Brood habitat use by Sage Grouse in Oregon

Habitat use by Sage Grouse ( Centrocercus urophasianus hens with broods was examined at Jackass Creek and Hart Mountain, Oregon, from 1989 through 1991. Sage Grouse hens initially selected low sagebrush ( Artemisia spp.) cover types during early brood-rearing, big sagebrush cover types later in the brood-rearing period, and ultimately concentrated use in and near lakebeds and meadows. Areas used by Sage Grouse broods typically had greater forb frequency than did random sites. Hens at Jackass Creek selected sites with forb cover similar to that generally available at Hart Mountain, but home ranges were larger at Jackass Creek because of lower availability of suitable brood-rearing habitat. Differences in habitat use by broods on the two areas were reflected in dietary differences; at Hart Mountain, chicks primarily ate forbs and insects, whereas at Jackass Creek most of the diet was sagebrush. Larger home ranges, differences in diets, and differences in availability of forb-rich habitats possibly were related to differences in abundance and productivity between areas.     

Changes in the distribution and status of sage-grouse in Utah

Sage-grouse ( Centrocercus spp.) were abundant in all of Utah's 29 counties at the time of European settlement wherever sagebrush ( Artemisia spp.) occurred. Greater Sage-Grouse ( C. urophasianus ) inhabited areas north and west of the Colorado River, and Gunnison Sage-Grouse ( C. minimus ) occupied suitable habitat south and east of the Colorado River. The largest Greater Sage-Grouse populations in Utah are currently restricted to suitable habitats in Box Elder, Garfield, Rich, Uintah, and Wayne Counties. A remnant breeding population of Gunnison Sage-Grouse occurs in eastern San Juan County. We stratified Greater Sage-Grouse populations (1971-2000) by counties where the 1996 to 2000 moving average for estimated spring breeding populations was > 500 (GT500) or < 500 (LT500). Males per lek declined in all populations from 1971 to 2000; however, there were consistently more males observed on GT500 than on LT500 leks. Juveniles per adult hen (including yearling hens) Greater Sage-Grouse in the 1973-2000 fall harvest in Box Elder, Rich, and Wayne Counties did not differ from 2.25, a ratio suggesting sustainable or increasing sage-grouse populations. Declines are attributed to loss, fragmentation, and degradation of sagebrush habitat. Sage-grouse conservation ultimately depends on management and enhancement of remaining sagebrush rangelands in Utah.     

Use and selection of brood-rearing habitat by Sage Grouse in south central Washington

Sage Grouse ( Centrocercus urophasianus ) brood-habitat use was examined during 1992 and 1993 at the Yakima Training Center in Yakima and Kittitas counties, Washington. During the 2 yr we followed 30 broods, of which 12 persisted to 1 August (￣ x = approximately 1.5 chicks/brood). Food forb cover was greater at all brood locations than at random locations. Hens with broods in big sagebrush/bunchgrass habitat ( Artemisia tridentata/Agropyron spicatum ) selected for greater food forb cover, total forb cover, and lower shrub heights; broods in altered big sagebrush/bunchgrass habitats selected greater tall grass cover and vertical cover height; broods in grassland showed no preference for any measured vegetation characteristics. During the early rearing period (post-hatching-6 wk) each year, broods selected sagebrush/bunchgrass. Broods in 1993 made greater use of grasslands than in 1992 and selected grassland during the late brood-rearing period (7-12 wk). Broods selected for sagebrush/bunchgrass during the midday, but 52% of brood locations in the afternoon were in grassland. Tall grass cover was greater at morning (0500-1000 h) and afternoon (1501-2000 h) brood locations than at midday (1001-1500 h) and random locations. Midday brood locations had greater shrub cover and height than morning and afternoon locations. Selection of habitat components was similar to the results of other studies, but habitat conditions coupled with a possible lack of alternate brood-rearing cover types resulted in low survival of chicks.     

Species diversity and habitat complexity: does vegetation organize vertebrate communites in the Great Basin?

