A sagebrush community type classification for mountainous northeastern Nevada rangelands

Vegetation characteristics of 15 sagebrush community types identified on the Humboldt National Forest, northeastern Nevada, are described. A total of 218 plant species were found over the 372 relatively undisturbed rangeland communities sampled. The dominant plant families encountered were the Asteraceae (45 taxa), Poaceae (32 taxa), Scrophulariaceae (14 taxa), and the Fabaceae (12 taxa). Average annual dry weight production of the community types ranged from about 400 kg/ha/yr on types with Artemisia nova as the dominant sagebrush species to 1,200 kg/ha/yr on some A. tridentata ssp. vaseyana community types. A general increase in species richness and vegetation plus litter ground cover was observed within community types as the dominant sagebrush species changed from A. nova to A. arbuscula to A. longiloba to A. tridentata spp. tridentata to A. tridentata ssp. vaseyana . Major differences in plant species production and constancy exist between the sagebrush community types studied.     

Mule deer foraging preference among five sagebrush ( Artemisia L.) taxa

The hypothesis that sagebrush taxa are equally utilized by mule deer ( Odocoileus hemionus hemionus ) on winter range was tested. Five taxa were studied for 10 years at 2 locations. The taxa were Artemisia tridentata ssp. tridentata (basin big sagebrush), A. t. ssp. wyomingensis (Wyoming big sagebrush), A. t. ssp. vaseyana (mountain big sagebrush), A. tripartita ssp. tripartita (tall threetip sagebrush), and A. arbuscula ssp. arbuscula (low sagebrush). Possible mule deer preferences were determined each year individually for the 2 sites. Utilization was high enough to conclude all taxa are important forage, but not excessive enough to mask preference. Artemisia tridentata ssp. vaseyana (34.4%) and A. arbuscula ssp. arbuscula (35.6%) were preferred over A. t. ssp. wyomingensis (10.9%) and A. t. ssp. tridentata (6.8%) at the Ashbough site. At the Scudder site there were few differences in preference for A. t. ssp. vaseyana (32.1%), A. t. ssp. wyomingensis (28.8%), and A. tripartita ssp. tripartita (32.0%). While ungulates often demonstrate a preference among taxa, all sagebrush taxa are a potentially valuable forage source.     

Correlation of neighborhood relationships, carbon assimilation, and water status of sagebrush seedlings establishing after fire

Interactions of Artemisia tridentata ssp. vaseyana (mountain big sagebrush) and neighboring herbs may affect community development following fire in sagebrush steppe. Dry mass, photosynthesis, and water relations were measured for seedlings of A. tridentata ssp. vaseyana occurring at different distances from neighboring herbs in the initial growing seasons following fire, when herbs dominate plant community cover. Seedling mass significantly increased as distance to neighboring herbs increased, although a low r 2 indicated that mass was also affected by other sources of variation. Carbon assimilation ( A net ) was also greater for A. tridentata ssp. vaseyana seedlings in microsites farther from herbs, except during those sampling dates when A net and water availability were at low levels. Contrary to our expectations, water status of seedlings was not correlated with their distances to neighboring herbs, and supplemental watering did not affect the slope of the relationship between seedling mass and distance to neighboring herbs. These findings suggest that negative relationships between A. tridentata ssp. vaseyana seedlings and herbs establishing after fire are not likely due to competition over water and may instead result from interactions over factors not measured here (e.g., nutrients).     

Soil characteristics of mountainous northeastern Nevada sagebrush community types

Soil physical and chemical properties were studied to identify habitat differences among 15 sagebrush dominated plant community types of the Great Basin. The sagebrush taxa studied followed an apparent gradient of increasing soil fertility, with Artemisia nova commonly occupying the lowest fertility sites. Sites of moderate soil fertility tended to support A. arbuscula , A. longiloba , and A. tridentata ssp. Wyomingensis . Artemisia tridentata ssp. Tridentata and A. tridentata ssp. Vaseyana were commonly found on sites with the highest soil fertility levels as indicated by maximum organic carbon, nitrogen, and phosphorus levels within such soils. Mollic epipedon depth, total depth, and water-holding capacity of the soil also increased as the dominant sagebrush taxon of a community type changed from A. nova to A. tridentata ssp. Vaseyana . Data from this study provide information concerning soil relationships within relatively undisturbed rangeland communities. Such data are required if future soil sampling efforts within the Great Basin are to have a basis for comparison.     

