Aboveground and Belowground Carbon Pools After Fire in Mountain Big Sagebrush Steppe

Postfire succession in mountain big sagebrush (Artemisia tridentata Nutt. subsp. vaseyana [Rydb.] Beetle) ecosystems results in a gradual shift from herbaceous dominance to dominance by shrubs. Determining the quality, quantity, and distribution of carbon (C) in rangelands at all stages of succession provides critical baseline data for improving predictions about how C cycling will change at all stages of succession under altered climate conditions. This study quantified the mass and distribution of above- and belowground (to 1.8-m depth) biomass at four successional stages (2, 6, 20, and 39 yr since fire) in Wyoming to estimate rates of C pool accumulation and to quantify changes in ecosystem carbon to nitrogen (C∶N) ratios of the pools during recovery after fire. We hypothesized that biomass C pools would increase over time after fire and that C∶N ratios would vary more between pools than during succession. Aboveground and live coarse roots (CR) biomass increased to 310 and 17 g C · m^?2, but live fine roots (FR) mass was static at about 225 g C · m^?2. Fine litter (≤ 1-cm diameter) accounted for about 70% of ecosystem C accumulation rate, suggesting that sagebrush leaves decompose slowly and contribute to a substantial soil organic carbon (SOC) pool that did not change during the 40 yr studied. Total ecosystem C (not including SOC) increased 16 g · m^?2 · yr^?1 over 39 yr to a maximum of 1 100 g · m^?2; the fastest accumulation occurred during the first 20 yr. C∶N ratios ranged from 11 for forb leaves to 110 for large sagebrush wood and from 85 for live CR to 12 for bulk soil and were constant across growth stages. These systems may be resilient to grazing after fire because of vigorous regrowth of persistent bunchgrasses and stable pools of live FR and SOC.     

Effects of Nitrogen Availability and Cheatgrass Competition on the Establishment of Vavilov Siberian Wheatgrass

Cheatgrass (Bromus tectorum L.) is the most widespread invasive weed in sagebrush ecosystems of North America. Restoration of perennial vegetation is difficult and land managers have often used introduced bunchgrasses to restore degraded sagebrush communities. Our objective was to evaluate the potential of ‘Vavilov’ Siberian wheatgrass (Agropyron fragile [Roth] P. Candargy) to establish on cheatgrass-dominated sites. We examined Vavilov establishment in response to different levels of soil nitrogen availability by adding sucrose to the soil to promote nitrogen (N) immobilization and examined cheatgrass competition by seeding different levels of cheatgrass. We used a blocked split-split plot design with two sucrose levels (0 and 360 g · m⁻²), two levels of Vavilov (0 and 300 seeds · m⁻²), and five levels of cheatgrass (0, 150, 300, 600, and 1 200 seeds · m⁻²). Seeding was conducted in fall 2003 and 2004, and measurements were taken in June 2004, 2005, and 2006. Sucrose addition decreased availability of soil nitrate but not orthophosphate. In the first year after seeding, sucrose reduced cheatgrass density by 35% and decreased both cheatgrass biomass per square meter and seed production per square meter by 67%. These effects were temporary, and by the second year after seeding, there was a sevenfold increase in cheatgrass density. As a result, the effects of sucrose addition were no longer significant. Sucrose affected Vavilov growth, but not density, during the first year after seeding. Vavilov density decreased as cheatgrass seeding density increased. Short-term reductions in N or cheatgrass seed supply did not have long-term effects on cheatgrass and did not increase Vavilov establishment. Longer-term reductions in soil N, higher seeding densities, or more competitive plant materials are necessary to revegetate areas dominated by cheatgrass.     

