Long-term vegetation dynamics in a cut western juniper woodland

Western juniper ( Juniperus occidentalis spp. occidentalis Hook.) expansion in the northern Great Basin has reduced shrubsteppe productivity and diversity. Chainsaw cutting of western juniper woodlands is commonly applied to remove tree interference and restore sagebrush plant communities. Studies assessing understory response following cutting have been limited to early successional stages and have not evaluated the effects of western juniper debris on plant succession. Cutting western juniper produces a large amount of debris which is commonly left on site, occupying a significant portion of treated areas. This study evaluated successional dynamics spanning 13 years after western juniper cutting. Four 0.45-ha blocks were selected on Steens Mountain in southeastern Oregon. Western juniper cover averaged 26% and mature tree density averaged 250 trees ? ha –1 . Blocks were cut in late summer 1991. Understory standing crop, cover, and density were compared among 3 locations: old canopy litter mats (canopy), interspace, and area underneath cut western juniper (debris). In the interspace, perennial grasses increased in cover and in standing crop relative to other functional groups. In canopy and debris locations, species composition shifted in the 6th year after cutting as annual grass cover, density, and standing crop increased. However, by 2003, perennial grass biomass was 2 times greater than annual grass biomass in canopy and debris locations. Because annual grasses increased in areas of debris accumulation, managers need to be cognizant of western juniper treatments that create safe sites that are favorable to the establishment of weedy species. Retaining western juniper debris on this site did not increase establishment and growth of perennial grasses compared to the interspace. A shift in perennial grass dominance from Thurber's needlegrass ( Achnatherum thurberianum [Piper] Barkworth) to bottlebrush squirreltail ( Elymus hystrix [Nutt.] Smith) occurred in areas of debris accumulation. Our results demonstrated that long-term vegetation evaluations are necessary to properly assess management activities and disturbance.     

Pinyon-juniper woodlands in Zion National Park, Utah

Juniperus osteosperma-Pinus monophylla or P. edulis (P-J) woodlands are the most widespread plant community in Zion National Park (ZNP), southwestern Utah. These woodlands dominate nearly half of the park's land area. Our study of this vegetational complex is based on a sample consisting of 115 macroplots (each 0.01 ha in area) objectively distributed across the entire area of ZNP. We recognize 3 subtypes within the P-J complex: Juniperus osteosperma (Utah juniper) alone, juniper with P. monophylla (single-leaf pinyon), and juniper with P. edulis (two-leaf pinyon). The 2 pinyon pines rarely occur together, and thus the foregoing subtypes do not overlap geographically to a significant extent. The first 2 subtypes occur primarily below 1800 m elevation, while the latter is most commonly found above that elevation. Because of the scarcity of sizable expanses (over ～ 10 ha) of well-developed soils in ZNP, the P-J complex occurs primarily on sites where exposed bedrock covers a large portion of the habitat. As a result, over 90% of stands assigned to the P-J complex support less than 50% tree canopy cover (64% have less than 25% tree cover). Shrub cover increases along the woodland successional gradient. Pinyon cover increases faster than juniper cover. Microbiotic soil crust cover is consistently greater in J. osteosperma-P. monophylla woodlands than in J. osteosperma-P. edulis woodlands, but total living cover increases significantly along the successional gradient in both communities. To enhance plant and animal biodiversity, we recommend that pinyon-juniper woodlands of Zion National Park be managed so that late seral stages do not dominate large tracts.     

Understory patterns in cut western juniper ( Juniperus occidentalis spp. occidentalis Hook.) woodlands

Western juniper ( Juniperus occidentalis spp. occidentalis ) has rapidly expanded into shrub steppe communities in the intermountain Northwest during the past 120 yr. Cutting juniper is a management tool used to restore shrub steppe communities. Response of the understory after cutting is strongly influenced by plant species composition existing prior to treatment. This study assessed distribution patterns of understory plants over 2 growing seasons after tree cutting in a western juniper woodland. Cover, density, and diversity of understory species were compared among 3 locations: interspaces, duff zones (previously under tree canopies), and debris zones (beneath cut trees). Plant cover density increased in all zones following tree cutting. Understory vegetation in cut woodlands exhibited strong zonal distribution. Cover and density of Poa sandbergii and Sitanion hystrix and canopy cover of annual forbs were greatest in duff zones ( P P < 0.05). Debris zones tended to have the lowest overall understory cover and plant density values. Under juniper debris many species common to interspaces were reduced in density, although plants that survived or established beneath debris grew larger than their counterparts in interspaces. Species increased in density and cover under debris were plants characteristic of duff zones and whose seeds are typically wind dispersed.     