Applicability of the universal soil loss equation for southeastern Idaho wildlands

Soil physical and chemical properties were studied to identify habitat differences among 15 sagebrush dominated plant community types of the Great Basin. The sagebrush taxa studied followed an apparent gradient of increasing soil fertility, with Artemisia nova commonly occupying the lowest fertility sites. Sites of moderate soil fertility tended to support A. arbuscula , A. longiloba , and A. tridentata ssp. Wyomingensis . Artemisia tridentata ssp. Tridentata and A. tridentata ssp. Vaseyana were commonly found on sites with the highest soil fertility levels as indicated by maximum organic carbon, nitrogen, and phosphorus levels within such soils. Mollic epipedon depth, total depth, and water-holding capacity of the soil also increased as the dominant sagebrush taxon of a community type changed from A. nova to A. tridentata ssp. Vaseyana . Data from this study provide information concerning soil relationships within relatively undisturbed rangeland communities. Such data are required if future soil sampling efforts within the Great Basin are to have a basis for comparison.     

Selection of big sagebrush by sage grouse

Feeding sites of wintering sage grouse ( Centrocercus urophasianus ) were located, one each in stands of three subspecies of big sagebrush ( Artemisia tridentata : ssp. tridentata , basin; ssp. vaseyana , mountain; and ssp. Wyomingensis , Wyoming). Evidences of differential use of plants within subspecies were observed. Whole leaves from fed-on and nonfed-on big sagebrush plants were examined for intrasubspecies chemical comparisons of crude protein, phosphorus, in vitro digestibility, and monoterpenoids. No significant differences were detected except for in vitro digestibility of Wyoming fed-on and nonfed-on big sagebrush and monoterpenoid content of basin big sagebrush. Nutritive content of all three subspecies was high, which may in part help to explain wintering sage grouse weight gains.     

Randomly amplified polymorphic DNA analysis (RAPD) of Artemisia subgenus Tridentatae species and hybrids

Species of Artemisia (subgenus Tridentatae ) dominate much of western North America. The genetic variation that allows this broad ecological adaptation is facilitated by hybridization and polyploidization. Three separate studies were performed in this group using randomly amplified polymorphic DNA (RAPD). Fifty-seven 10-mer primers generated nearly 400 markers from genomic DNA obtained from leaf tissue. These studies were (1) a measure of the variability of plants within and between populations and between subspecies using 5 A. tridentata ssp. wyomingensis populations, 2 A. cana ssp. cana populations, and 1 A. cana ssp. viscidula population; (2) an examination of the hypothesis that tetraploid (4 x ) Artemisia tridentata spp. vaseyana derives de novo from diploid (2 x ) populations via antopolyploidy; and (3) an examination of the validity of the status of putative hybrids that have been produced by controlled pollination. These later hybrid combinations- A. tridentata ssp. tridentata × A. t. ssp. vaseyana , A. t. ssp. wyomingensis × A. tripartita , and A. cana ssp. cana A. tridentata ssp. wyomingensis - were made to combine traits of parental taxa in unique combinations with possible management application. RAPD marker data were subjected to similarity and UPGMA clustering analyses. RAPD markers were effective in measuring genetic diversity at different systematic levels. Individual plants within a population were approximately 55% to > 80% similar to one another; populations within subspecies gave corresponding values of similarity, probably a result of the combined effects of large population sizes and wind pollination. The 2 subspecies of A. cana were approximately 45% similar. At least some 4 x populations of A. tridentata ssp. vaseyana apparently derive de novo from 2 x plants based on their being embedded in 2 x phenogram groups, thus reinforcing evidence that autopolyploidy plays an important role in Tridentatae population biology. Two ( A. tridentata ssp. tridentata × A. t. ssp. vaseyana and A. cana ssp. cana × A. tridentata ssp. wyomingensis ) of the 3 putative hybrid combinations were confirmed to include hybrids. These hybrids may have potential in management applications. Additional use of RAPD technology combined with other techniques may be useful in delimiting genetic characteristics and in guiding artificial selection in Tridentatae .     