Mineral Nitrogen in a Crested Wheatgrass Stand: Implications for Suppression of Cheatgrass

Cheatgrass (Bromus tectorum L.) is an exotic annual grass causing ecosystem degradation in western US rangelands. We investigated potential mechanisms by which crested wheatgrass (Agropyron cristatum L. Gaertn., Agropyron desertorum [Fisch. {Ex Link} Scult.]) suppresses the growth and invasibility of cheatgrass. Research focused on monthly mineral soil N availability and the proportional concentration of NH₄⁺-N in a crested wheatgrass community by microsite (crested wheatgrass, unvegetated interspace, shrub subcanopy) and soil depth (0–15, 15–30 cm) over a 1-yr period. Mineral soil N in crested wheatgrass microsites ranged from 0.24 to 1.66 mmol · kg^-1 and was not appreciably lower than the other microsites or other ecosystems we have measured in the Great Basin. The molar proportion of NH₄⁺-N in the mineral N pool of crested wheatgrass averaged over 85% for the year and is significantly higher than the other microsites and far greater than other plant communities we have measured in the Great Basin. We conclude that crested wheatgrass does not suppress cheatgrass by controlling mineral N below a threshold level; rather, we hypothesize that it may limit nitrification and thereby reduce NO₃^--N availability to the nitrophile cheatgrass.     

Cheatgrass Invasion in Salt Desert Shrublands: Benefits of Postfire Reclamation

In 1998, fires burned more than 11 330 ha of rangeland on Dugway Proving Ground in Utah's west desert. Postfire revegetation was implemented in 2 affected salt desert shrub communities (greasewood; Sarcobatus vermiculatus Hook. and black sagebrush/shadscale; Artemisia nova A. Nels; Atriplex confertifolia Torr. & Frem.) to deter cheatgrass (Bromus tectorum L.) encroachment. We monitored cheatgrass densities for 3 years after the fire in burned drill seeded, burned not-seeded, and unburned plots to assess the rate of invasion and determine the impact on cheatgrass of drill seeding perennial species. Cheatgrass invaded quickly in both shrub sites following the fires. In the greasewood site, drill seeded species germinated but did not establish. This was likely due to a combination of soil salinity and extremely dry weather conditions during the second year of the study. Drill seeded species in the black sagebrush site germinated and established well, resulting in the establishment of 16.5 perennial grasses · m^-2 and 1 356 shrubs · ha^-1. Cheatgrass densities were consistently lower in drill seeded versus not-seeded plots, although these were not always statistically different when Bonferroni comparisons were considered. The initial decrease in cheatgrass densities in drill seeded plots may have resulted from soil disturbance coupled with extremely low precipitation rather than competitive effects. Nevertheless, as seeded species mature and increase their competitive ability, we predict long-term suppression of cheatgrass in the absence of further disturbance.     

Disturbance Type and Sagebrush Community Type Affect Plant Community Structure After Shrub Reduction

Treatments to reduce shrub cover are commonly implemented with the objective of shifting community structure away from shrub dominance and toward shrub and perennial grass codominance. In sagebrush (Artemisia L.) ecosystems, shrub reduction treatments have had variable effects on target shrubs, herbaceous perennials, and non-native annual plants. The factors mediating this variability are not well understood. We used long-term data from Utah’s Watershed Restoration Initiative project to assess short-term (1 − 4 yr post-treatment) and long-term (5 − 12 yr post-treatment) responses of sagebrush plant communities to five shrub reduction treatments at 94 sites that span a range of abiotic conditions and sagebrush community types. Treatments were pipe harrow with one or two passes, aerator, and fire with and without postfire seeding. We analyzed effect sizes (log of response ratio) to assess responses of sagebrush, perennial and annual grasses and forbs, and ground cover to treatments. Most treatments successfully reduced sagebrush cover over the short and long term. All treatments increased long-term perennial grass cover in Wyoming big sagebrush (A. tridentata Nutt. ssp. wyomingensis Beetle & Young) communities, but in mountain big sagebrush (ssp. vaseyana [Rydb.] Beetle) communities, perennial grasses increased only when seeded after fire. In both sagebrush communities, treatments generally resulted in short-term, but not long-term, increases in perennial forb cover. Annual grasses (largely invasive cheatgrass, Bromus tectorum L.) increased in all treatments on sites dominated by mountain big sagebrush but stayed constant or decreased on sites dominated by Wyoming big sagebrush. This result was unexpected because sites dominated by Wyoming big sagebrush are typically thought to be less resilient to disturbance and less resistant to invasion than sites dominated by mountain big sagebrush. Together, these results indicate some of the benefits, risks, and contingent outcomes of sagebrush reduction treatments that should be considered carefully in any future decisions about applying such treatments.     