In this study, we have examined the effect of vegetation structure on the three major vertebrate taxa in Great Basin habitats of southwestern Utah. The effect of increasing vegetation heterogeneity, both horizontally and vertically, on the diversities of lizards, rodents, and postbreeding birds was investigated. We found no statistically significant relationship between diversity of all animal taxa and horizontal vegetation heterogeneity, although lizard diversity tended to decrease with increasing heterogeneity and rodent diversity tended to increase. Bird species diversity was positively correlated with vertical habitat heterogeneity. Abundances were highest for rodents in pinyon/juniper habitat and highest for lizards and birds in areas with the highest grass cover. Species richness was highest in sagebrush habitat for rodents but highest for lizards and birds in pinyon/juniper. Evenness values were relatively similar and high for birds and rodents and were relatively high for lizards in all habitats except for pinyon/juniper, which had an evenness value of 0.38. For rodents and lizards, abundance was significantly correlated with the index for horizontal habitat heterogeneity. After logarithmic transformation, abundance of lizards was positively correlated with increasing vegetation complexity. Combined abundance of lizards and rodents was also positively correlated with vegetation complexity. Rodent and lizard abundances, however, were affected by different aspects of the habitat. After logarithmic transformation, lizard abundances increased significantly with increasing grass cover, whereas rodent abundances increased significantly with increasing shrub cover.     

Comparison of habitat attributes at sites of stable and declining long-billed curlew populations

Long-billed curlew populations were studied in the upper Green River Basin of Wyoming. Sites were selected where curlew populations appear constant in numbers and declining in numbers. Results show that while few habitat differences were found between the two areas, disturbances such as grazing and dragging during nesting reduced productivity. Nest failures were also correlated with field fertilization and early season grazing. Within each area curlews nested successfully on field sites that were elevated and had adequate grass cover but not tall grass.     

Vegetation response to prescribed fire in Dinosaur National Monument

Much of western North America is dominated by dense, monotypic, late seral stands of big sagebrush ( Artemisia tridentata Nutt.). These stands often have depauperate understories with limited species richness, diversity, and herbaceous cover. The National Park Service at Dinosaur National Monument, Colorado, is using both strategic and natural prescribed fire in Wyoming big sagebrush ( Artemisia tridentata ssp. wyomingensis Beetle and Young) communities to foster intra-community (α -scale) and landscape diversity. This study analyzed an accumulated foliar cover data set between paired burn and control areas on 6 different sites during the last 20 years. Across the monitoring period, mean total vegetation cover of all combined sites was 44% control and 42% burn. Total vegetation cover in burn areas was higher than or equal to paired control areas within 2-3 years post-burn. Shrubs were essentially eliminated in burn areas, but perennial grass cover was 10-35% higher. Mean number of species on all sites and years combined was 17 control and 18 burn. Species richness was different on only 1 site-year, Dry Woman 1995 ( P = 0.001, 15 control, 9 burn). Species similarity by site and between treatments ranged from 44% to 75%. Differences in Shannon-Weiner diversity index values between paired sites occurred in 6 of 20 years ( P < 0.05). Index value differences on these 6 sites were due to a large annual grass component in burn areas. Prescribed burning successfully shifted late successional sagebrushdominated communities to earlier herbaceous-dominated successional stages without lowering total vegetation cover, while maintaining -scale diversity and species richness.     

Suitability of shrub establishment on Wyoming mined lands reclaimed for wildlife habitat

Restoring coal mined land to pre-mining shrub cover, density, height, community composition, and diversity to renew wildlife habitat quality is a priority for reclamation specialists. Long-term shrub reestablishment success on reclaimed mined land in Wyoming and suitability of these lands for wildlife habitat are unknown. Fourteen reclaimed study sites, 10 yr old or older, were selected on 8 mines in Wyoming to evaluate shrub reestablishment and wildlife habitat value for antelope ( Antilocapra americana ) and sage grouse ( Centrocercus urophasianus ). Five sites were categorized as fourwing saltbush ( Atriplex canescens ) sites and 9 as fourwing saltbush/big sagebrush ( A. canescens/Artemisia tridentata spp. wyomingensis ) sites. Published data describing antelope and sage grouse-preferred habitat requirements in sagebrush-grassland steppe ecosystems were used to evaluate shrub community value of sampled sites for wildlife habitat. Mean shrub canopy cover, density, and height for fourwing saltbush sites were 5.8%, 0.23 m -2 , and 41.6 cm, respectively, compared to 5.6%, 0.61 m -2 , and 31.1 cm for fourwing saltbush/big sagebrush sites. Two fourwing saltbush and 4 fourwing saltbush/big sagebrush sites provided sufficient cover for antelope, while 2 fourwing saltbush and 4 fourwing saltbush/big sagebrush sites were adequate for sage grouse. Only 1 fourwing saltbush/big sagebrush site provided high enough shrub densities for sage grouse. One fourwing saltbush and 7 fourwing saltbush/big sagebrush sites provided ample shrub heights for antelope, while 1 fourwing saltbush and 8 fourwing saltbush/big sagebrush sites were sufficient for sage grouse. One fourwing saltbush and 1 fourwing saltbush/big sagebrush site provided enough grass, forb, and shrub composition for antelope, while no site in either reclamation type was satisfactory for sage grouse. Shrub diversity was 3 times higher for fourwing saltbush/big sagebrush sites (0.984) than for fourwing saltbush sites (0.328). Individually, sites seeded with multiple shrub species had higher canopy cover, density, and diversity compared with single-species shrub seedings. Achieving premining shrub cover, density, height, community composition, and diversity within existing bond-release time frames is unrealistic, considering that some native shrublands require 30-60 yr to reach maturity.     