Seasonal patterns of arthropod diversity and abundance on big sagebrush, Artemisia tridentata

The sagebrush biotype is the largest in the western United States. This vast sagebrush community is thought to harbor equally vast and diverse arthropod communities, but these remain little explored. Our objective was to examine the diversity, abundance, and seasonal phenology of arthropod taxa found on the dominant shrub of the sagebrush ecosystem, big sagebrush ( Artemisia tridentata ). We wanted to improve understanding of this little-studied arthropod assemblage that may play significant roles in the dynamics of sagebrush populations and the sagebrush ecosystem. We sampled free-living and gall-forming arthropods from a stratified random sample of sagebrush plants at the Barton Road Ecological Research Area, Idaho, resulting in a sample of over 8000 individuals and 232 morphospecies. Species richness and abundance declined from May to August, and abundance of most taxa similarly declined over the summer. A few taxa, including Acari (mites), were notably more abundant in August. Fluid feeders were the most diverse and abundant free-living feeding guild during all months and comprised up to 79% of morphospecies. The gall formers included 4713 individuals of 12 species of gall flies ( Rhopalomyia spp.), primarily (97%) R. ampullaria. Abundance of galls increased from small to large (presumably young to old) plants. Overall, A. tridentata was host to a high diversity of arthropods, some of which have potential to cause or mitigate significant damage to their host plant. Arthropods seem likely to have the greatest impact on sagebrush early in the growing season, when they are most diverse and abundant. Documentation of the full diversity of arthropods associated with sagebrush required samples taken throughout the growing season, but a single sample early in the growing season captured a high proportion of taxa.     

A new combination and a new variety in Artemisia tridentata

The combination Artemisia tridentata Nutt. ssp. spiciformis (Osterhout) Goodrich & McArthur comb. nov. is made. This high elevation taxon was originally described at the species level and more recently has generally been treated as a form of A. tridentata ssp. vaseyana . The subspecies designation is supported by its parallel nature to the other A. tridentata  subspecies and by its relatively widespread and locally abundant populations. Also, a new variety of A. tridentata ssp. vaseyana is proposed.         

Planting depth of 'Hobble Creek' mountain big sagebrush seed

We conducted a greenhouse study in which 'Hobble Creek' mountain big sagebrush ( Artemisia tridentata ssp. vaseyana ) seeds were planted at various depths in soil to determine the optimal planting depth. Results showed that the optimal planting depth is 5 mm or less.     

Controls on radial growth of mountain big sagebrush and implications for climate change

Mountain big sagebrush ( Artemisia tridentata Nutt. ssp. vaseyana ) covers large areas in arid regions of western North America. Climate-change models predict a decrease in the range of sagebrush, but few studies have examined details of predicted changes on sagebrush growth and the potential impacts of these changes on the community. We analyzed effects of temperature, precipitation, and snow depth on sagebrush annual ring width for 1969 to 2007 in the Gunnison Basin of Colorado. Temperature at all times of year except winter had negative correlations with ring widths; summer temperature had the strongest negative relationship. Ring widths correlated positively with precipitation in various seasons except summer; winter precipitation had the strongest relationship with growth. Maximum snow depth also correlated positively and strongly with ring width. Multiple regressions showed that summer temperature and either winter precipitation or maximum snow depth, which recharges deeper soil horizons, are both important in controlling growth. Overall, water stress and perhaps especially maximum snow depth appear to limit growth of this species. With predicted increases in temperature and probable reduced snow depth, sagebrush growth rates are likely to decrease. If so, sagebrush populations and cover may decline, which may have substantial effects on community composition and carbon balance.     

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.     