Prediction of Cheatgrass Field Germination Potential Using Wet Thermal Accumulation

Invasion and dominance of weedy species is facilitated or constrained by environmental and ecological factors that affect resource availability during critical life stages. We compared the relative effects of season, annual weather, site, and disturbance on potential cheatgrass (Bromus tectorum L.) germination in big sagebrush (Artemisia tridentata Nutt.) communities. Soil water status and temperature in the seedbed were measured continuously for 4 years on 9 big sagebrush sites in Nevada and Utah. Field plots at lower-, middle-, and upper-elevation sites were either undisturbed, or were burned, sprayed with herbicide, or both sprayed and burned. Spraying removed perennial herbaceous vegetation, whereas burning removed sagebrush. We used thermal-germination data from laboratory incubation studies of 18 cheatgrass seedlots and field soil moisture and temperature measurements to model and predict potential germination in the field plots for periods when seedbeds were continuously wet (above -0.5, -1, or -1.5 MPa) and across intermittent wet and dry periods. Season had the greatest effect on potential cheatgrass germination, followed by annual weather, and site variables (elevation and location); the effects of disturbance were minimal. Potential germination was predicted for most sites and years in spring, a majority of sites and years in fall, and few sites or years in winter. Even though disturbance has limited effects on potential germination, it can increase cheatgrass invasion and dominance by reducing perennial herbaceous species resource use and allowing increased cheatgrass growth and reproduction.     

Native Plant Growth and Seedling Establishment in Soils Influenced by Bromus Tectorum

The invasion of 40 million hectares of the American West by cheatgrass (Bromus tectorum L.) has caused widespread modifications in the vegetation of semi-arid ecosystems and increased the frequency of fires. In addition to well-understood mechanisms by which cheatgrass gains competitive advantage, it has been implicated in reducing arbuscular mycorrhizal fungi (AMF) abundance and taxa diversity. We evaluated this possibility at a high elevation site in a two-pronged approach. To test whether cheatgrass changed native AMF communities in ways that affected subsequent native plant growth, we grew cheatgrass and native plants in native soils and then planted native plants into these soils in a greenhouse experiment. We found that cheatgrass-influenced soils did not inhibit native plant growth or AMF sporulation or colonization. To test whether soils in cheatgrass-dominated areas inhibited establishment and growth of native plants, cheatgrass was removed and six seeding combinations were applied. We found that 14.02 ±  seedlings · m⁻² established and perennial native plant cover increased fourfold over the three years of this study. Glyphosate reduced cheatgrass cover to less than 5% in the year it was applied but did not facilitate native plant establishment or growth compared with no glyphosate. We conclude that cheatgrass influence on the soil community does not appear to contribute to its invasion success in these high elevation soils. It appears that once cheatgrass is controlled on sites with sufficient native plant abundance, there may be few lingering effects to inhibit the natural reestablishment of native plant communities.     

Can Imazapic Increase Native Species Abundance in Cheatgrass (Bromus tectorum) Invaded Native Plant Communities?

