Centrocercus minimus, C. urophasianus

• INTRODUCTORY
• WILDLIFE DISTRIBUTION AND OCCURRENCE
• BIOLOGICAL DATA AND HABITAT REQUIREMENTS
• FIRE EFFECTS AND USE
• FIRE CASE STUDIES
• REFERENCES

INTRODUCTORY

AUTHORSHIP AND CITATION FEIS ABBREVIATION SYNONYMS COMMON NAMES TAXONOMY ORDER/CLASS FEDERAL LEGAL STATUS OTHER STATUS
	greater sage-grouse © Jessica Young	Gunnison sage-grouse © Louis Swift

In this report "sage-grouse" refers to both greater sage-grouse and Gunnison sage-grouse. Where the literature refers to a subspecies of greater sage-grouse or to Gunnison sage-grouse, they will be identified as such.

There is no evidence that the 2 species of sage-grouse interbreed [26], but sage-grouse do rarely hybridize with both sharp-tailed grouse (Tympanuchus phasianellus) and dusky grouse (Dendragapus obscurus) [65]. Hybrids of sharp-tailed grouse and greater sage-grouse have been found in Montana [44], Saskatchewan [61], and North Dakota [77]. A hybrid of dusky grouse and sage-grouse was found in Utah [100].

ORDER:
Galliformes

CLASS:
Bird

FEDERAL LEGAL STATUS:
As of this writing (2002), Gunnison sage-grouse in Colorado and western sage-grouse in Washington are listed by the U.S. Fish and Wildlife Service as category "C", a candidate taxon ready for listing [117].

OTHER STATUS:
As above, as of 2002, sage-grouse are noted on the Audubon Society's Blue List as a species of special concern in Oregon, Washington, and adjacent Canada [114]. The California Department of Fish and Game also lists sage-grouse as a species of special concern [22] and the Washington Department of Fish and Wildlife [125] has recommended that western sage-grouse be designated a state-threatened species. Nevada Fish and Wildlife [88] has placed sage-grouse on its sensitive animal list. The Colorado Division of Wildlife has recommended that the Gunnison sage-grouse be listed as a species of state special concern [26].

The Utah Department of Natural Resources [118] has assigned sage-grouse a state rank of S2, which is imperiled due to rarity and/or other demonstrable factors. New Mexico [89] classifies sage-grouse as being extirpated in the state.

WILDLIFE DISTRIBUTION AND OCCURRENCE

• GENERAL DISTRIBUTION
• ECOSYSTEMS
• STATES
• BLM PHYSIOGRAPHIC REGIONS
• KUCHLER PLANT ASSOCIATIONS
• SAF COVER TYPES
• SRM (RANGELAND) COVER TYPES
• PLANT COMMUNITIES

Greater sage-grouse are distributed from north-central Oregon, southern Idaho, and southern Alberta and Saskatchewan south to eastern California and extreme western North and South Dakota. Isolated populations also occur in eastern Washington [65,86,121]. Western sage-grouse occur only in eastern Washington and Oregon. The ranges of western and eastern sage-grouse overlap in Oregon. Eastern sage-grouse occur in all states and provinces within the range of sage-grouse except Washington [1,24].

Gunnison sage-grouse occur in 7 counties in southwestern Colorado and 1 county in southeastern Utah [26].

ECOSYSTEMS [51]:
FRES29 Sagebrush
FRES30 Desert shrub
FRES35 Pinyon-juniper

STATES:

Sage-grouse occur throughout the range of big sagebrush (A. tridentata), except on the periphery of big sagebrush distribution or in areas where it has been eliminated [24]. Sage-grouse prefer mountain big sagebrush (A. t. ssp. vaseyana) and Wyoming big sagebrush (A. t. spp. wyomingensis) communities to basin big sagebrush (A. t. spp. tridentata) communities.

Sagebrush cover types other than big sagebrush can fulfill sage-grouse habitat requirements; in fact, sage-grouse may prefer other sagebrush cover types to big sagebrush. Sage-grouse in Antelope Valley, California, for example, use black sagebrush (A. nova) cover types more often than the more common big sagebrush cover types [107]. Drut and others [39] found hens with broods on the National Antelope Refuge in Oregon were most frequently found (54-67% of observations) in low sagebrush (A. arbuscular) cover. Desert shrub habitat may also be utilized by sage-grouse [120].

Sagebrush communities not included in SRM Cover Types but supporting sage-grouse include silver sagebrush and fringed sagebrush (A. frigida) [97], [122]. Sage-grouse use of less common sagebrush communities (e.g. Bigelow sagebrush (A. bigelovii)) may occur but is not documented in the current literature.

BIOLOGICAL DATA AND HABITAT REQUIREMENTS

• TIMING OF MAJOR LIFE HISTORY EVENTS
• PREFERRED HABITAT
• COVER REQUIREMENTS
• FOOD HABITS
• PREDATORS
• MANAGEMENT CONSIDERATIONS

Brooding: Chicks fly by 2 weeks of age, although their movements are limited until they are 2 to 3 weeks old [121]. They can sustain flight by 5 to 6 weeks of age. Juveniles are relatively independent by the time they have completed their first molt at 10 to 12 weeks of age [65].

Seasonal movements: Fall movements to wintering areas are driven by weather conditions and usually occur gradually. After late winter or spring lekking activity, sage-grouse may move to higher elevations or down to irrigated valleys for nesting and feeding. Brooding ranges may be a considerable distance from winter ranges or spring nesting grounds. Schlatterer [106] reported that in southern Idaho, brooding grounds were 13 to 27 miles (21-43 km) from the nesting grounds. Males may also move long distances over the seasons. During winter in Wyoming, Patterson [91] recovered a male greater sage-grouse 75 air miles (120 km) from where he had banded it the previous summer.

PREFERRED HABITAT:
Sage-grouse are totally dependent on sagebrush-dominated habitats [11]. Sagebrush is a crucial component of their diet year-round, and sage-grouse select sagebrush almost exclusively for cover [91]. Because sage-grouse habitat and cover requirements are inseparably tied to sagebrush, they will be discussed together.

Breeding: Open areas such as swales, irrigated fields, meadows, burns, roadsides, and areas with low, sparse sagebrush cover are used as leks [71]. Of 45 leks, Patterson [91] reported that 11 were on windswept ridges or exposed knolls, 10 were in flat sagebrush, 7 were in bare openings, and the remaining 17 were on various other site types. Leks are usually surrounded by areas with 20 to 50% sagebrush cover, with sagebrush no more than 1 foot (30.5 cm) tall.

When not on the lek, sage-grouse disperse to the surrounding areas [121]. Wallestad and Schladweiler [124] studied habitat selection of male greater sage-grouse in central Montana during breeding season and recorded sagebrush height and canopy cover at 110 daytime feeding and loafing sites of cocks. Eighty percent of the locations occurred in sagebrush with a canopy cover of 20-50%. In another Montana study [45], sagebrush cover averaged 30% on a cock-use area, and no cocks were observed in areas of less than 10% canopy cover.

Some females probably travel between leks. In Mono County, California, the home range of marked female greater sage-grouse during 1 month of the breeding season was 750 to 875 acres (300-350 ha), enough area to include several active leks [17].

Nesting: Within a week to 10 days following breeding, the hen builds a nest in the vicinity of the lek [4]. Hens usually nest near the lekking grounds [106], but some hens have been noted to fly as far as 12 to 20 miles (19-32 km) to favorable nesting sites [55,104]. Distances greater sage-grouse hens traveled from the lek to nest in central Montana in a study by Wallestad and Pyrah [123] were:

In Idaho, Autenrieth [4] found the proximity of nests to leks to be:

Quality of nesting habitat surrounding the lek is the single most important factor in population success [5]. Adequacy of cover is critical for nesting. There can be too little: where 13% was the average percent total crown cover on Idaho range, nests were located where average cover was 17%. No greater sage-grouse hens nested in the most arid, open areas with less than 10% total shrub cover. There can be too much: average shrub cover at 87 nest sites was 18.4%, and in more dense cover, sage-grouse did not nest where total shrub cover was greater than 25% [70]. In Utah no nests occurred where threetip sagebrush cover exceeded 35% [97].

Sagebrush forms the nesting cover for most sage-grouse nests throughout the West with concealment being the basic requirement [23]. Rabbitbrush (Chrysothamnus spp.) is occasionally used for nesting cover with greasewood (Sarcobatus vermiculatus) and shadscale (Atriplex canescens) being rarely used [91].

Sage-grouse prefer relatively tall sagebrush with an open canopy for nesting. In Utah, 33% of 161 nests were under silver sagebrush that was 14 to 25 inches (36-63.5 cm) tall, while big sagebrush of the same height accounted for 24% of nests [97]. In a threetip sagebrush (A. tripartata) habitat averaging 8 inches (20 cm) in height, hens selected the tallest plants for nesting cover. Similarly, Patterson [91] reported that in Wyoming, 92% of greater sage-grouse nests in Wyoming big sagebrush were in areas where vegetation was 10 to 20 inches (25-51 cm) tall and cover did not exceed 50%.