Ecological classification and monitoring model for the Wyoming big sagebrush shrubsteppe habitat type of northeastern Wyoming

The purposes of this study were to develop a multivariate statistical model related to plant succession, to classify by seral stage, and to monitor succession in Wyoming big sagebrush shrubsteppe habitat ( Artemisia tridentata ssp. wyomingensis / Agropyron smithii–Bouteloua gracilis ) in Thunder Basin National Grassland, Wyoming. This model can be used by range and wildlife managers to evaluate management alternatives by assessing changes in plant species cover and composition within and between seral stages. Four ecological seral stages that represent early to late succession were quantitatively identified with an estimated 92% accuracy. Three key plant species provided the necessary information to define seral stages and monitor trends. Percent canopy cover and percent frequency (used to calculate index values: % canopy cover × % frequency of occurrence) of Wyoming big sagebrush ( Artemisia tridentata ssp. wyomingensis ), western wheatgrass ( Agropyron smithii ), and blue grama ( Bouteloua gracilis ) are the only field measurements required for this model.     

Predation of artificial Sage Grouse nests in treated and untreated sagebrush

We measured predation on 120 artificial Sage Grouse ( Centrarcus urophasianus ) nests in montane sagebrush grassland in northern Utah. We examined nests in areas that had been chained and seeded 25 years previously (treated areas) and in areas that were untreated. Predation rates of artificial nests were higher in areas of untreated sagebrush, even though these areas had greater sagebrush cover, taller shrubs, and greater horizontal plant cover. These results differ from those previously hypothesized for treated sagebrush habitat and may reflect a greater abundance of other potential prey species, especially lagomorphs, in untreated areas that attracted greater densities of predators. In addition, over 80% of nests were depredated by mammals, which hunt using olfaction and are less likely than avian predators to be affected by nest cover. We conclude that, after treated sagebrush has recovered to some degree, predation rates of Sage Grouse nests may be lower in treated sagebrush. Consequently, factors other than nest predation (e.g., winter food, thermal cover, insects, perennial forb abundance) may be more important reasons for preserving mature sagebrush stands for Sage Grouse.     

Comparison of Sage and Sharp-tailed Grouse leks in south central Wyoming

Columbian Sharp-tailed Grouse ( Tympanuchus Phasianellus columnianus ) and Sage Grouse ( Centro-cercus urophasianus ) leks were compared in an area of sympatry in south central Wyoming. Sharp-tailed Grouse leks had more (P < .05 shrub cover, taller shrubs, more forb, grass, and shrub species, and less visibility than did Sage Grouse leks. Reduction in shrub cover or the diversity of herbaceous species could potentially have greater influence on the use of lek sites by Columbian Sharp-tailed Grouse than by Sage Grouse in areas of sympatry in Wyoming.     

