. Prairie Naturalist. 21(3): 113-118. [15530]

188. Ward, George H. 1953. Artemisia, section Seriphidium, in North America: a cytotaxonomic study. Contributions from the Dudley Herbarium. 4(6): 155-205. [2454]

189. Wasser, Clinton H. 1982. Ecology and culture of selected species useful in revegetating disturbed lands in the West. FWS/OBS-82/56. Washington, DC: U.S. Department of the Interior, Fish and Wildlife Service. 347 p. [4837]

190. Weber, William A.; Wittmann, Ronald C. 1996. Colorado flora: eastern slope. 2nd ed. Niwot, CO: University Press of Colorado. 524 p. [27572]

191. Welsh, Stanley L.; Atwood, N. Duane; Goodrich, Sherel; Higgins, Larry C., eds. 1987. A Utah flora. The Great Basin Naturalist Memoir No. 9. Provo, UT: Brigham Young University. 894 p. [2944]

192. Wendtland, Kyle J.; Dodd, Jerrold L. 1992. The fire history of Scotts Bluff National Monument. In: Smith, Daryl D.; Jacobs, Carol A., eds. Recapturing a vanishing heritage: Proceedings, 12th North American prairie conference; 1990 August 5-9; Cedar Falls, IA. Cedar Falls, IA: University of Northern Iowa: 141-143. [24731]

193. West, Neil E. 1988. Intermountain deserts, shrub steppes, and woodlands. In: Barbour, Michael G.; Billings, William Dwight, eds. North American terrestrial vegetation. Cambridge; New York: Cambridge University Press: 209-230. [19546]

194. White, Richard S.; Currie, Pat O. 1983. Response of silver sagebrush to burning, mechanical and chemical control practices. Bulletin of the Ecological Society of America. 64(2): 110; 1983. [2539]

195. White, Richard S.; Currie, Pat O. 1983. The effects of prescribed burning on silver sagebrush. Journal of Range Management. 36(5): 611-613. [2540]

196. Whitson, T. D.; Nix, K. J.; Cox, Roger. 1989. Control of silver sagebrush (Artemisia cana) in rangeland. Western Society of Weed Science. Research Progress Report: 144. [41552]

197. Winward, Alma H. 1980. Taxonomy and ecology of sagebrush in Oregon. Station Bulletin 642. Corvallis, OR: Oregon State University, Agricultural Experiment Station. 15 p. [2585]

198. Winward, Alma H. 2001. Sagebrush taxonomy and ecology workshop; 1999 October 5-6; Logan, UT [Online]. Logan, UT: U.S. Department of Agriculture, Forest Service, Region 4, Wasatch-Cache National Forest (Producer). Available: http://www.fs.fed.us/wcnf/unit/eco/sagebrush_workshop/sagebrush_ecology.htm [2002, October 3]. [42051]

199. Wright, Henry A.; Bailey, Arthur W. 1982. Fire ecology: United States and southern Canada. New York: John Wiley & Sons. 501 p. [2620]

200. Wright, Henry A.; Neuenschwander, Leon F.; Britton, Carlton M. 1979. The role and use of fire in sagebrush-grass and pinyon-juniper plant communities: A state-of-the-art review. Gen. Tech. Rep. INT-58. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 48 p. [2625]

201. Young, James A.; Evans, Raymond A. 1981. Demography and fire history of a western juniper stand. Journal of Range Management. 34(6): 501-505. [2659]

202. Young, James A.; Evans, Raymond A.; Major, Jack. 1977. Sagebrush steppe. In: Barbour, Michael G.; Major, Jack, eds. Terrestrial vegetation of California. New York: John Wiley & Sons: 763-796. [4300]

203. Youngblood, Andrew P.; Padgett, Wayne G.; Winward, Alma H. 1985. Riparian community type classification of eastern Idaho - western Wyoming. R4-Ecol-85-01. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Region. 78 p. [2686]




FEIS Home Page