Klott and others [76] measured variables at greater sage-grouse nests by habitat type in Idaho. Shrub cover and height were greater at nest sites than random sites in Wyoming big sagebrush and low sagebrush:

In Montana, Wallestad and Pyrah [123] compared sagebrush characteristics around 31 successful and 10 unsuccessful nests. Successful nests had greater than average sagebrush cover surrounding the nest and were located in stands with a higher average canopy cover (27%) than unsuccessful nests (20%). Difference was significant at the 0.005 level. They also found the average height of sagebrush cover over all nests was 15.9 inches (40.4 cm) as compared to an average height of 9.2 inches (23.4) cm in adjacent areas (significant at 0.005 level).

During the nesting season, cocks and hens without nests use "relatively open" areas for feeding, and roost in "dense" patches of sagebrush [70,71].

Brood rearing: Sagebrush is an essential part of sage-grouse brood habitat. An interspersion of sagebrush densities, from scattered to dense, are utilized by broods throughout the summer. Broods can be grouped into 2 categories: those that remain in sagebrush types through the summer and those that shift from sagebrush types in mid-summer and later return to sagebrush.

Throughout the summers of 1968-1969 in a study in Montana, areas that received the greatest amount of utilization by greater sage-grouse broods were areas of sagebrush density characterized as scattered (1-10%) and common (10-25%). Scattered sagebrush received heaviest utilization in June. "Common" sagebrush was utilized heavily throughout the summer. "Dense" sagebrush had greatest use during late August and early September; "rare" sagebrush cover received greatest use in July and August.

Combined data for both years of the study at brood sites showed an average sagebrush cover of 14% during June, 12% during July, 10% during August and 21% during September, which reflects the vegetational types utilized by broods during the summer. Height of sagebrush at brood sites ranged mainly between 6 to 18 inches (15.2-45.7 cm) [120]. In 158 Montana locations, young greater sage-grouse broods used areas of low plant height 9 to 15 inches (23-38 cm)) and density, while older broods and adults used areas where plants were taller (7 to 25 inches (18-63.5 cm) [81].

Early in summer the size of the area used by greater sage-grouse hens with broods in Idaho seemed to depend upon the interspersion of sagebrush types that provided an adequate amount of food and cover. Areas with sagebrush in scattered densities, with occasional clumps in the common to dense categories, appeared to be preferred. In their daily activity, broods tended to use more open sites for feeding and to seek more dense clumps of sagebrush for roosting.

Throughout early summer, daily movements of broods were relatively long, reflecting the great daily activity required of broods to meet their nutritional needs. Klott and others [76] found that greater sage-grouse hens with broods had the following home ranges in Idaho:

Cover types used by hens with broods typically had greater availability of forbs during periods of high use, but differences in availability between areas influenced use of cover types, movements, and diets.

In Oregon, the greater sage-grouse hens at Jackass Creek selectively used sites with forb cover greater than typically found there and similar to that generally available to broods at Hart Mountain National Antelope Refuge. This amount of forb cover (12-14%) may represent the minimum needed for greater sage-grouse brood habitat in Oregon [39].

Succulence of their favored foods appears to be a key to sage-grouse movements [70]. As plants mature and dry, the grouse move to areas still supporting succulent vegetation. A delay in maturing of forbs has a noticeable effect on bird movements [71].

Broodless: A study by Gregg and others [56] in Oregon revealed differences in chronology of summer movements and cover types used between broodless greater sage-grouse hens and greater sage-grouse hens with broods. Broodless hens gathered in flocks and remained separate from but in the vicinity of hens with broods during early summer. However, broodless hens moved to meadows earlier in summer and used a greater diversity of cover types than hens with broods perhaps because dietary needs of broodless hens might be less specific than those of hens with broods.

Winter: A winter-use area appears to be both a key habitat segment and a major factor in sage-grouse distribution over a large area [45]. The best winter habitat is below snowline, where sagebrush is available all winter [104]. Dalke and others [34] reported wintering grounds of greater sage-grouse in Idaho were usually where snow accumulation was less than 6 inches (15 cm). In areas of deep snow, greater sage-grouse winter where sagebrush has grown above the snow level [6].

Sage-grouse appear to select areas of little or no slope. In a Colorado study, nearly 80% of Gunnison sage-grouse winter use of 500 square miles (1,252 km²) of sagebrush was on less than 35 square miles (87 km²): on flat areas where sagebrush projected above the snow, or on south- or west-facing sites of less than 5% slope, where sagebrush was sometimes quite short but still accessible [65]. In Montana, prime wintering areas were flat, large expanses of dense sagebrush; winter home ranges of 5 greater sage-grouse females in Montana varied from 2,615 to 7,760 acres (1,050-3,100 ha) during 2 different years [45].

Winter-use areas are determined by amount of snow rather than affinity to a particular site [102]. Majority of winter observations are in sagebrush with more than 20% canopy coverage. Species and subspecies of sagebrush that seem to be preferred by grouse in the winter are black sagebrush, low sagebrush, and some subspecies of big sagebrush [4,33].

COVER REQUIREMENTS:
See Preferred Habitat

FOOD HABITS:
Adults: The importance of sagebrush in the diet of adult sage-grouse is impossible to overestimate. Numerous studies have documented its year-round use by sage-grouse [9,23,24,71,91,107,109,121,122]. A Montana study, based on 299 crop samples, showed that 62% of total food volume of the year was sagebrush. Between December and February it was the only food item found in all crops. Only between June and September did sagebrush constitute less than 60% of the greater sage-grouse diet [121]. Sage-grouse select sagebrush species differentially. Greater sage-grouse in Antelope Valley, California, browsed black sagebrush more frequently than the more common big sagebrush [107]. Young and Palmquist [131] state the browse of black sagebrush is highly preferred by greater sage-grouse in Nevada. In southeastern Idaho, black sagebrush was preferred as forage [33].

Among the big sagebrush subspecies, basin big sagebrush is less nutritious and higher in terpenes than either mountain or Wyoming big sagebrush. Sage-grouse prefer the other two subspecies to basin big sagebrush [6]. In a common garden study done in Utah, Welch, Wagstaff and Robertson [126] found sage-grouse preferred mountain big sagebrush over Wyoming and basin big sagebrush. However, when leaves and buds of the preferred species became limited, the birds shifted to the lesser-liked plants. The authors concluded the birds, while expressing preference, are capable of shifting their eating habits.

Sage-grouse lack a muscular gizzard and cannot grind and digest seeds; they must consume soft-tissue foods [121]. Apart from sagebrush, the adult sage-grouse diet consists largely of herbaceous leaves, which are utilized primarily in late spring and summer [42]. Additionally, sage-grouse use perennial bunchgrasses for food [8].

Sage-grouse are highly selective grazers, choosing only a few plant genera. Dandelion (Taraxacum spp.), legumes (Fabaceae), yarrow (Achillea spp.) and wild lettuce (Lactuca spp.) account for most of their forb intake [6,109]. Martin [81] found that from July to September, dandelion comprised 45% of greater sage-grouse intake; sagebrush comprised 34%. Collectively, dandelion, sagebrush, and two legume genera (Trifolium and Astragalus) contributed more than 90% of the sage-grouse diet. Insects are a minor diet item for adult sage-grouse. Martin and others [80] reported insects comprised 2% of the adult greater sage-grouse diet in spring and fall and 9% in summer. Sagebrush made up 71% of the year-round diet.

Prelaying females: Herbaceous dicots are used heavily by females before egg laying and may be essential for sage-grouse nutrition because of their high protein and nutrient content [8].

Favored foods of prelaying and brood-rearing greater sage-grouse hens in Oregon are [130]:

common dandelion (Taraxacum officinale)
goatsbeard (Tragopogon dubuis)
western yarrow (Achillea millifolium)
prickly lettuce (Lactuca serriola)
sego lily (Calcochortus macrocarpus)

In southeastern Oregon, Barnett and Crawford [8] studied prelaying nutrition of greater sage-grouse hens, March to April, 1990 and 1991.

Juveniles: In their 1st week of life, sage-grouse chicks consume primarily insects, especially ants and beetles [91]. Their diet then switches to forbs, with sagebrush gradually assuming primary importance. In a Utah study, forbs composed 54 to 60% of the summer diet of juvenile sage-grouse, while the diet of adult birds was 39 to 47% forbs [116].

In a Wyoming study, Johnson and Boyce [66] evaluated effects of eliminating insects from the diet of newly-hatched greater sage-grouse chicks. All chicks hatched in captivity and not provided insects died between the ages of 4 and 10 days, whereas all chicks fed insects survived the 1st 10 days. Captive greater sage-grouse chicks required insects for survival until they were at least 3 weeks old. Greater sage-grouse chicks > 3 weeks old survived without insects, but their growth rates were lowered significantly, indicating insects were still required for normal growth after 3 weeks of age. As quantity of insects in the diet increased, survival and growth rates also increased up to 45 days, the length of the experiment.

In a study conducted in Idaho, Klebenow and Gray [74] measured food items for juvenile greater sage-grouse for each age class, classes being defined by weeks since birth. In the 1st week insects were very important - 52% of the total diet. Beetles, primarily family Scarabaeidae, were the main food item. Beetles were taken by all other age classes of chicks, but in smaller amounts. All ages fed upon ants and while the volume was generally low, ants were found in most of the crops. After week 3, insect volume dropped and stayed at a lower level throughout all the age classes, fluctuating but always under 25%.