Tree size and understory phytomass production in a western juniper woodland

Understory phytomass production in a western juniper ( Juniperus occidentalis ) woodland was examined relative to tree size in central Oregon in 1983 and 1984. Vegetation was sampled in two zones, the canopy zone (beneath the canopy) and the intercanopy zone (the space between canopies), on two adjacent sites—a lower slope site with shallow soil and an upper slope site with deeper soil. Sampling was stratified into three tree size classes. Individual species production was significantly affected by tree size and location relative to tree canopy. Production of bottlebrush squirreltail, bluebunch wheatgrass, cheatgrass, miscellaneous annual grasses, perennial forbs, and annual forbs increased with increasing tree size. Sandberg bluegrass production was greater in the intercanopy than the canopy zone, while production of bottlebrush squirreltail, bluebunch wheatgrass, miscellaneous annual grasses, and both perennial and annual forbs was greater in the canopy zone. Production of cheatgrass was determined by the interaction of tree size and zone. Phytomass relationships were expressed to a greater degree on the upper slope site, where total production exceeded that of the lower slope site by approximately 50% the second year of the study. Individual trees appear to exert a great influence on associated vegetation as western juniper woodlands progress from the seedling (tree establishment) phase to closed stands of mature trees. Original community dominants appear to be spatially segregated beneath tree canopies and associated with large trees, while formerly less common species, such as cheatgrass, come to dominate the entire site.     

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.     

Understory response to tree harvesting of singleleaf pinyon and Utah juniper

Fifteen widely separated sites within the pinyon - juniper woodlands of the Great Basin were cleared of trees. Understory response was recorded for 2 to 4 years. The array of postharvest plant assemblages were classified into one of four phytosociological groups using discriminant analysis. Pre - and postharvest plant assemblages from the same site appeared in the same phytosociological group, which indicates postharvest response could be predicted from the preharvest plant assemblage. Initial postharvest response appears cyclic in nature, and cycles are controlled by both residual plants and the rapid immigration of shrub species. Perennial plant density generally declined following tree harvest, but the fewer remaining plants produced significantly more cover than in preharvest stands. &nbsp;&nbsp;&nbsp;     

Plant community patterns in unburned and burned blackbrush ( Coleogne ramosissima Torr.) shrublands in the Mojave Desert

The blackbrush vegetation type is dominated by Coleogyne ramossisima , which is thought to preclude the coexistence of many other plant species. Fire can remove blackbrush cover and possibly increase plant species richness and evenness. Fire also may increase the frequency and cover of alien annual grasses, thereby intensifying landscape flammability. We tested these predictions in unburned and burned (6-14 years postfire) blackbrush at 3 sites spanning the range of this vegetation type in the Mojave Desert. Species richness in unburned blackbrush was similar to published values for vegetation types in western North America, bur richness varied significantly among the 3 sites and 4 spatial scales (1, 10, 100, and 1000 m 2 ). Richness values declined in order from annual forbs, woody perennials, herbaceous perennials, annual grasses, cacti, to perennial grasses. Fire reduced Coleogyne cover, thus boosting species evenness. In contrast, species richness decreased after burning, although the results varied among spatial scales. Total cover was unaffected by fire because cover of wood perennials decreased, while cover of annual forbs, annual grasses, herbaceous perennials, and perennial grasses increased. Native species richness and cover decreased, whereas alien richness and cover increased after burning especially where the alien forb Erodium cicutarium was present. Fire had no effect on frequency and variable effects on cover of alien annual grasses. These results indicate that in blackbrush species richness can vary among sites and local spatial scales, and effects of fires can vary among plant life-forms and between natives and aliens.     