Native plant communities invaded by cheatgrass (Bromus tectorum L.) are at risk of unnatural high intensity fires and conversion to cheatgrass monocultures. Management strategies that reduce cheatgrass abundance may potentially allow native species to expand and minimize further cheatgrass invasion. We tested whether the selective herbicide imazapic is effective in reducing cheatgrass and “releasing” native species in a semiarid grassland and shrub steppe in north-central Oregon. The experiment consisted of a completely randomized design with two treatments (sprayed with 70 g ai · ha^?1 of imazapic and unsprayed) and three replicates of each treatment applied to either 2.5 or 4 ha plots. We repeated this experiment in three different sites dominated by the following native species: 1) bluebunch wheatgrass (Pseudoroegneria spicata [Pursh] A. Löve ssp. spicata) and needle and thread (Hesperostipa comata [Trin. & Rupr.] Barkworth), 2) needle and thread and Sandberg bluegrass (Poa secunda J. Presl), and 3) big sagebrush (Artemisia tridentata Nutt.). Nested frequency of all plant species in 1-m² quadrats was collected for 1  yr pretreatment and 4  yr posttreatment. In all three sites, cheatgrass frequencies were significantly lower in sprayed plots than unsprayed plots for 3–4  yr posttreatment (P < 0.1). Other annual plant species were also impacted by imazapic, but the effects were highly variable by species and site. Only two native perennial species, hoary tansyaster (Machaeranthera canescens [Pursh] Gray) and big sagebrush, increased in sprayed plots, and increases occurred only at two sites. These results suggest that a short-term reduction in cheatgrass alone is not an effective strategy for increasing the abundance of most native perennial plant species.     

Simulating Current Successional Trajectories in Sagebrush Ecosystems With Multiple Disturbances Using a State-and-Transition Modeling Framework

Disturbances and their interactions play major roles in sagebrush (Artemisia spp. L.) community dynamics. Although impacts of some disturbances, most notably fire, have been quantified at the landscape level, some have been ignored and rarely are interactions between disturbances evaluated. We developed conceptual state-and-transition models for each of two broad sagebrush groups—a warm-dry group characterized by Wyoming big sagebrush (Artemisia tridentata Nutt. subsp. wyomingensis Beetle & Young) communities and a cool-moist group characterized by mountain big sagebrush (Artemisia tridentata Nutt. subsp. vaseyana [Rydb.] Beetle) communities. We used the Vegetation Dynamics Development Tool to explore how the abundance of community phases and states in each conceptual model might be affected by fire, insect outbreak, drought, snow mold, voles, sudden drops in winter temperatures (freeze-kill), livestock grazing, juniper (Juniperus occidentalis var. occidentalis Hook.) expansion, nonnative annual grasses such as cheatgrass (Bromus tectorum L.), and vegetation treatments. Changes in fuel continuity and loading resulted in average fire rotations of 12 yr in the warm-dry sagebrush group and 81 yr in the cool-moist sagebrush group. Model results in the warm-dry sagebrush group indicated postfire seeding success alone was not sufficient to limit the area of cheatgrass domination. The frequency of episodes of very high utilization by domestic livestock during severe drought was a key influence on community phase abundance in our models. In the cool-moist sagebrush group, model results indicated at least 10% of the juniper expansion area should be treated annually to keep juniper in check. Regardless, juniper seedlings and saplings would remain abundant.     

Suppression of Cheatgrass by Perennial Bunchgrasses

Long-term control of the invasive annual grass cheatgrass is predicated on its biological suppression. Perennial grasses vary in their suppressive ability. We compared the ability of a non-native grass (“Hycrest” crested wheatgrass) and two native grasses (Snake River wheatgrass and bluebunch wheatgrass) to suppress cheatgrass. In a greenhouse in separate tubs, 5 replicates of each perennial grass were established for 96 d, on which two seeds of cheatgrass, 15 cm apart, were then sown in a semicircular pattern at distances of 10 cm, 30 cm, and 80 cm from the established perennial bunchgrasses. Water was not limiting. After 60 d growth, cheatgrass plants were harvested, dried, weight recorded, and tissue C and N quantified. Soil N availability was quantified at each location where cheatgrass was sown, both before sowing and after harvest. Relative to cheatgrass grown at 80 cm, all perennial grasses significantly reduced aboveground biomass at 30 cm (68% average reduction) and at 10 cm (98% average reduction). Sown at 10 cm from established perennial grasses, cheatgrass aboveground biomass was inversely related with perennial grass root mass per unit volume of soil. All cheatgrass sown at 10 cm from “Hycrest” crested wheatgrass died within 38 d. Before sowing of cheatgrass, soil 10 cm from established perennial grasses had significantly less mineral N than soil taken at 30 cm and 80 cm. Relative to cheatgrass tissue N for plants grown at 80 cm, cheatgrass nearest to the established perennial grasses contained significantly less tissue N. All perennial grasses inhibited the NO₂⁻ to NO₃⁻ nitrification step; for “Hycrest” crested wheatgrass, soil taken at 10 cm from the plant had a molar proportion of NO₂⁻ in the NO₂⁻ + NO₃⁻ pool of > 90%. In summary, a combination of reduced nitrogen availability, occupation of soil space by perennial roots, and attenuation of the nitrogen cycle all contributed to suppression of cheatgrass.     