Forbs were the major plant food of the chicks. Harkness gilia (Linanthus harknessii) was the main forb species in the 1st week and then steadily decreased. It was not found in the diet after 6 weeks. Loco (Arabis convallarius) and common dandelion were important food items for most of the collection period and occurred with generally high frequencies. Common dandelion was the most abundant food item and the mainstay of the sage-grouse chicks. At 6 weeks of age, goatsbeard reached its peak in the diet and sego lily was found in greatest volume a week later. These 5 species were the most important forbs.

The only shrub of importance was big sagebrush. It appeared in the diet at 4 weeks of age and as the ages progressed, the volume increased steadily. These 6 plants comprised 83% of the total sample and are listed in order of decreasing percent total volume [74]:

With plants like common dandelion and goatsbeard, all aboveground parts of the plant were sometimes eaten. The stems, however, were not of main importance. The reproductive parts, mainly buds, flowers, and capsules, were the only parts taken from some of the other species. Conversely, leaves were the only parts of sagebrush found in the crops. Leaves and flowers of the species listed above and other dicots contained higher amounts of crude protein, calcium, and phosphorus than sagebrush and may be important in greater sage-grouse diets for these reasons [8].

In a central Montana study, Peterson [94] analyzed crop contents of juvenile greater sage-grouse. Data indicate increasing use of sage species and decreasing use of insects. Percent frequency and percent volume of food items commonly utilized by 1- to 12-week-old greater sage-grouse collected during 1966 and 1968 were:

Five most preferred¹ plant species in each 2-week age division of juvenile sage-grouse, 1966 and 1968 are listed below. Note only 1 sagebrush species is listed and that it is not preferred until weeks 7-8 [94]:

Water: Sage-grouse apparently do not require open water for day-to-day survival if succulent vegetation is available. They utilize free water if it is available, however. Sage-grouse distribution is apparently seasonally limited by water in some areas. In summer, sage-grouse in desert regions occur only near streams, springs, and water holes. In winter in Eden Valley, Wyoming, sage-grouse have been observed regularly visiting partially frozen streams to drink from holes in the ice [23].

PREDATORS:
Predators are commonly believed to negatively impact sage-grouse populations and of most importance is timing of death. Nest loss to predators is most important when potential production of young and recruitment are seriously impacted [19]. Lack of adequate nesting and brooding cover may account for high juvenile losses in many regions [68].

Studies by Gregg and others [57], Delong and others [37] in Oregon, and Sveum and others [112] in Washington suggest that nest success is related to herbaceous cover near the nest site. Taller, more dense herbaceous cover apparently reduces nest predation and likely increases early brood survival [19]. Although predators were the proximate factor influencing nest loss, the ultimate cause may relate to the vegetation available to nesting sage-grouse [57]. Tall dense vegetation may provide visual, scent, and physical barriers between predators and nests of ground-nesting birds. Greater amounts of both tall grass and medium-height shrub cover were associated collectively with a lower probability of nest predation [37]. In a series of Nevada studies, artificial nest predation experiments were conducted. Artificial nests experienced 100% mortality with the loss of 1,400 eggs in 200 simulated sage-grouse nests in 2 weeks in 1 study, 84% of the nests were destroyed in 3 days in another study, while just 3% of the nests were destroyed in 10 days in an area of significantly better cover (t test, P < 0.05) [69].

Generally, quantity and quality of habitats used by sage-grouse control the degree of predation. Thus, predation would be expected to be most important as habitat size and herbaceous cover within sagebrush decreases [19].

A decline in preferred prey may also result in increased predation on sage-grouse. Kindschy [68] suggested that in southeastern Oregon, a decline in black-tailed jackrabbit (Lepus californicus) numbers may have caused predators to switch to sage-grouse as their primary prey.

Predator species: Coyote (Canis latrans) [68,115], bobcat (Lynx rufus) [7], badger (Taxidea taxus) [76], falcons (Falconidae) [93], and hawks, kites, and eagles (Accipitridae) [9,40,95] prey on adult and juvenile sage-grouse. Crows and ravens (Corvus spp.) and magpies (Pica spp.) consume juvenile birds [68,115]. Coyote, ground squirrels (Spermophilus spp.), and badger are the most important mammalian nest predators. Among bird species, magpies and ravens commonly prey on sage-grouse nests [62,65,121].

Sage-grouse are a popular game bird. Mortality due to hunting is generally considered to be compensatory [19,34] and replacive [19], where until mortality reaches a "threshold value" it has no effect on population levels. Autenrieth and others [6] state data are not available to suggest that closed or restricted hunting seasons will materially affect overall sage-grouse population levels on their primary range.

In a study on hunting in a low density greater sage-grouse population in Nevada, Stigar [110] concluded low sage-grouse populations may be a result of factors other than hunting. Protecting 1 sage-grouse population from hunting while doubling the birds harvested in a 4-year period on another population showed that despite low recruitment, both populations increased to nearly the same density.

In an Oregon study, no relationship existed between the rate of summer recruitment (chicks/adult) and harvest by hunters. Nor was any significant relationship found between the size of the fall harvest and population trends during the subsequent spring. Decline in greater sage-grouse populations was apparently unrelated to harvest because no relationship was found between the magnitude of the harvest and the size of subsequent spring populations [31].

Braun and Beck [20] demonstrated no relationship between harvest rates and subsequent breeding population size in Colorado. Gill [54], also in a Colorado study, concluded that hunter harvest had little effect on spring populations of male Gunnison sage-grouse.

MANAGEMENT CONSIDERATIONS:
Future of the sage-grouse depends largely on people's ability and willingness to maintain habitat vital to its needs. Sage-grouse are habitat-specific to 1 particular plant type in meeting their life requirements. Destruction of habitat has been the basic cause of sage-grouse decrease throughout the West [23].

Sage-grouse once occurred virtually everywhere there was sagebrush. Sage-grouse have declined primarily because of loss of habitat due to overgrazing, elimination ofsagebrush, and land development [59]. Sage-grouse populations began declining from 1900 to 1915, when livestock utilization of sagebrush rangeland was heavy [91]. In the 1950's and 1960's, land agencies adopted a policy of aggressive sagebrush control to convert sagebrush types to grassland. Chaining, frequent fire, and herbicide treatments reduced sagebrush by several million acres, and sage-grouse numbers plummeted drastically [23,84]. Conversion of sagebrush types to grassland has since been brought into question as a management practice for both wildlife and livestock [64,65,107,121].

Daubenmire [35] questions removal of sagebrush as a management tactic and lists 6 concerns:

1. There is little evidence to indicate the extent to which the desired grass increase (measured shortly after shrub eradication) is maintained.

2. Protection afforded many grass plants by dense clumps of shrubs is the sole reason why any perennial grass remains on much of the depleted ranges. Sagebrush elimination opens the way to complete destruction of perennial grass by overuse.

3. Studies in Washington [36] have shown that for more than 4 months in the summer, big sagebrush uses only water that has percolated through the soil profile below the reach of grass roots. Removal of sagebrush allows some minerals to migrate permanently below the reach of grass roots.

4. Removing sagebrush eliminates certain elements of avifauna. While rearing their young, these birds drain insect populations. Increased insects may damage the residual grass.

5. When sagebrush is removed by herbicidal sprays, certain perennial broadleaved forbs are heavily damaged or eliminated.

6. In some areas sagebrush promotes the uniform accumulation of snow and delays its melting.

Call [23] states:

"Any land use practice which has as its objective the permanent elimination of sagebrush and establishment of grasses in the Mountain West will ultimately reduce the collective carrying capacity of that range for livestock, elk, mule deer, antelope, sage-grouse, and many smaller species of wildlife."

For detailed information on individual species of sagebrush and effects of various control measures on those species, see the summary for each species in FEIS.

FIRE EFFECTS AND USE

• DIRECT FIRE EFFECTS ON ANIMALS
• HABITAT-RELATED FIRE EFFECTS
• FIRE USE

Breeding: Leks, where breeding occurs, are usually small open areas from 0.1 to 10.0 acres (0.4-4.0 ha) in size, but may be as large as 100 acres (40.5 ha) [23]. Effects of fire on the lek itself are not as much a concern as they are on the area around the lek [5]. This area is used by sage-grouse for loafing, feeding, and escape cover [4,72]. A study by Wallestad and Schladweiler [124] in Montana recorded sagebrush cover and height at 110 daytime sites of cock greater sage-grouse and found 80% of the locations occurred in sagebrush with a canopy cover of 20-50%. Sagebrush canopy cover for the 110 sites averaged 36%. Also in Montana, Eng and Schladweiler [45] found that a cock-use area during the breeding season had canopy coverage of sagebrush averaging 30%. In Idaho, Autenrieth [4] found 80% of cock greater sage-grouse locations occurred in sagebrush stands of 20-50% canopy coverage.