Relationship between environmental and vegetational parameters for understory and open-area communities

Ten individuals from each of four tree species were selected, and their associated understory and adjacent open-area communities were sampled for both environmental and vegetational parameters, including light intensity, pH, litter depth, soil depth, and percentages of exposed rock, litter cover, living cover, shrubs, forbs, grasses, and annuals. The four tree species were ponderosa pine, Rocky Mountain juniper, Gambel oak, and snowbrush ceanothus. The study site was in the lower Uinta Mountains about 10 miles east of Kamas, Utah. Correlations among the various biotic and abiotic parameters were examined. The interplay of these factors in differentiating the understory and open-area communities is discussed. &nbsp;     

Habitat of three rare species of small mammals in juniper woodlands of southwestern Wyoming

Southwestern Wyoming constitutes the northern limit of the ranges of the cliff chipmunk ( Tamias dorsalis ), pinyon mouse ( Peromyscus truei ), and canyon mouse ( P. crinitus ). In addition to trying to determine their presence in the region, we wanted to identify habitat characteristics commonly used by each of these species. We used Sherman live-traps to sample 14 sites representing 2 distinct habitat types in 1998 and 1999: juniper-rocky slopes and juniper cliffs. Seventeen habitat characteristics were measured at capture locations for each species and compared with randomly located points. Best subsets multiple logistic regression was used to construct models that distinguish between used and available habitat for each species. The cliff chipmunk occurred in both rocky slopes and cliffs. The pinyon mouse was also captured in rocky slopes and cliffs and was most often captured in locations in the interior of the juniper woodland with high tree canopy cover, high forb cover, and low density of rock outcrops. The canyon mouse was captured only in cliffs at sites consisting of high forb cover, high rock cover, and high tree density.     

Understory seed rain in harvested pinyon-juniper woodlands

Seed rain was collected on six paired tree harvest and undisturbed plots in singleleaf pinyon ( Pinus monophylla )&ndash;Utah juniper ( Juniperus osteosperma ) stands. Approximately 14,600 seeds were collected during four years. Seed rain in undisturbed plots was similar to levels in mixed forest communities. Seed rain on harvest plots was similar to disturbed sites and grasslands. Seed rain levels reflect the current successional stage rather than the climax community type for the site. Seed rain increased in numbers and seed production per unit of plant cover following tree removal and especially on transition soil microsites. Only three to four of the plant species present on a site contributed greater than 10% of the total seed rain. Seed rain composition was similar on harvest and undisturbed plots (Jaccard Similarity Index Values = 47% to 67%) and explains in part the rapid reestablishment of predisturbance understory communities.     

Late Pleistocene-early Holocene history of coniferous woodlands in the Lucerne Valley region, Mohave Desert, California

The composition of four radiocarbon-dated, late Pleistocene woodrat middens is reported and analyzed. A date of 11,850 &plusmn;550 BP records the first reported macrofossil occurrence in this region of late Pleistocene Pinus monophylla-Juniperus osteosperma woodlands. A 7,800 &plusmn;350 BP date documents the most recent J. osteosperma woodlands in this presently coniferless desert area, while a date of 12,100 &plusmn;400 BP is the oldest record of juniper woodlands among the four middens. Other juniper and creosote bush desert flora radiocarbon dates along with six pollen profiles were obtained. The research suggests that as recently as 7,800 BP this part of the Mohave Desert was subject to a cooler, moister climate than at present, and that the aboriginal food resources of pinyon seeds and juniper berries were probably available to early prehistoric man in this area.     