Long-term effects of tebuthiuron on Bromus tectorum

Use of herbicides to thin dense stands of Artemisia spp. (sagebrush) can free up resources for herbaceous plants and increase forage production, but may also facilitate weed invasion. We revisited a sagebrush thinning experiment in a north central Wyoming big sagebrush–grassland 11 years after application of tebuthiuron (N-[5-(1,1- dimethylethyl)-1,3,4-thiadiazol-2-yl]-N-N′-dimethylurea) to determine the long-term responses of shrubs, available soil resources, perennial grasses, and Bromus tectorum L. (downy brome). Tebuthiuron reduced shrub cover by more than half, from 31% in untreated plots to 15% in treated plots ( P = 0.002), and increased downy brome cover approximately 4-fold, from 0.9% in untreated plots to 3.5% in treated plots ( P = 0.02). Treatment with tebuthiuron also resulted in marginally significant increases in cover of perennial grasses (from 9% to 12.3%; P = 0.07) and bare ground (from 39.1% to 43.9%; P = 0.08). In comparisons of resource availability among microsites, available NO 3 was higher under dead sagebrush than under live sagebrush ( P = 0.03). No significant differences in soil water content were detected. The relatively recent expansion of downy brome populations at this site and the high NO 3 –N levels observed under dead sagebrush suggest that conditions facilitating downy brome invasion may persist for many years following sagebrush thinning. We demonstrate that sagebrush thinning can cause increases in downy brome populations years after initial treatment and suggest that managers should use caution when considering thinning sagebrush if downy brome is present, even if initial populations are small.     

Comparative habitat use in a juniper woodland bird community

We compared vegetation structure used by 14 bird species during the 1998 and 1999 breeding seasons to determine what habitat features best accounted for habitat division and community organization in Utah juniper ( Juniperus osteosperma ) woodlands of southwestern Wyoming. Habitat use was quantified by measuring 24 habitat variables in 461 bird-centered quadrats, each 0.04 ha in size. Using discriminant function analysis, we differentiated between habitat used by 14 bird species along 3 habitat dimensions: (1) variation in shrub cover, overstory juniper cover, mature tree density, understory height, and decadent tree density; (2) a gradient composed of elevation and forb cover; and (3) variation in grass cover, tree height, seedling/sapling cover, and bare ground/rock cover. Of 14 species considered, 9 exhibited substantial habitat partitioning: Mourning Dove ( Zenaida macroura ), Bewick's Wren ( Thryomanes bewickii ), Blue-gray Gnatcatcher ( Polioptila caerulea ), Mountain Bluebird ( Sialia currucoides ), Plumbeous Vireo ( Vireo plumbeus ), Green-tailed Towhee ( Pipilo chlorurus ), Brewer's Sparrow ( Spizella breweri ), Dark-eyed Junco ( Junco hyemalis ), and Cassin's Finch ( Carpodacus cassinii ). Our results indicate juniper bird communities of southwestern Wyoming are organized along a 3-dimensional habitat gradient composed of woodland maturity, elevation, and juniper recruitment. Because juniper birds partition habitat along successional and altitudinal gradients, indiscriminate woodland clearing as well as continued fire suppression will alter species composition. Restoration efforts should ensure that all successional stages of juniper woodland are present on the landscape.     

Influence of fire and juniper encroachment on birds in high-elevation sagebrush steppe

We examined relationships between high-elevation sagebrush ( Artemisia spp.) steppe habitats altered by prescribed fire and western juniper ( Juniperus occidentalis ) encroachment on breeding distributions of Brewer's Sparrows ( Spizella breweri ), Vesper Sparrows ( Pooecetes gramineus ), Green-tailed Towhees ( Pipilo chlorurus ), and Sage Thrashers ( Oreoscoptes montanus ) on Steens Mountain in southeastern Oregon. In 2000 we conducted fixed-radius point count surveys at 172 sites encompassing burned and unburned sagebrush habitat and a range of juniper densities. For each bird species we developed habitat models using local variables measured in the field and landscape variables derived from remotely sensed data. Akaike's Information Criterion (AIC c ) was used to select the best-approximating model from a suite of a priori candidate models. Brewer's Sparrows, Sage Thrashers, and Green-tailed Towhees were positively related to increasing local sagebrush cover or percent sagebrush in the landscape, whereas Vesper Sparrows were negatively associated with sagebrush cover and positively related to increases in sagebrush fragmentation at local and landscape scales. Including a measure of juniper encroachment substantially improved models for all species in the analysis. Green-tailed Towhees showed a curvilinear response to the amount of juniper in the landscape. All other species showed a strong negative relationship with juniper. Our results indicate that, although changes in sagebrush habitat associated with fire had a negative influence on sagebrush birds, juniper encroachment due to fire suppression also impacted this high-elevation sagebrush bird community.     