An Idaho study by Martin [83], after wildfires in 1981, found greater sage-grouse continued to use leks located within extensive burns. Attendance of males at burned leks did not decrease (after adjusting for population declines) despite extensive reduction of surrounding shrub cover. During this study, males and females attended leks in spite of the loss of vegetative cover immediately surrounding leks. Greater sage-grouse seemed to adapt to loss of sagebrush in travel corridors and in principal breeding season habitat of cocks. Cocks and hens appeared to feed, loaf, and roost primarily in unburned habitat and fly into burned leks. Martin found greater sage-grouse attended burned and unburned leks within perimeters of large burns and suggested there are 2 factors that account for why greater sage-grouse attended burned leks in his study area: 1) The summer wildfire burn occurred in a mosaic pattern and left many unburned islands, so abundant nesting habitat was available near all leks within the burn. 2) The long-term population within the burn area was well below average population when the burn occurred, allowing the smaller unburned portions of the study area to supply necessary habitat requirements. Gates [52] observed greater sage-grouse displaying and mating on a burned area in Idaho during the 1st 2 breeding seasons following a burn. These observations also took place during a population decline.

Fischer [47] studied lek attendance patterns after a late-summer prescribed burn on the upper Snake River Plain of southwestern Idaho. He could detect no differences attributable to fire after 2 years. However, an additional year of data on lek attendance and further analysis of lek data from Fischer's study provide evidence that fire had a negative influence on the breeding population in the treatment area [28]. The notion that fire had negative effects on the greater sage-grouse breeding population in the treatment area is supported by 4 findings: 1) The treatment area had a higher loss of leks (-58%) than did the control area (-35%). 2) Changes in attendance in major leks by males were similar in treatment and control areas during the preburn period but the treatment area had a greater decline in attendance (-90%) than the control area (-63%) during the postburn period. 3) Average lek attendance at the 2 largest leks in both areas was greater in the treatment (67 males) than the control (59 males) area during the preburn period. However, the situation reversed during the postburn period, and average attendance at the 2 treatment leks (22 males) was less than the average attendance at the control leks (36 males). 4) The mean number of male greater sage-grouse per lek was similar in treatment and control areas during the preburn period but was much lower in the treatment area (6 males) than in the control area (17 males) during the postburn period.

In a study in Idaho, Connelly and others [30] found some greater sage-grouse leks persist even after sagebrush areas have been burned. Data from the study suggest that greater sage-grouse continue to use leks in altered areas because some hens nest successfully under nonsagebrush plants. The authors caution that persistence of leks in burned areas should not be interpreted as evidence that fire has little effect on greater sage-grouse populations. Greater sage-grouse use of nonsagebrush shrubs for nest sites may allow populations to persist at low levels in a burned area until the area recovers. However, this behavior may only slow, but not prevent, the birds' ultimate disappearance from the burned area.

Leks used by migratory populations of sage-grouse appear to be more susceptible to deleterious habitat-related fire effects because they are usually dominated by Wyoming big sagebrush [129] which re-establishes slowly following fire [85]. Degradation of breeding range because of fire, drought, grazing, or herbicides will likely reduce sage-grouse populations because of low nesting effort, low nest success, or poor chick survival. Regardless of the method used to eliminate or reduce sagebrush cover in xeric sage-grouse breeding habitat, these actions have the potential for reducing breeding populations of grouse. It is recommended that prescribed burning be avoided in relatively xeric habitats used by breeding migratory sage-grouse. If a Wyoming big sagebrush habitat type supports a breeding population of sage-grouse, it is recommended a high priority be given to suppressing wildfires in these habitats during drought [28].

Nesting: After mating, hens disperse to nest [72]. Most nest near leks, and areas within a 1.9 mile (3 km) radius of leks is considered the most important for nesting [21]. Some hens may fly as far as 32 to 48 miles (20-30 km) to nest [104]. Nesting habitat is often a belt between summer and winter range [70]. Often there is a complex of habitat types centered around a lek. In nonmigratory populations of sage-grouse, this complex may provide for year-round needs of the birds. In migratory populations, this suite of habitat types often includes breeding, nesting, early brood-rearing, and winter habitats, and sage-grouse migrate to fill their summer needs [50].

Gill [54] found that Gunnison sage-grouse nested under all types of shrub cover on his study area in Colorado, but preferred sagebrush, with 92.3% of 117 nests located under sagebrush. Shrub heights preferred for nesting varied from 11.8 to 23.6 inches (30-60 cm), with 7.9 to 31.5 inches (20-80 cm) adequate and shrub canopy coverage > 17% [24]. In Idaho, Klebenow [70] found no greater sage-grouse nested in the most arid, open areas (< 10% total shrub cover, mean shrub canopy coverage at 87 nest sites = 18.4%). In more dense cover, greater sage-grouse did not nest where total shrub cover was > 25% [71].

Complete removal of sagebrush in burned areas could reduce nesting, hiding, and wintering cover for sage-grouse [21]. Sage-grouse show fidelity to leks and/or nesting areas [13,48]. A study on the Big Desert, in Idaho, by Fischer and others [48], found that because greater sage-grouse hens appear to seek suitable habitat within a relatively small area, nest area fidelity may reduce nesting if large areas of nesting habitat are destroyed. Hens may nest in unsuitable areas and experience lower nest success [30].

Klebenow [70] states the cover shrubs provide is necessary for nesting greater sage-grouse, and complete removal of shrubs can only result in elimination of greater sage-grouse from the area. Fire eliminates potential winter and nesting habitat according to Robertson [103] and Fischer [47]. Connelly and Braun [27] feel fire may negatively impact sage-grouse populations by eliminating or fragmenting relatively large blocks of wintering or nesting habitat. During the 1st few years after burning, nesting habitat is essentially destroyed [87].

Fischer [47] studied the effects of fire on migratory greater sage-grouse breeding, nesting, and brood-rearing habitat in southeastern Idaho. Nests inside the burned area were only found in unburned patches, suggesting that removal of sagebrush by fire reduces nesting cover for grouse. Although no immediate differences in use of burned and unburned habitat were demonstrated in this study for 3 postfire years, long-term response of nesting greater sage-grouse to fire may be dependent on both the scale of sagebrush removal and intensity of fidelity to nesting areas. Greater sage-grouse that show nesting-area fidelity may subsequently return to the same area and attempt a nest even after the habitat has been manipulated. Attenuation of nest-area fidelity as site-tenacious individuals die may decrease production in the burned area. Fire may also reduce the number of suitable nesting sites by removing shrub cover for nests.

Brood rearing: The single unifying theme threatening sage-grouse across their geographical range is the universality of brood habitat loss [38]. Drought and increased fire frequency may be the primary agents causing a decline in brood-rearing habitat for sage-grouse. Moreover, an unfavorable situation due to drought and an increase in wildfire may have been made worse in many areas by vigorous prescribed burning [27]. Brood habitat typically has 15-25% shrub canopy closure but at least 10-20% cover of live forbs and grasses [38]. In southern Idaho, the percent canopy cover of big sagebrush at brood sites was 8.5%, significantly less than the average for the entire area, 14.3% [70].

Broods are tied to food in addition to cover [91]. After hatching, before chicks can fly and when mortality is highest [4,91], broods need food in close proximity to escape cover [111]. Diet of sage-grouse chicks is chiefly insects early in life, shifting to succulent forbs and shrub foliage as chicks grow older [74,91,94]. Abundant food forbs in close proximity to unburned sagebrush cover could benefit sage-grouse broods by providing additional food with adequate cover, and fire-enhanced flowering may improve forage availability for sage-grouse during the brood rearing period [130]. Sime [109] states openings in sagebrush canopy may increase forbs and improve brood habitat, and Klebenow [71] suggests reduced shrub cover on burned sites may increase accessibility of forbs and insects to chicks. In a 2-year study of greater sage-grouse responses to wildfire in Idaho, Martin [83] found that burning improved habitat for sage-grouse broods. During both postburn years, forb crown cover was significantly higher in burned habitats than in unburned habitats. However, Nelle and others [87] found that during the 1st few years after burning on a site in southeastern Idaho, greater sage-grouse brood-rearing habitat was not improved.

After a prescribed burn on the Snake River plain of southeastern Idaho, Connelly and others [28] found no difference in the use of burned and unburned areas by a migratory population of greater sage-grouse. These results suggest fire does not improve brood-rearing habitat in relatively low precipitation areas dominated by Wyoming big sagebrush. The authors were unable to show an increase in forbs following the fire, so fire may have caused an overall decline in brood-rearing habitat, perhaps contributing to the decline in greater sage-grouse population following the fire. Decreased food abundance following fire in the treatment area may have indirectly affected survival by increasing chick movement. Fire in Wyoming big sagebrush has not been reported to increase the length of the growing season for forbs important in the diet of sage-grouse [130].

Fischer's [47] research in southeastern Idaho provided evidence for excluding fires that eliminate large blocks of vegetation in brood habitat for migratory greater sage-grouse within xeric regions, because of its impact on insects. Fire appeared to negatively impact insect abundance in 1 of the 3 orders that are most important in sage-grouse diets. The fire created a mosaic of sagebrush areas interspersed with open areas having abundant grasses and forbs, but there was no positive response of greater sage-grouse to the burned area. Broods require food, mainly forbs, in addition to cover. Both may be present in a given acre of land, but the birds also use and appear to seek areas where there is an interspersion of habitat types. They only feed in areas where cover is nearby [71].

A study by Fischer and others [49] in southeastern Idaho indicated that short-term effects of a prescribed fire in a xeric environment did not enhance brood-rearing habitat and may have been detrimental to grasshoppers which are important in sage-grouse diets. Gates and Eng [53] examined use of burned areas by migratory greater sage-grouse after prescribed burns in 1981 and 1982 at the Idaho National Engineering Laboratory. They suggested the patchy burn which occurred could enhance early brood-rearing habitat. If availability of forbs and insects were enhanced on burned areas, then an abundant food supply would be available to broods in close proximity to escape cover.