Late quaternary plant zonation and climate in southeastern Utah

Plant macrofossils from packrat middens in two southeastern Utah caves outline development of modern plant zonation from the late Wisconsin. Allen Canyon Cave (2195 m) and Fishmouth Cave (1585 m) are located along a continuous gradient of outcropping Navajo Sandstone that extends from the Abajo Mountains south to the San Juan River. By holding the site constant, changes in the floral composition for a plot of less than one hectare can be observed, even if sporadically, over tens of millennia. At Allen Canyon Cave, Engelmann spruce-alpine fir forest was replaced by the present vegetation consisting of pinyon-juniper woodland on exposed ridgetops and cliffside stands of Douglas fir, ponderosa pine, and aspen. Xerophytic woodland plants such as pinyon, plains prickly pear, and narrowleaf yucca arrived sometime in the middle Holocene between 7200 and 3400 B.P. At Fishmouth Cave, Utah juniper in Holocene middens replaced blue spruce, limber pine, Douglas fir, and dwarf and Rocky Mountain junipers in late Wisconsin samples. Disharmonious associations for the late Wisconsin occur only at the lower site with the xerophytes Mormon tea, plains prickly pear, and narrowleaf yucca growing alongside subalpine conifers. One possible explanation involves the late Wisconsin absence of ponderosa and pinyon pines from the Colorado Plateaus. Released from competition at their lower limits, subalpine conifers were able to expand into lower elevations and mix with xerophytic plants found today in understories of pinyon-juniper and ponderosa pine woodlands. Quantitative climatic estimates are derived for the late Wisconsin by applying vertical gradients for temperature and precipitation to the amount of vegetation depression. The Fishmouth Cave sequence indicates a minimum lowering of 850 m for blue spruce, limber pine, and dwarf juniper. A depression of at least 700 m for Engelmann spruce and alpine fir is suggested for the Allen Canyon locality. Use of conservatively low gradients for stations below 2080 m yields a 3&ndash;4 C cooling from present mean annual temperature and 35 to 60 percent more rainfall than today. Steeper gradients associated with more mountainous terrain suggest a 5 C lowering in temperature and up to 120 percent increase over modern precipitation. &nbsp;&nbsp;     

Herbaceous succession after burning of cut western juniper trees

The expansion of western juniper ( Juniperus occidentalis spp. occidentalis Hook.) in the northern Great Basin has resulted in the wide-scale conversion of sagebrush-steppe communities to juniper woodlands. Prescribed fire and mechanical cutting are the 2 main methods used to remove juniper and restore sagebrush steppe. Mechanical treatments commonly leave cut juniper on site. Disadvantages of leaving cut juniper are the increased fuel hazard and the potential for increased establishment and growth of invasive species. This study evaluated the response of herbaceous plants to winter burning of cut western juniper. Vegetation response was compared among 2 burning treatments (burning trees the first winter after cutting and burning the second winter after cutting), a control (cut-unburned juniper), and the interspace between cut trees. To minimize fire impacts to herbaceous perennials, cut trees were burned in the winter when soils and ground litter were frozen and/or soils were at field capacity. Only felled trees were burned, as fire did not carry into interspaces or litter mats around western juniper stumps. We hypothesized that winter season burning would increase herbaceous perennials and would reduce cheatgrass establishment when compared to the cut-unburned control. After 10 years, total herbaceous and perennial grass cover was 1.5- to 2-fold greater, respectively, in burned treatments compared to cut-unburned controls. Perennial grass density was 60% greater in the burned treatments than in the cut-unburned treatment and the interspace. Cheatgrass cover was twice as great in the control than in the 2 burn treatments and the interspace. We concluded that burning cut western juniper when soils were wet and frozen in winter enhanced community recovery of native perennials compared to leaving cut juniper unburned.     

Differential effects of cattle and sheep grazing on high mountain meadows in the Strawberry Valley of central Utah

Species diversity, niche metrics, cover, frequency, and soil relationships were studied on high mountain meadows on adjacent cattle and sheep allotments in Strawberry Valley, Wasatch County, Utah. The cattle allotment vegetation was predominantly Mountain bluebell (Mertensia ciliata) , and the sheep allotment vegetation was predominantly Smallwing sedge (Carex microptera) . Other species of importance on both areas included Letterman needlegrass (Stipa lettermanii) , Mountain bromegrass (Bromus carinatus) , and Yarrow (Achillea millefolium) . Tall forbs were most abundant on the cattle allotment, and low forbs, perennial grasses, and sedges were most abundant on the sheep allotment. Vegetation composition on the two allotments was significantly different. &nbsp;&nbsp;&nbsp;     

Estimates of the incidence of mistletoes in pinyon-juniper woodlands of the Coconino National Forest, Arizona