Vegetal change on a northern Utah foothill range in the absence of livestock grazing between 1948 and 1982

Reexamination of a semiarid foothill rangeland, first evaluated in 1948, indicated that secondary succession continues to shift toward a perennial grass-forb community formerly dominated by xeric shrubs, particularly big sagebrush ( Artemisia tridentata spp. vaseyana ). The direct role in livestock grazing in establishment and maintenance of shrub-dominant plant communities appears confirmed in the decline of shrubs upon cessation of livestock grazing in summer and continued browsing by mule deer in winter. The reduction of shrub forages on mule deer winter ranges is a major factor in population declines.     

Plant age/size distributions in black sagebrush ( Artemisia nova ): effects on community structure

The demography of black sagebrush ( Artemisia nova Nelson) was investigated in the Buckskin Mountains of western Nevada to determine patterns of stand renewal in sagebrush communities currently free from wildfires. Biomass sampling was conducted to develop growth classes that reflected apparent age of the shrubs. The density of black sagebrush plants was twice that of basin big sagebrush ( A. tridentata ssp. tridentata Nutt.) in adjacent communities on contrasting soils (2.2 versus 1.1 plants per m 2 ). Black sagebrush accumulated only 75% as much woody biomass as big sagebrush. Regression equations were developed and tested for predicting total woody biomass, current annual growth (CAG), and leaf weight of black sagebrush plants. Apparent age classes were developed both for the black sagebrush plants and the sub-canopy mounds on which they grew. Discriminant analysis was used to test this classification system. Plant succession, apparently controlled by nitrate content of the surface soil, appeared to eliminate the successful establishment of black sagebrush seedlings on the mounds. After the shrubs die, the mounds eventually deflate. We propose that mounds reform around shrub seedlings, but because seedling establishment is so rare in these communities, this could not be verified.     

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%).     

Spatial relationships among young Cercogarpus ledifolius (curlleaf mountain mahogany)

This study analyzed spatial location patterns Cercocarpus ledifolius Nutt. (cutleaf mountain mahogany) plants, classified as current-year seedling, established seedling, juvenile, and immature individuals, at a central Nevada study site. Most current-year seedlings were located in mahogany stands in which large, mature individuals had the greatest abundance. These stands had greater litter cover and a thicker layer of litter than areas with few current-year seedlings. Most established young Cercocarpus were located in adjacent Artemisia tridentata spp. vaseyana (mountain big sagebrush) communities, or in frequent canopy gaps between relatively few large, mature Cercocarpus . We discuss potential roles of plant litter, root growth characteristics, nurse plants, and herbivory in the establishment and renewal of Cercocarpus communities.     

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.     

Long-term fire history in Great Basin sagebrush reconstructed from macroscopic charcoal in spring sediments, Newark Valley, Nevada

We use macroscopic charcoal analysis to reconstruct fire history in sagebrush ( Artemisia tridentata var. wyomingensis and A. tridentata var. tridentate ), in Newark Valley, Nevada. We analyzed charcoal at continuous 1-cm intervals (～7-127 years), and pollen at 2- to 10-cm intervals (～70-263 years) in a core spanning the last 5500 cal yr BP (calendar years before present). A charcoal peak in the historic period was associated with a >1400-ha fire dated to 1986 that burned in the watershed. We reconstructed the prehistoric fire history by inferring fires from similar charcoal peaks that were significantly greater than the background charcoal accumulation. Our results suggest the fire regime is climate and fuel driven. During periods of wetter climate, sagebrush increased and fires were more abundant, and during extended dry periods when sagebrush decreased, fires were less frequent. Our method does not allow calculation of a fire-return interval; however, our results support models that estimate a mean fire-return interval of up to a century in Artemisia tridentata var. wyomingensis . The charcoal record indicates that fires have increased within the historic period. This contrasts with pinyon/juniper studies that indicate an expansion of woodland associated with fewer fires in the historic period. We suggest that in the central Great Basin, a regime of frequent fires in sagebrush that limits woodland expansion is true for the sagebrush-woodland ecotone, but in sagebrush-dominated valleys with lower fuel loads, fires have always been less frequent. Protecting sagebrush-dominated valleys from frequent fire would appear to be consistent with the prehistoric fire regime.