Summer: Sage-grouse have been reported to be attracted to burn areas during summer [73,83]. Summer habitat is characterized by shrub canopy cover of at least 15% and at least 10% live forb cover [38]. Sagebrush and forbs are essential components of summer sage-grouse habitat [41]. If fire increases availability of photosynthesizing, succulent foods in uplands during the late summer, available sage-grouse habitat may be increased [130].

Connelly and others [28] studied migratory greater sage-grouse response to a controlled burn on the upper Snake River Plain of southeastern Idaho. They found fire apparently had no influence on the timing of migration of female greater sage-grouse. However, they found the summer population associated with the burn area declined 75% while that in the control area declined 52%. A similar but more pronounced decline occurred at the largest lek in the burned area. Estimated summer population associated with this lek declined 98% during the 9-year study while the population associated with a similar lek in the unburned area only declined 45%.

Migratory greater sage-grouse in a xeric habitat were studied by Fischer [47] in Idaho. He found that cover of forbs important in greater sage-grouse summer diets was similar in burned and unburned habitats. Although the fire created a mosaic of sagebrush areas interspersed with open areas having abundant grasses and forbs, there was no movement of greater sage-grouse to the burned area.

Winter: Many researchers [10,45,91] describe winter habitat as probably the most limiting seasonal habitat and thus perhaps the most critical [99]. The majority of winter observations are in sagebrush with more than 20% canopy coverage [102]. Connelly and others [28] describe sage-grouse wintering habitat as open to moderately dense sagebrush with 10-20% canopy cover. For sedentary populations of sage-grouse, the wintering area is often located within 2 miles (3.2 km) of the strutting ground and nesting area [102].

Fire may negatively impact sage-grouse populations by eliminating or fragmenting relatively large blocks of wintering habitat [27] and at times, severe fires destroy important wintering areas for sage-grouse [24]. Sage-grouse show affinity for particular winter ranges [13,45], and known sage-grouse wintering areas should receive priority attention in the control of wildfires [6]. Lack of protection of critical winter habitat has resulted in sage-grouse population declines [13], and Sime [109] states it is imperative that winter sage-grouse habitat be protected. Braun and others [21] caution that complete removal of sagebrush in burned areas could reduce nesting, hiding, and wintering cover for sage-grouse. Gates and Eng [53] suggest elimination of sagebrush by burning will destroy wintering habitat for grouse.

In areas where winter habitat is limiting, loss of contiguous sagebrush rangeland could negatively impact sage-grouse by removing food and cover. Robertson [103] conducted a study of a migratory greater sage-grouse population in a xeric habitat in Idaho. He found burning decreased the structural components of greater sage-grouse winter habitat, and use of these areas decreased dramatically. This does not imply that that burning is detrimental to greater sage-grouse with large expanses of winter habitat. If burning takes place on critical winter range, and heavy snow fall covers much of the remaining habitat during the following winter, food and cover may be severely limited. Klebenow [71] suggests there is little place for fire on wintering sites of migratory greater sage-grouse in xeric habitat since there is nearly a complete reliance on shrubs for food and cover.

FIRE USE:
Use of prescribed fire as a management tool for sage-grouse habitat is controversial [21,27,34,127]. Specific concerns relate to sagebrush removal as a means to manage for a sagebrush obligate and the unknown long-term sagebrush re-establishment rates [32]. A major danger in using prescribed fire for management of sage-grouse habitat is increasing fire frequency beyond the rate at which the sagebrush component of the ecosystem can recover. If fire return intervals become frequent enough to exceed the capacity of sagebrush to repopulate, sagebrush cover would be reduced far below historical levels which could be very detrimental to sage-grouse [130]. Matisse [84] suggests wildfire and prescribed burns have had a detrimental effect on quality and quantity of sage-grouse habitat.

Miller and Eddleman [85] acknowledge the debate on benefits of fire to enhance sage-grouse habitat. They state 4 factors determine negative or positive outcome of fire on sage-grouse habitat: 1) site potential, 2) site condition, 3) functional plant group(s) that is(are) limiting, and 4) pattern/size of the burn. Fire is a useful tool to enhance native perennial grass and forbs, particularly in areas where sagebrush is abundant, a "good" population of native forbs is present, and exotic species are limited.

This most often applies to mountain big sagebrush communities where shrub cover can exceed 35% and perennial forbs can increase 2-3 fold following fire [96]. There is little evidence that fire will enhance sage-grouse habitat where there is already a balance of native shrubs and perennial grasses and forbs. Fire should not be used where sagebrush cover is the limiting factor for sage-grouse or where the understory lacks perennial forbs and grasses and introduced annuals are present.

Response of native understory species to fire is usually determined by moisture spectrum and condition of the site. On mesic sites, fire is useful for increasing amounts, nutrient quality, and season of succulence of perennial grasses and annual forbs important in sage-grouse diet. However, in the drier sagebrush cover types, perennial forbs are not increased by fire.

Fire can open up dense stands of sagebrush and increase landscape level heterogeneity, or homogenize large landscapes by removing the shrub layer and promoting dominance of introduced annuals. Blaisdell and others [16] state primary use of fire on sagebrush-grass ranges should be to control dense stands of sagebrush so more desirable species can increase. The goal should be roughly consistent with the climax cover that can be attained in a particular habitat type or site. In addition, they state grazing management after burning is essential. Most burns should be completely protected from livestock grazing for at least 1 and possibly 2 growing seasons. Benson and others [11] recommend a nongrazing period of 2-3 years. On areas where cheatgrass (Bromus tectorum) is abundant, special measures may be necessary to prevent recurrent fires [16].

Braun and others [21] provide the following guidelines to be used when altering sagebrush habitat:

1. The state wildlife agency will be notified of each specific proposal to control vegetation a minimum of 2 years in advance of treatment by means of an "Environmental assessment." In situations where it is not possible to provide such notice (i.e. private lands), the state wildlife agency should be notified as soon as the project is proposed. Lead time is necessary to evaluate control projects during all seasons of the year. a.) The public land-management agency will provide the state wildlife agency with maps on which the proposed treatment areas are located along with detailed plans as to the type of treatment and expected results. b.) The state wildlife agency will plot sage-grouse use leks, nesting areas, wintering sites, and meadows and summer range or brooding areas on the maps. c.) Representatives of the cooperating agencies will meet on the proposed project area for an on-the-ground inspection following completion of the maps. d.) No sagebrush will be treated or removed until a comprehensive multiple-use management plan has been completed for the area. e.) Project plans for sagebrush control will include provisions for long-term quantitative measurements of vegetation before and after control to acquire data on the effects on wildlife habitats, and to ascertain whether objectives of the project were accomplished. The land-management agency should bear responsibility for evaluation of the project as it relates to habitat, while the state wildlife agency should assume responsibility of measuring the effects of the project on the sage-grouse resource.

2. No sagebrush control work will be considered where live sagebrush cover is less than 20%, or on steep (20% or more gradient) upper slopes with skeletal soils where big sagebrush is 12 inches (30 cm) or less in height.

3. The breeding complex (leks and nesting areas) will be considered as all lands within a 1.9 mile (3 km) radius of an occupied lek (in some situations, depending on the quality of the nesting habitat, this radius may well exceed 1.9 miles (3 km)). Control of vegetation within the breeding complex will not be undertaken within 1.9 miles (3 km) of leks or on nesting and brood areas. On-site investigations by land-management and state wildlife agency personnel will be essential to determine inviolate areas. Areas to be protected from treatment will be marked on maps.

4. There will be no sagebrush control attempted in any area known to have supported important wintering concentrations of sage-grouse within the preceeding 10 years.

5. No control will be attempted along streams, meadows, or secondary drainages (dry or intermittent). A 109 yard (100 m) strip (minimum) of living sagebrush will be retained on each edge of meadows and drainages. On-site inspections by land-management and wildlife agency personnel will be made to assess the desirability of increasing or decreasing the width of untreated strips in specific areas.

6. When sagebrush control is found to be unavoidable in sage-grouse range, all treatment measures should be applied in irregular patterns using topography and other ecological considerations to minimize adverse effects to sage-grouse. Widths of untreated areas can vary for the convenience of application technique, except treated areas will not be wider than 33 yards (30 m) and untreated areas will be at least as wide as treated areas. Untreated areas will not be treated until food and cover plants in the treated areas attain comparable composition to the treated areas.

7. Where possible, spraying will be done with a helicopter or ground equipment. No spraying will be done when wind velocity exceeds 6.2 miles per hour (10 km/hr.)

8. Whenever possible, complete kill or removal of sagebrush in treated areas should be avoided. Partial kill or removal of sagebrush may enhance the area for livestock, prevent loss of all snow cover in winter, and allow for some use of the disturbed area by sage-grouse.