The relative incidence of mistletoes found in pinyon-juniper woodlands is estimated for the Coconino National Forest, Arizona, using a roadside survey. Approximately 50% of the pinyon-juniper woodlands surveyed were infested with juniper mistletoes ( Phoradendron juniperinum or P. capitellatum ). In contrast, only about 12% of the areas surveyed were infested with pinyon dwarf mistletoe ( Arceuthobium divaricatum ). In these infested woodlands, 70% with juniper mistletoes and 44% with pinyon dwarf mistletoe were lightly infested. Most areas where pinyon dwarf mistletoe populations were observed (90%) also had juniper mistletoe ( P. juniperinum ) present, but many juniper mistletoe populations observed had no pinyon dwarf mistletoe (50%), although pinyon pine was common at those locations. Therefore, pinyon dwarf mistletoe is frequently co-distributed with juniper mistletoe as has been suggested by other investigators, but juniper mistletoe frequently occurs where there is no pinyon dwarf mistletoe present.     

Livestock and vegetation responses to rotational grazing in short-grass steppe

There is a paucity of information concerning livestock and vegetation responses to rotational grazing in short-grass steppe. We compared effects between 1995 and 2003 of time-controlled, short-duration rotational grazing against season-long continuous grazing, at a moderate stocking rate (1.95 ha ? animal unit month –1 ), on livestock gains and on foliar and basal cover in short-grass steppe. Livestock average daily gains, grazing-season gains, and beef production did not differ between grazing systems. We observed no relationship between precipitation (annual or growing-season) and average daily gain. In contrast, both grazing-season gains and beef production exhibited a significant curvilinear response to both growing-season and annual precipitation. Basal and foliar cover of all plant functional groups (C 3 annual grasses, C 3 perennial grasses and grass-likes, C 4 perennial grasses, cactus, annual forbs, perennial forbs, and shrubs/subshrubs) did not differ between grazing systems. Litter and bare ground were also unaffected by grazing system, with litter increasing and bare ground decreasing over the duration of this experiment. Land managers in shortgrass steppe who are implementing rotational grazing at moderate stocking rates should not expect increased livestock gains or changes in basal and foliar cover of the plant community. Rotational grazing may be a useful management tool for achieving conservation goals through modifications of seasonality and intensity of use in paddocks within larger management units to accommodate spatial variability in vegetation and temporal variation in precipitation.     

Tree densities on pinyon-juniper woodland sites in Nevada and California

The densities of sing le leaf pinyon and Utah juniper trees in four diameter classes (1&ndash;9, 10&ndash;19, 20&ndash;29, and &ge; 30 cm) were measured on 522 plots of 1/10 ha each throughout the Great Basin. Density distribution patterns of pinyon and juniper varied with aspect, elevation, and eastern (EGB) versus western Great Basin (WGB) locations. On most locations north and, to a lesser extent, west slopes supported higher densities of pinyon than south and east slopes, with high relative densities of small diameter trees on north slopes and large diameter trees on west slopes. Pinyon densities were higher on EGB than on WGB sites and on higher elevation than on lower elevation sites. Juniper densities were higher on EGB than on WGB sites and on lower elevation than on higher elevation sites. Juniper densities on low-elevation WGB sites were higher on south and west aspects than on north and east, with higher relative densities in the 20&ndash;29 cm diameter class than in other diameter classes. On low elevation EGB sites, east and south slopes supported higher juniper densities than did north and west slopes, with comparatively higher relative densities in the 10&ndash;19 cm diameter class. Differences in relative densities between diameter classes were not significant among aspects on high elevation sites.     

Lagomorphs and the dispersal of seeds into communities dominated by exotic annual weeds