Soil Carbon Pools in California’s Annual Grassland Ecosystems

Rangeland ecosystems cover approximately one-third of the land area in the United States and half of the land area of California. This large land area, coupled with the propensity of grasses to allocate a considerable proportion of their photosynthate belowground, leads to high soil carbon (C) sequestration potential. Annual grasslands typical of the Mediterranean climates of the western United States differ in their life history strategies from the well-studied perennial grasslands of other regions and thus may also differ in their soil C pools and fluxes. In this study we use the literature to explore patterns in soil C storage in annual grass-dominated rangelands in California. We show that soil C is highly predictable with depth. Cumulative soil C content increased to 2–3-m depth in rangelands with a woody component and to at least 1-m depth in open rangelands. Soil C within a given depth varied widely, with C content in the top 1-m depth spanning almost 200 Mg C · ha^?1 across sites. Soil C pools were not correlated with temperature or precipitation at a regional scale. The presence of woody plants increased C by an average of 40 Mg · ha^?1 in the top meter of soil. Grazed annual grasslands had similar soil C content as ungrazed grassland at all depths examined, although few details on grazing management were available. Soil C pools were weakly positively correlated with clay content and peaked at intermediated levels of aboveground net primary production. Our results suggest that annual grasslands have similar soil C storage capacity as temperate perennial grasslands and offer an important resource for mitigation of greenhouse gas emissions and climate change.     

Prescribed Summer Fire and Seeding Applied to Restore Juniper-Encroached and Exotic Annual Grass-Invaded Sagebrush Steppe

Western juniper (Juniperus occidentalis Hook.) encroachment and exotic annual grass (medusahead [Taeniatherum caput-medusae L. Nevski] and cheatgrass [Bromus tectorum L.]) invasion of sagebrush (Artemisia L.) communities decrease ecosystem services and degrade ecosystem function. Traditionally, these compositional changes were largely confined to separate areas, but more sagebrush communities are now simultaneously being altered by juniper and exotic annual grasses. Few efforts have evaluated attempts to restore these sagebrush communities. The Crooked River National Grassland initiated a project to restore juniper-encroached and annual grass-invaded sagebrush steppe using summer (mid-July) applied prescribed fires and postfire seeding. Treatments were unburned, burned, burned and seeded with a native seed mix, and burned and seeded with an introduced seed mix. Prescribed burning removed all juniper and initially reduced medusahead cover but did not influence cheatgrass cover. Neither the native nor introduced seed mix were successful at increasing large bunchgrass cover, and 6 yr post fire, medusahead cover was greater in burned treatments compared with the unburned treatment. Large bunchgrass cover and biological soil crusts were less in treatments that included burning. Exotic forbs and bulbous bluegrass (Poa bulbosa L.), an exotic grass, were greater in burned treatments compared with the unburned treatment. Sagebrush communities that are both juniper encroached and exotic annual grass invaded will need specific management of both juniper and annual grasses. We suggest that additional treatments, such as pre-emergent herbicide control of annuals and possibly multiple seeding events, are necessary to restore these communities. We recommend an adaptive management approach in which additional treatments are applied on the basis of monitoring data.     