Prescribed fire has been used by the U.S. Forest Service on sagebrush lands in the 12-inch (300 mm) or greater precipitation zones. Results have usually been positive in terms of forb response. In the 8- to 12-inch (200-300 mm) precipitation zones, best results seem to occur with April burns when fine fuels from the previous growing season carry a relatively "cool" fire. Small burns in irregular patterns or strips not exceeding 50 yards (45 m) in width and 100 yards (90 m) long are best. Where wildfire is a concern during summer months, burned strips perpendicular to the prevailing summer winds serve as fire breaks [6].

After a study of sage-grouse use of burned, unburned, and seeded sites, Sime [109] summarized several factors researchers feel should be considered before using fire to manage for sage-grouse: 1) Radio telemetry rather than observation should be used to insure proper identification of seasonal ranges of specific population segments as some segments may be sedentary while others are migratory [13]. 2) Openings in the overhead canopy should be restricted to 1 to 10 acres (0.4-4.0 ha) [24]. 3) Fire should only be used when desirable vegetation is dormant so as not to harm the plant species grouse rely on for food and/or cover [128]. 4) The burn site should not be grazed until a vigorous stand of self-perpetuating vegetation is well established [24]. Fire would be most effective in areas where annual precipitation enables rapid revegetation and in areas with a limited potential for invasion of the site by weedy, undesirable species. When few propagules from the native vegetation community are available or when fire is used in areas of low precipitation, the disturbed site may require artificial seeding. Seeding crested wheatgrass (Agropyron cristatum) in sage-grouse range is not recommended.

Call and Maser [24] suggest the following list to consider when using fire to manage sage-grouse habitats:

1) Fire is an inexpensive tool that can be used for habitat manipulation, but all projects must be carefully evaluated, planned, and supervised. A set of clear objectives is essential.

2) Fire is best used in a manner that results in a mosaic pattern of shrubs and open areas, where the resultant openings range in size from 1 to 10 acres (0.4-4.0 ha).

3) Large, severe fires may remove an excessive amount of cover or may sterilize the soil.

4) Burning within an area should be done on a rotational basis, different patches burned every few years, with as long as 20 years between burning treatments on each site. This will produce a variety of habitats within the general area. Timing of treatments over the years will depend on sagebrush response and growth rates at specific sites.

5) Best results from burning occur in late April and early May when dry grasses and other herbaceous fuels from the previous growing season will carry a relatively "cool" fire. This will leave some sagebrush and still create openings for additional growth of grasses. Fires in late spring and early summer, however, could destroy many nesting birds and other young wildlife, including sage-grouse, so use of fire is preferable when young are capable of escaping. At times, it may be necessary to use prescribed fire in the spring to obtain the desired changes in habitat. Strip burns that do not exceed 50 yards (45 m) in width and 100 yards (90 m) in length create desirable openings for sage-grouse [4].

6) Sagebrush habitats identified as important wintering areas that are still in vigorous condition should remain intact. Sage-grouse depend on sagebrush leaves in such areas, not on potential development of grass and forbs in the understory or interspersed openings. If important stands of sagebrush used by sage-grouse for wintering have deteriorated because of such things as insects, old age, or livestock grazing, the manager should initiate measures to rejuvenate the stand by light chaining, by reduced grazing, or other means.

7) Four primary elements needed for a successful burn in areas with more than 11.8 inches (300 mm) of precipitation are (1) wet soil; (2) windspeed in excess of 8 mph (12.8 km/hr) and gusty; (3) fine fuels of 612-704 pounds/acre (278-320 kg/ha); and 4) no burning after spring grass growth reaches 2 inches (5 cm) unless burning is to improve the forbs in the community. Prescribed burns should be conducted when plants preferred as food by sage-grouse are dormant [128].

8) Livestock concentrate on burned areas and eat the new growth, so they need to be carefully managed. Haphazard burning and heavy grazing accelerate sagebrush reinvasion, soil erosion, and loss of forage plants desirable for both grouse and livestock. Grazing use must therefore be regulated to prevent excessive reinvasion by sagebrush (more than 10,000/acre (24,700/ha)) and to prevent removal of more than 50% of the annual herbaceous growth (by weight).

9) Sagebrush taxa and their value to grouse need to be identified prior to burning. Some subspecies, such as mountain big sagebrush, may invade an area immediately after burning and may not be as desirable for sage-grouse as the original species or subspecies [60].

Breeding: Regardless of the method used to reduce or eliminate sagebrush cover in sage-grouse breeding habitat, control actions have the potential for reducing breeding populations of sage-grouse. Moreover, sagebrush reduction programs may exacerbate the negative effects of natural phenomena such as drought, causing extreme declines in the sage-grouse populations [28]. Martin [82] found 80% of all cock sage-grouse and over 80% of all hen locations were in sagebrush stands with canopy cover exceeding 20%. He comments this is also the range of sagebrush canopy cover where control is most likely to occur.

Before making any alterations of sagebrush habitat in the vicinity of leks, Ellis and others [43] recommend the following: 1) When possible, protect all sagebrush within 1.86 miles (3 km) radius of a lek as suggested by Braun and others [21]. 2) If alteration is unavoidable within 1.86 miles (3 km) of a lek, identify day-use areas. Once the core-use area has been identified, protect it and try to provide a buffer zone around it if possible. Because sage-grouse often walk to day-use areas, it is recommended that a continuous strip (i.e. travel lane) no less than 656 feet (200 m) be maintained between the lek and the day-use areas. 3) If day-use areas are identified and it is concluded that they cannot be protected, the manager should determine their physical sagebrush characteristics. Preservation of statistically similar adjacent stands along with an adjoining travel lane may provide male grouse with necessary habitat to continue using the lek. Any planned alterations should be done after males have moved to summer or fall ranges. Wallestad and Pyrah [123] suggest a 2-mile (3.2 km) buffer zone around a lek should be protected from all sagebrush eradication if the manager has not identified the wintering-nesting complexes associated with it.

A fire in the right place at the right time, and of the correct size and intensity can create an opening that can be used as a lek. Such openings, 1 to 10 acres (0.4-4 ha) in size and at the elevations used for breeding, may be beneficial to sage-grouse in homogeneous sagebrush habitats [34].

Nesting: Sage-grouse prefer light to moderate sagebrush densities for nesting. Where sagebrush is dense (in excess of 20-40% canopy cover) and greater than 2 feet (61 cm) in height, controlled burning to create a mosaic of sagebrush and grassland with a variety of sagebrush heights would probably be beneficial to the birds. Repeated burning could be adverse in this case, as would large "hot" fires that remove an excessive amount of cover [24,71]. Where cover is all ready limited, fires could cause adverse conditions for the birds [58,71].

The U.S. Fish and Wildlife Service currently uses fire as a management tool to improve sage-grouse nesting and brood-rearing habitat at the Hart Mountain National Antelope Refuge (HMNAR) in Oregon [32]. To evaluate recovery of burned vegetation with respect to sage-grouse nesting and brood-rearing requirements, Nelle and others [87] recommend that scheduling when a habitat is burned should not be predetermined by the length of time since the area last burned. By measuring 3 variables (sagebrush height, percent sagebrush canopy cover, and percent total shrub cover) at permanent, randomly located transects within burned vegetation, a fast and efficient monitoring program to follow the recovery of vegetation could be implemented.

Also on the HMNAR, DeLong and others [37] studied nesting habitat in relation to predation. They suggest prescribed fire would reduce shrub cover and may increase herbaceous cover. Sagebrush reduction, however may negatively affect sage-grouse nesting habitat in the short term [30], and should only be implemented in areas where other suitable nesting habitat exists nearby. In the long term, once sagebrush re-establishes in treatment areas, sage-grouse nesting habitat may be enhanced by an improved balance of shrub and grass components available to sage-grouse. They recommend land management practices that increase cover and height of native grasses in sagebrush communities with medium-height shrubs as a means to enhance sage-grouse nesting success and productivity [37].

Pyle and Crawford [96] report increased production of forbs and perennial grasses following fire in sagebrush. Increases in perennial grass cover should improve nesting habitat by providing hens with more cover from predators. Greater forb availability should provide hens with more opportunities to meet their physiological demands for nesting and renesting. Burning may be used to increase herbaceous cover or decrease shrub cover but the shrub component is critical to a sagebrush obligate such as sage-grouse [25]. Frequent burns that promote invasion by exotics such as cheatgrass or large burns that remove the sagebrush overstory may not be used by nesting or brood-rearing sage-grouse until sagebrush becomes re-established [30,70]. Managers must consider size of burns, juxtaposition of habitat components, and frequency of burns if they want to improve habitat while still maintaining all components needed by sage-grouse year round [25].

Burning to remove sagebrush on the Upper Snake River Plain is not justifiable as a sage-grouse management practice because only unburned vegetation in the area offers suitable nesting habitat. No brush control work should occur where live sagebrush cover is less than 20% [21] as it is on the Upper Snake River Plain. From 1980 to 1996, 28,840 acres (11,676 ha) were burned by prescribed fire on the Upper Snake River Plain, which accounted for 29% of the study area, none of which was suitable nesting habitat in 2000 [87]. Connelly and others [28] found no evidence that nest success, and thus nest habitat, increased in the treatment area after a prescribed burn on the Upper Snake River Plain and recommend that prescribed burns be avoided in relatively xeric habitats used by wintering and breeding sage-grouse. During drought periods, they recommend prescribed burns be totally prohibited in these habitats.