Brood-rearing habitat use by Rio Grande Wild Turkeys in Oregon

Wild turkey ( Meleagris gallopavo ) brood-rearing sites have been described for portions of their range, but brood-rearing habitat use and characteristics of brood-rearing sites used by Rio Grande Wild Turkeys ( M. g. intermedia ) in the Pacific Northwest were unknown. We described cover types at 362 brood-rearing sites and measured habitat characteristics at 64 of these sites used by a recently established Rio Grande Wild Turkey population in southwestern Oregon during May-September 1989 and 1990. Hens with broods used 9 of 10 available cover types. Meadows, mixed hardwood/conifer woodlands, and savannas were used more often than expected (47% of observations, P ≤ 0.05). Broods used mature mixed conifer and dense sapling/pole mixed conifer cover types less than expected and did not use brush-fields. Many brood-rearing sites were characterized by a parklike appearance. Understory vegetation averaged P ≤ 0.05). We suggest that land managers maintain mixed hardwood/conifer woodland and savanna cover types adjacent to meadows on south slopes to provide brood-rearing habitat for Rio Grande Wild Turkeys in southwestern Oregon.     

Habitat use by breeding male sage grouse: a management approach

Radio telemetry was used to study habitat use of breeding male sage grouse ( Centrocercus urophasianus ) at a lek in northeastern Utah during 1983 and 1984. Objectives were to determine if grouse day-use areas differed significantly in sagebrush characteristics from adjacent nonuse areas and to establish a simplified method for use by land managers in identifying grouse use areas. We determined that male grouse used areas of greatest sagebrush height and cover. Our methods provide a means for land managers to identify habitat associated with a lek that is suitable for male sage grouse day use in the event sagebrush alteration is planned within 3 km of a lek.     

Habitat use patterns within the home range of pygmy rabbits ( Brachylagus idahoensis ) in southeastern Idaho

Pygmy rabbits ( Brachylagus idahoensis ) are a small sagebrush ( Artemisia tridentata ) obligate lagomorph found within the Great Basin of northwestern United States. Because of its reliance on sagebrush, this species is thought to be experiencing a major range reduction as a result of loss of sagebrush habitat. To aid in conservation of this species, we need to better understand its use of the sagebrush environment. We estimated summer home range use patterns by relocating 5 radio-collared pygmy rabbits (3 females and 2 males) over a 24-hour cycle. We then compared soil texture, shrub density, height, and canopy cover between areas close to burrow entrances and areas of high use and low use. Mean home range sizes of female and male rabbits were 37.2 and 67.9 ha, respectively. Rabbits had disproportionate amounts of time (68.4% ± 9.1, s￣x ) and travel (63.0% ± 5.7, s￣x ) in areas within a 60-m radius of their burrows. Soil texture did not differ among the 3 areas, but shrub density, specifically big sagebrush, and forb density were significantly higher close to the burrow than in the high- and low-use areas. We conclude that pygmy rabbits are possibly burrow obligates and that their abundance and distribution are likely limited by available burrow sites.     

Summer habitat use and selection by female Sage Grouse ( Centrocercus urophasianus ) in Oregon

Cover types and vegetative characteristics (e.g., grasses, forbs, shrubs) used by female Sage Grouse ( Centrocercus urophasianus ) during summer were compared with available habitat on two study areas in southeastern Oregon. Broodless hens, which constituted 114 of the 125 (91%) radio-marked hens studied, selected big ( Artemisia tridentata ) subspp.) and low sagebrush ( A. arbuscular ) cover types at both study areas. At Hart Mountain, broodless hens did not select specific vegetative characteristics within cover types. However, at Jackass Creek, forb cover was greater ( P = .004) at broodless hen sites than at random locations. Differences in habitat use by broodless hens between study areas were associated with differences in forb availability. Broodless hens used a greater diversity of cover types than hens with broods. Broodless hens gathered in flocks and remained separate from but near hens with broods during early summer. By early July broodless hens moved to meadows while hens with broods remained in upland habitats.     