Site,Competition, and Plant Stock Influence Transplant Success of Wyoming Big Sagebrush

Within the sagebrush steppe ecosystem, sagebrush plants influence a number of ecosystem properties, including nutrient distribution, plant species diversity, soil moisture, and temperature, and provide habitat for a wide variety of wildlife species. Recent increases in frequency and size of wildfires and associated annual grass expansion within the Wyoming big sagebrush alliance have increased the need for effective sagebrush restoration tools and protocols. Our objectives were to quay the success of Wyoming big sagebrush transplants relative to transplant stock (nursery seedlings vs. wildlings) across different ecological sites and vegetation types and to test the hypothesis that reduction of herbaceous vegetation would increase survival of transplanted sagebrush. We used a randomized block (reps = 5) design at each of three sites—1) cheatgrass dominated, 2) native plant dominated, and 3) crested wheatgrass dominated—near Elko, Nevada. Treatments included plant stock (nursery stock or locally harvested wildlings) and herbicide (glyphosate) to reduce competition from herbaceous vegetation. Transplants were planted in the spring of 2009 and 2010 and monitored for survival. Data were analyzed for site and treatment effects using mixed-model ANOVA. Surviving plant density at and 2 yr postplanting was generally highest (up to 3-fold) on the native site (P < 0.05). Density of surviving transplants was almost 3-fold higher for nursery stock on most sites for the 2009 planting, but differences in survival by planting stock were minimal for the 2010 planting. Glyphosate application increased surviving plant density up to 300% (depending on site) for both years of planting. High labor and plant material investments (relative to traditional drilling or broadcasting) may limit the size of projects for which sagebrush transplants are practical, but these costs may be partially offset by high success relative to traditional methods. Our data indicate that sagebrush transplants can be effective for establishing sagebrush on depleted sites.     

Resilience and Resistance of Sagebrush Ecosystems: Implications for State and Transition Models and Management Treatments

In sagebrush ecosystems invasion of annual exotics and expansion of piñon (Pinus monophylla Torr. and Frem.) and juniper (Juniperus occidentalis Hook., J. osteosperma &lsqb;Torr.] Little) are altering fire regimes and resulting in large-scale ecosystem transformations. Management treatments aim to increase resilience to disturbance and enhance resistance to invasive species by reducing woody fuels and increasing native perennial herbaceous species. We used Sagebrush Steppe Treatment Evaluation Project data to test predictions on effects of fire vs. mechanical treatments on resilience and resistance for three site types exhibiting cheatgrass (Bromus tectorum L.) invasion and/or piñon and juniper expansion: 1) warm and dry Wyoming big sagebrush (WY shrub); 2) warm and moist Wyoming big sagebrush (WY PJ); and 3) cool and moist mountain big sagebrush (Mtn PJ). Warm and dry (mesic/aridic) WY shrub sites had lower resilience to fire (less shrub recruitment and native perennial herbaceous response) than cooler and moister (frigid/xeric) WY PJ and Mtn PJ sites. Warm (mesic) WY Shrub and WY PJ sites had lower resistance to annual exotics than cool (frigid to cool frigid) Mtn PJ sites. In WY shrub, fire and sagebrush mowing had similar effects on shrub cover and, thus, on perennial native herbaceous and exotic cover. In WY PJ and Mtn PJ, effects were greater for fire than cut-and-leave treatments and with high tree cover in general because most woody vegetation was removed increasing resources for other functional groups. In WY shrub, about 20% pretreatment perennial native herb cover was necessary to prevent increases in exotics after treatment. Cooler and moister WY PJ and especially Mtn PJ were more resistant to annual exotics, but perennial native herb cover was still required for site recovery. We use our results to develop state and transition models that illustrate how resilience and resistance influence vegetation dynamics and management options.     