Brood rearing: Prescribed burning may have different long-term effects as species diversity of vegetation and invertebrate food resources change with succession [49] and could result in suboptimal nesting and brood-rearing habitat. The cumulative effect of burning upon critical nesting and brood-rearing habitats could be seriously detrimental to a sage-grouse population if vegetation over a large area remains in suboptimal condition for many years. Burn programs should be planned to avoid creating a landscape of adjacent young burns, especially of burns younger than 14 years. Inherent variation in vegetation responses to burning can render predicting the long-term effects of fire difficult [87].

Spot burns in which several patches of a few acres are burned can produce suitable brood-rearing areas. Spot burns along the edges of meadows where sagebrush is encroaching may also enhance brood-rearing areas, although care should be taken to leave an adequate sagebrush-meadow ecotone to provide necessary cover for sage-grouse. Spring burns during the nesting period must be avoided, but late fall burns may be an alternative where brood-rearing habitats are needed. Burning should seldom be allowed on areas where nesting habitat is limited [6]. Wrobleski [130] states increases in percent of annual forb production in burned areas will likely provide a rich forage base for prelaying females and young chicks. Abundant food forbs in close proximity to unburned sagebrush cover could benefit sage-grouse broods by providing additional food with adequate cover. In a study in Idaho, Martin [83] found burning improved habitat for sage-grouse broods. During both postburn years of his study, forb crown cover was significantly higher in burned habitats than in unburned habitats.

Klebenow [71] states broods seek out areas where their favorite food forbs are abundant. He states most sites like this have been depleted by excessive grazing and require rejuvenation, and fire can be an important part of management to improve these lands. Burning small areas to achieve a mosaic of food and cover areas should produce a pattern most suitable for sage-grouse. Different stages of successional growth would be desirable in order to produce the greatest variety of food forb items. Burning should probably be done on a rotational basis, burning different patches each year or every few years, possibly with as long as 20 years between burning treatments on each site. A diversity of habitat types, in terms of both food and cover, should be an objective for this habitat. Using fire to open dense sagebrush canopies in occupied sage-grouse range may enhance brood-rearing areas by increasing forb production according to Sime [109]. She also suggests prescribed burning could be used to mitigate for spring/summer habitat lost to agricultural, industrial, or human development and would benefit both sexes and all ages of a sage-grouse population.

Miller and Eddleman [85] have compiled the following information from various authors on the relative response of forbs common to the sagebrush biome and used as food by sage-grouse:

(S = severely damaged, O = 0 to slight damage, U = undamaged, + = increases, - = declines).

The mountain big sagebrush/antelope bitterbrush habitat type is a key brood-rearing habitat of sage-grouse [70,75]. Fall and spring prescribed fire in dense sagebrush-bitterbrush stands did not adversely influence most primary insects and forbs available on burned sites. Indeed, the food supply on burned sites was enhanced based on the increased frequency of dandelion taxa, total forb cover, total forb diversity, and habitat heterogeneity. However, sage-grouse broods require sagebrush for cover on a daily basis [41], so burning or other practices conducted in montane settings could reduce foraging habitat if an interspersion of treated and untreated sites is not maintained [6]. Pyle and Crawford [96] also found that fall burning increased frequency of dandelion taxa on HMNAR, but apparently had no effect on other primary foods after a study of prescribed burning from 1987 to 1989. June and darkling beetles, primary foods of sage-grouse chicks, were unaffected by prescribed fire. Total forb cover ranged from 13 to 15% on burned plots to 8% on unburned controls during the 2nd growing season. The same prescribed fire reduced total shrub cover from 1987 to 1989. Authors stated fall-burned sites were perhaps most suitable to broods because they supported the greatest amount of dandelion taxa. However, the prescribed fire reduced sagebrush cover which diminished its site value as a food and cover source for broods. Brood cover was changed but not eliminated by prescribed fire. Cover was altered from dominance by shrubs before burning to dominance by forbs, grasses, and sprouting shrubs after burning.

Wrobleski [130] studied greater sage-grouse brood-rearing habitat on the HMNAR in Oregon. He compared burned and unburned plots and found:

1. (Morphological) Reproductive effort by individual plants of at least 5 species was greater in 1st-year burned areas, indicating that fire-enhanced flowering occurred in some of the herbaceous dicot species used by sage-grouse. This increased response was observed in both annual (Phlox spp.) and perennials (Lomatium spp., Modoc hawksbeard, shaggy milkvetch, and longleaf phlox).

2. (Phenological) There was a longer period of active photosynthesis observed in all selected species in burned plots.

3. (Arthropod abundance) Populations of arthropods appear to be able to maintain themselves through disturbance by fire.

Greater numbers of inflorescences and flowers provide prenesting hens and broods with increased quantities of higher quality foods. By extending the period of potential photosynthesis and plant succulence on these herbaceous dicot species, forage quality will be higher later into the dry season [130].

Connelly and others [28] warn against using fire in relatively low precipitation areas dominated by Wyoming big sagebrush to improve brood-rearing habitat for migratory greater sage-grouse. A study by Fischer and others [49] indicated the short-term effects of prescribed fire in a xeric (less than 10 inches (250 mm) of annual precipitation) environment did not enhance brood-rearing habitat and may have been detrimental to grasshoppers, which are important in sage-grouse diets. Abundance of grasshoppers was significantly lower in burned habitat the 2nd (P=0.003) and 3rd (P=0.0001) postburn years.

Summer: In a study by Connelly and others [28] on migratory greater sage-grouse in a xeric region of Idaho, fire apparently did not improve or create summer habitat adjacent to the breeding range. Continuation of long distance movements from the breeding area to summer range up to 4 years after the fire appears to confirm this.

Winter: Any proposed type conversion which includes winter-use areas must be carefully considered. Approval should not be given prior to an on-the-ground inspection of sage-grouse winter distribution during peak snow conditions. What may appear to be an excessive amount of sagebrush during summer months may provide only minimal amounts available above the snow for wintering birds. Burning should seldom be allowed on winter use areas and reseeding may be needed in xeric zones [6]. Critical winter habitats are very vulnerable to treatment since big sagebrush control has traditionally been directed towards dense stands on flat to gentle slopes [45,102]. Braun and others [21] state quality of sage-grouse wintering habitat after sagebrush removal treatments is directly related to the amount of sagebrush remaining. In light of the number of months of use by sage-grouse and the potentially large area that a winter range may influence, removal of sagebrush from these segments would greatly reduce sage-grouse populations over large areas [45]. Loss of sagebrush on a relatively small area, but a relatively large portion of wintering habitat, was followed by a large decline in sage-grouse numbers in Montana [113]. Beck [10] states disturbance of sage-grouse winter habitat in identified winter-use areas should be avoided or kept to a minimum.

Connelly and others [28] suggest that in some instances, especially where the herbaceous understory is badly depleted or junipers (Juniperus spp.) have invaded, fire may be used to improve the ecological condition of a site. If fire is used in this manner, treatment should be designed to preserve sage-grouse winter habitat and allow rapid recolonization of the burned area by sagebrush.

Robertson [103] studied the effects of an August, 1989, prescribed burn on the winter ecology of migratory greater sage-grouse in southeastern Idaho. Before the prescribed fire, greater sage-grouse were located on the proposed treatment area 42% (1988) and 34% (1989) of the time. Following the fire in fall of 1989, only 6% of greater sage-grouse locations were found within the burned area. The prescribed burning of winter range probably caused this change in distribution. The mosaic effect of the vegetation created by the burn should still maintain stands of adequate habitat within the burn, but other factors such as inability to occupy snow roosts, theremoregulatory stress, and intraspecific competition, might preclude the use of these areas as suitable cover. Burning did not appear to adversely influence greater sage-grouse on the study area except for redistributing them. If adequate habitat is not available within 6.21 miles (10 km) (the limit of movements from leks of capture during the study) sage-grouse may suffer higher rates of predation and/or starvation, or sage-grouse may emigrate to areas where habitat is more readily available. Moreover, even if critical wintering habitat is not lost in a fire, loss of habitat required during the breeding/brood-rearing season is possible. The guideline for protecting habitat within 1.86 miles (3 km) of a lek [21] may not provide adequate protection of winter range for a migratory population, because sage-grouse were located over twice that distance from their lek of capture. Since no differentiation on a large scale can be made between winter and spring use areas, sagebrush management should only be initiated when all factors are considered. Any prescribed burning on environments having migratory populations of sage-grouse should be initiated only after critical use areas are identified. Critical wintering areas can be defined as those areas sustaining use by sage-grouse in winters of "above average" snowfall. Habitat may be plentiful in years of average snowfall; however, wintering sage-grouse could be negatively impacted should prescribed burning take place on critical seasonal-use areas.

In another Idaho study conducted in xeric, migratory greater sage-grouse habitat, Connelly and others [28] determined burning likely has a detrimental effect on greater sage-grouse populations, and areas affected by prescribed burns may burn again because of wildfire. Results of their study do not support use of fire to improve rangelands that provide satisfactory sage-grouse habitat. The authors urge managers to refrain from burning in low precipitation (10.4 inches (< 260 mm)) sagebrush habitats that are used by breeding greater sage-grouse.

For specific information on use of fire in various types of sagebrush habitats, see the Fire Effects section of sagebrush reviews on this website.