Passerine response to roads associated with natural gas extraction in a sagebrush steppe habitat

Natural gas extraction and field development are pervasive throughout the sagebrush steppe of Wyoming. We conducted this study to determine how roads associated with natural gas extraction affect the distribution of breeding songbirds in sagebrush steppe habitat. The study encompassed dirt and paved roads in the Jonah Field II and Pinedale Anticline Project Area in Sublette County, Wyoming. Sites are dominated by Wyoming big sagebrush ( Artemisia tridentata ), and common passerines include sagebrush obligates: Brewer's Sparrows ( Spizella breweri ), Sage Sparrows ( Amphispiza belli ), and Sage Thrashers ( Oreoscoptes montanus ); and non-obligates: Horned Larks ( Eremophila alpestris ) and Vesper Sparrows ( Pooecetes gramineus ). Species relative density was measured using 50-m-radius point counts during spring 1999 and 2000. Four roads with low traffic volumes (700-10 vehicles per day) were surveyed and point counts were centered at variable distances from the road surface such that relative densities were measured 0-600 m from the road's edge. Density of sagebrush obligates, particularly Brewer's and Sage Sparrow, was reduced by 39%-60% within a 100-m buffer around dirt roads with low traffic volumes (700-10 vehicles per day). While a 39%-60% reduction in sagebrush obligates within 100 m of a single road may not be biologically significant, the density of roads created during natural gas development and extraction compounds the effect, and the area of impact can be substantial. Traffic volume alone may not sufficiently explain observed declines adjacent to roads, and sagebrush obligates may also be responding to edge effects, habitat fragmentation, and increases in other passerine species along road corridors. Therefore, declines may persist after traffic associated with extraction subsides and perhaps until roads are fully reclaimed.     

Grazing impacts on sagebrush communities of central Utah

Twenty-three study sites were established in the sagebrush communities bordering Utah Lake. Relic (ungrazed) and grazed stands were represented in the sample. Differences in species composition, vegetation, and soil characteristics were assessed. Major differences in species composition and vegetative characteristics were due to the influence of grazing. Major changes were loss of native perennial grass cover, and increases in cover from introduced annual species. Differences in the soil characteristics were due to habitat rather than grazing influences.     

Effects of browsing by native ungulates on the shrubs in big sagebrush communities in Yellowstone National Park

The effects of elk ( Cervus elaphus ), pronghorn ( Antilocapra americana ), and mule deer ( Odocoileus hemionus ) browsing on shrubs in big sagebrush ( Artemisia tridentata ) communities were monitored over a 31-year period in Yellowstone National Park. Ungulates were restricting Wyoming big sagebrush (spp. wyomingensis ) heights, size, and recruitment on the lower-elevation stratum only, while no such suppression was observed on the high-elevation stratum. Parallel increases in mountain big sagebrush (spp. vaseyana ) densities and cover occurred over the study period on both browsed and unbrowsed sites at the higher-elevation stratum, although big sagebrush, green rabbitbrush ( Chrysothamnus viscidiflorus ), and horsebrush ( Tetradymia canescens ) were slightly taller and crown sizes were slightly larger on unbrowsed than browsed sites. Wyoming big sagebrush utilization (percent leader use) was eight times higher (￣ x = 87 ± 7.2% by pronghorns, mule deer, and elk) on the low-elevation winter ranges stratum (the Boundary Line Area [BLA] portion of the winter range), while mostly mountain big sagebrush with leader use averaged only 11 ± 4.1% (nearly all by elk) on the high-elevation range stratum. In addition, annual aboveground biomass production of big sagebrush did not differ between browsed and unbrowsed study sites on the high-elevation stratum of the winter range. Population turnover was higher on browsed plots versus unbrowsed plots. No difference was observed in percent dieback of big sagebrush adult plants between browsed and unbrowsed plots at the higher stratum. Browsing did not influence the number of leaves or seedstalks per plant ( P > .05), but leaves averaged 45% longer and seedstalks 42% longer on browsed big sagebrush. Ungulate browsing, however, apparently suppressed production, germination, and survival of Wyoming big sagebrush on the low-elevation stratum. Numbers of Wyoming big sagebrush declined 43% and cover declined 29%, 1957-1990, on browsed sites on the BLA. Annual biomass production on browsed sites at the low-elevation stratum was only 6-35% that of unbrowsed sites, and big sagebrush recruitment was less on browsed sites. Percent leader use of big sagebrush did not differ between the period of ungulate reductions, 1962-1969, and the 1980s on the lower stratum (￣ x = 87% leader use), but utilization was less on higher portions of the winter range during the period of elk reductions (￣ x = 2%) than during the 1980s following cessation of elk controls (￣ x = 11%).