Large areas of western rangeland are presently dominated by alien annual weeds such as Bromus tectorum (cheatgrass). These communities resist succession to perennial communities primarily because the annuals are competitively superior to establishing perennial seedlings and they promote fires that favor weeds over perennials. Succession may be further slowed, however, by low rates of seed dispersal into annual grasslands. We investigated the role of lagomorphs ( Sylvilagus nuttalli , Nuttall's cottontail; S. audubonii , desert cottontail; and Lepus californicus , black-tailed jackrabbit) in seed dispersal across an ecotone: 50 m into woodland, border, and 20 m, 50 m, and 100 m into grassland. We searched pellets for juniper seeds visually and for any other species through germination from crushed pellets after cold, moist stratification. Pellets were not even distributed across transects, but there was no trend with respect to position of transect. Juniperus osteosperma (Utah juniper) was the most abundant seed. Both the number of juniper seeds and the proportion of pellets with juniper seeds decreased steadily from a high in woodland to absence at 100 m into grassland. Only 2 dicot seedlings emerged from pellets, 1 Salsola pestifer and 1 unknown that died prior to identification. Consequently, there was little seed movement into grassland; 72% of all seeds were collected from either woodland or border transects. Lagomorphs apparently do not effectively replenish the native perennial seed pool of cheatgrass-dominated disturbances at Dugway.     

Winter and early spring bird communites in grasslands, shrubsteppe, and juniper woodlands in central Oregon

We compared winter (December, January, and February) and early spring (March and April) bird communities among 4 successional stages that included grassland, shrubsteppe, juniper-shrubsteppe, and old-growth juniper woodland in central Oregon. Birds were surveyed monthly from December through April in 1998/1999 and 1999/2000 using the point count method to estimate relative abundance of birds (individuals ? transect –1 year –1 ). We used distance sampling to correct for potential bias in estimating abundance and density among successional stages. A total of 4513 birds (32 species) were detected. Relative abundance of total birds (all species combined) was similar in the juniper-shrubsteppe and old-growth woodland (48.7 and 48.9 individuals ? transect –1 , respectively) but 6 times greater than in the grassland and shrubsteppe (8.0 and 7.7 individuals ? transect –1 , respectively) during the 5-month period. Total density of birds was greater in old-growth juniper woodland and juniper-shrubsteppe than in grassland or shrubsteppe. However, median bird species richness in 1998/1999 was highest in grassland and lowest both in shrubsteppe and juniper-shrubsteppe, and in 1999/2000 it was highest in shrubsteppe and lowest in juniper-shrubsteppe and oldgrowth juniper woodland. American Robins and Townsend's Solitaires were the most abundant species in junipershrubsteppe and old-growth juniper woodland during the winter months. Sage Sparrows and Horned Larks were the most abundant species in shrubsteppe during winter, and Horned Larks were most abundant in grasslands during the early spring transition period prior to nesting. Our results indicate that a different suite of species use these successional stages during the nonbreeding season. If avifauna conservation is a part of long-term management goals, a broad range of successional stages should be maintained on the landscape to provide habitat for a variety of avian species throughout the year.     

Foliage biomass and cover relationships between tree- and shrub-dominated communities in pinyon-juniper woodlands

Woodlands dominated by singleleaf pinyon ( Pinus monophylla Torr.and Frem.) and Utah ( Juniperus osteosperma [Torr.] Little) cover extensive areas in the Great Basin and Southwest. Both species are aggressive and can nearly eliminate the previous shrub-dominated community. Successional pathways from shrub-dominated communities before tree establishment to the tree-dominated communities that follow are known only for a few specific sites. How site growing conditions affect successional patterns needs further study. We compared the relationship of foliage biomass and percentage of cover between paired shrub-dominated and tree-dominated plots over several sites. Sites studied are from different elevation and topographic conditions on one mountain range. Foliage biomass in shrub-dominated plots had about a three-to-one variation over the range of site conditions sampled. Tree-dominated plots varied by about two-to-one. Cover in shrub-dominated plots had a four-to-one variation; cover in the tree-dominated plots varied by about two-to-one. Total foliage biomass in both tree- and shrub-dominated plots correlated best with the site index of height at 200 years of age. Variation in percentage of cover in both tree and shrub-dominated plots correlated best with elevation. Foliage biomass variation in shrub-dominated plots was proportional to the variation in the paired tree-dominated plots. A similar proportional relationship was present for percentage of cover between paired tree- and shrub-dominated plots. Foliage biomass was more sensitive to topographic differences than to cover. Variation in plant species sampled in the shrub-dominated plots correlated with total foliage biomass of the same plots. Species sampled also correlated with pinyon height at 200 years of age and total foliage biomass in the paired tree-dominated plots.