FIRE CASE STUDIES

• FIRE CASE STUDY CITATION
• REFERENCE
• SEASON/SEVERITY CLASSIFICATION
• STUDY LOCATION
• PREFIRE HABITAT
• SITE DESCRIPTION
• FIRE DESCRIPTION
• FIRE EFFECTS ON ANIMAL SPECIES AND HABITAT
• FIRE MANAGEMENT IMPLICATIONS

Fires were ignited 23-28 September, 1997. Mean estimate of fireline intensity was 1,321 kWm^-1, and reaction intensity was 302 kW/m².

STUDY LOCATION:
Hart Mountain National Antelope Refuge, Oregon

PREFIRE HABITAT:
Total shrub cover before treatment was 26 ± 1.6% in treatment plots and 24 ± 1.5% in control plots.  Sagebrush (Artemisia spp.) cover (22%) on all plots before treatment comprised 87% of all shrub cover. Other common shrubs on the study plots before treatment (2.7% total cover) were spiny hopsage (Atriplex spinosa) and broom snakeweed (Gutierrezia sarothrae). Small communities dominated by low sage (Artemisia arbuscula) and basin big sagebrush (A. tridentata ssp. tridentata) were found within the study area but were not sampled. Prefire shrub density in burned plots ranged from 39,690 to 71,880 individuals acre^-1 (16,063 to 29,091 ha^-1).

There were 8 species of grasses observed before the fire. Sandberg bluegrass (Poa secunda) and bottlebrush squirreltail (Elymus elymoides) both had 45% prefire frequency and were the dominant grasses. Cheatgrass (Bromus tectorum) had a prefire frequency of 18%.

Of 20 annual forb species observed on the study area, only weevil prairie-dandelion (Nothocalais troximoides) with 13% prefire frequency and blue-eyed Mary (Collinsia parviflora) with 37% prefire frequency were found frequently enough before fires to be tested for significance. A total of 40 species of perennial forbs was observed. Perennial forb species tested for significance included Modoc hawksbeard (Crepis modocensis), longleaf phlox (Phlox longifolia), wild onion (Allium spp.), milk-vetch (Astragalus spp.), dwarf yellow fleabane (Erigeron chrysopsidis), and desertparsley (Lomatium spp.).

SITE DESCRIPTION:
Plant response to prescribed fire was evaluated on 4 burned and 4 unburned plots in a Wyoming big sagebrush (A. t. ssp. wyomingensis) ecosystem. Four plots were chosen randomly for prescribed fire treatment. Plots were approximately 988 acres (400 ha) in size and located in the northeastern portion of Hart Mountain National Antelope Refuge. Elevation of the study plots ranged from 5,085 to 5,299 feet (1,550-1,615 m) with level topography. Soils were a cobbly clay-loam of the Ratto-Coglin complex. Mean annual precipitation at refuge headquarters was 11.42 inches (290 mm). Precipitation was 9.8 and 18 inches (249 and 456 mm) in 1997 and 1998, respectively.

FIRE DESCRIPTION:
The prescribed fire plots were ignited with a heli-torch September 23-28, 1997, utilizing strip-head or ring-fire ignition patterns. Ambient temperatures during fires ranged from 66 to 82 degrees Fahrenheit (18.7^o-27.5^oC), relative humidity from 17 to 24%, and wind speed from 4 to 6 miles h^-1 (6.4 to 9.7 km h^-1). Cloud cover was < 10%, with no precipitation for more than 2 weeks. Understory fuel moistures for the dead herbaceous vegetation ranged from 4.4-6.5% and moisture of 10-hour dead fuels from 5.5-8.0%. Flame height was 6.6 to 12 feet (2.0-3.7m), flame length 6.6 to 14.4 feet (2.0-4.4m), flame depth from 6.6 to 25.6 feet (2.0-7.8m), rate of spread ranged from 15 to 39.4 feet/min (4.6-12m/min), and residence time from 0.6 to 2.6 minutes.

Management objective of the prescribed burn was to ascertain effects of prescribed fire on Wyoming big sagebrush and implications for ecological restoration of sage-grouse habitat.

FIRE EFFECTS ON ANIMAL SPECIES AND HABITAT:
Prescribed fire created habitat areas with different species compositions on the landscape. Although structure was altered by the loss of sagebrush dominance, species composition of the burned plots changed very little throughout the study, and no significant differences were observed in alpha or gamma diversity. Frequency of none of the common herbaceous species changed significantly after prescribed fire.

Presence or absence of most species in the plots remained unchanged, but percent cover of species changed dramatically. Fire caused burn plots to have higher cover of annual forbs and sprouting herbaceous species than shrubs.

The 3 most abundant shrub species, Wyoming big sagebrush, spiny hopsage, and broom snakeweed are partially classified as avoiders. They were eliminated from areas that burned (with the exception of postfire sagebrush seedlings and a few sprouting hopsage individuals). Fire killed all sagebrush plants within burned portions of the treated plots (47 ± 12% of the area). However, in burned areas, 49 to 297 sagebrush seedlings acre^-1 (121-734 ha ^-1) were present the 1st postfire year. Although none were sampled in the transects, scattered spiny hopsage individuals were observed germinating, and spiny horsebrush (Tetradymia spinosa) seedlings were also observed. Number of reproductive shoots (257 vs. 80), vegetative shoots (510 vs. 149), and total shoots (767 vs. 229) per plant were significantly greater among plants along burn edges than among plants in unburned interior (Wilcoxon sign rank p=0.0078). Prescribed fire elevated the number of total shoots of sagebrush individuals along burn edges by 243 ± 29% than sagebrush individuals in unburned interior.

Prescribed fire decreased overall cover of grasses (MANOVA, p=0.13). Cover of perennial grasses declined by 4% (9% in 1997 to 5% in 1998) in burned plots, while there was only a 1% change (10% in 1997 to 9% in 1998) in control plots (p=0.027). Burning had no significant effect on cover of tall grasses (< 2% decrease in burn vs. no change in control plots) or annual grasses (1-2% all treatments and years). Density of tall perennial bunchgrasses (2.7 m^-2 in burn, 2.2 in control) in the 1st posttreatment year (1998) was not significantly different between burned and unburned control plots. Of 8 species of grasses observed, there was no significant difference between burn and control plots.

There were significant increases in the percent cover of forbs (forbs selected by sage-grouse, perennial and annual forbs) (MANOVA, p=0.012). Prescribed fire increased cover of annual forbs by 16% (3% in 1997 to 19% in 1998) in burned plots, while only a 5% increase (3% in 1997 to 8% in 1998) was observed in the control plots (p=0.003). No significant differences were detected in cover of grouse forbs or perennial forbs. Fire caused no significant changes in frequency of any annual or perennial forb species.

FIRE MANAGEMENT IMPLICATIONS:
Sagebrush cover on the study area (and in the surrounding area) was high (>25%) due to livestock grazing and fire suppression. There was little interspersion of other habitats in this "sea of sagebrush," yet sage-grouse populations on the refuge continue to decline. Thus, it is likely that on the study area sagebrush is not a limiting factor for sage-grouse populations. Reducing cover of sagebrush and increasing understory vegetation may bring the ecosystem closer to the historic conditions with which sage-grouse evolved.

Nesting habitat for sage-grouse may not be reduced for 3 reasons: Sage-grouse may have evolved with lower sagebrush cover; greater than half of the burned plots did not actually burn; and higher levels of cover exist along burn edges due to elevated vigor of sagebrush. As sagebrush seedlings within the burned areas mature and reproduce, they will begin to add to sage-grouse nest and hiding cover within burned areas. Additionally, with herbaceous cover along burn edges at least equal to control levels, and sagebrush cover along burn edges elevated, these burn edges may prove to be superior nesting habitat for sage-grouse.

Quality of sage-grouse wintering habitat after sagebrush removal treatment is directly related to the amount of sagebrush remaining [21]. This prescribed fire treatment left > 50% (in an irregular mosaic pattern) of the treatment area with fully intact sagebrush cover (>25%). This cover, sagebrush in the control plots, and sagebrush surrounding the study area will likely provide more than adequate wintering cover.

A major danger in using prescribed fire for management of sage-grouse habitat is increasing the fire frequency beyond the rate at which the sagebrush component of the ecosystem can recover. If fire return intervals become frequent enough to exceed the capacity of sagebrush to repopulate, sagebrush cover would be reduced far below historical levels which would be very detrimental to sage-grouse.

Increases in percent cover of annual forbs will likely provide a rich forage base for pre-laying sage-grouse females and young chicks. Within the burned areas, greater forb cover will enhance feeding opportunities. Abundant food forbs in close proximity to unburned sagebrush cover could benefit sage-grouse broods by providing additional foods with adequate cover. Reintroducing prescribed fire to sagebrush ecosystems with "unnaturally" high sagebrush cover may improve habitat quality for sage-grouse by increasing forage quality and quantity, and providing greater overstory and understory cover along burn edges.

Some pronounced effects of prescribed fires occurred at the burn/unburn ecotone. Immediately after the fires the wind moved large quantities of soil and ash to downwind burn edges. Increased density of reproductive and vegetative shoots along edges may be partially a result of this additional soil and ash. Drastic increases in cover of grasses and forbs observed along some edges may be the result of high levels of soil and ash or decreased competition in the adjacent areas.

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