Invasion is Contingent on Species Assemblage and Invasive Species Identity in Experimental Rehabilitation Plots

Ecological studies often suggest that diverse communities are most resistant to invasion by exotic plants, but relatively few local species may be available to a rehabilitation practitioner. We examine the ability of monocultures and diverse assemblages to resist invasion by an exotic annual grass (cheatgrass) and an exotic biennial forb (dyer's woad) in experimental rehabilitation plots. We constructed seven assemblages that included three monocultures of grass, forb, or shrub; three four-species mixtures of grasses, forbs, or shrubs; and a three-species mixture of one species from each growth form in an experimental field setting to test resistance to invasion. Assemblages were seeded with cheatgrass and dyer's woad for two consecutive years and quantified as biomass and density of individuals from each exotic species. Soil NO₃^- and leaf-area index were examined as predictors of invasive plant abundance. Cheatgrass invasion was greatest in forb and shrub assemblages, and least in mixed grass or grass monoculture; dyer's woad invasion was greatest into mixed grass or grass monoculture, but least into monoculture or mixed-species assemblages composed of forbs or shrubs. The community composed of grasses, forbs, and shrubs suppressed invasion by both species. Consequently, assemblages were most resistant to invasion by species of the same growth form. Moreover, these monocultures and mixtures were generally similar in conferring resistance to invasion, but a monoculture of big sagebrush was more resistant than a mixture of shrubs. Soil NO₃^- was correlated with invasion by cheatgrass, whereas LAI was correlated with invasion by dyer's woad, suggesting these species were more limited by belowground and aboveground resources, respectively. Overall, increasing diversity with limited species did not necessarily enhance resistance to invasion.     

Effects of Sagebrush Restoration and Conifer Encroachment on Small Mammal Diversity in Sagebrush Ecosystem

Conifer encroachment in sagebrush ecosystems reduces habitat heterogeneity, niche space, and resource availability, all of which negatively affect many wildlife populations. Sagebrush restoration is recommended as a management action to mitigate conifer encroachment and restore wildlife across millions of hectares in the Great Basin. Despite this recommendation, the effects of conifer encroachment and sagebrush restoration are unknown for most wildlife species. Small nonvolant mammal communities include keystone species, consumers and prey; facilitate energy flow and ecological function; and provide important ecological goods and services. We assessed causal relationships between conifer encroachment and sagebrush restoration (conifer removal and seeding native plants) on small mammal communities over 11 yr using a Before-After-Control–Impact design. Sagebrush habitat supported an additional small mammal species, twice the biomass, and nearly three times higher densities than conifer-encroached habitat. Sagebrush restoration increased shrub cover, decreased tree cover, and density but failed to increase native herbaceous plant density. Restoration caused a large increase in the non-native, invasive annual cheatgrass (Bromus tectorum L.). Counter to prediction, small mammal diversity did not increase in response to sagebrush restoration, but restoration maintained small mammal density in the face of ongoing conifer encroachment. Piñon mice (Peromyscus truei), woodland specialists with highest densities in conifer-encroached habitat, were negatively affected by sagebrush restoration. Increasing cheatgrass due to sagebrush restoration may not negatively impact small mammal diversity, provided cheatgrass density and cover do not progress to a monoculture and native vegetation is maintained. The consequences of conifer encroachment, a long-term, slow-acting impact, far outweigh the impacts of sagebrush restoration, a short-term, high-intensity impact, on small mammal diversity. Given the ecological importance of small mammals, maintenance of small mammal density is a desirable outcome for sagebrush restoration.     

Plant Establishment in Masticated Utah Juniper Woodlands

Juniper (Juniperus spp.) encroachment into sagebrush (Artemisia spp.)-bunchgrass communities has reduced understory cover on millions of hectares of semiarid rangelands. Mechanical masticators shred trees to restore desirable vegetation and reduce the potential for catastrophic wildfire. Mechanical mastication where juniper density is high and perennial grass cover is low brings a risk of invasive weed dominance unless perennial species are established. To determine whether juniper mastication favors annual- or perennial-grass establishment, we compared seedling emergence, tillers, and aboveground biomass of cheatgrass (Bromus tectorum L.) and Anatone bluebunch wheatgrass (Pseudoroegneria spicata [Pursh] A. Löve). Comparisons were made among hand-planted rows between and under juniper canopies of masticated and adjacent untreated control areas at three locations in Utah. Bluebunch wheatgrass had 16% (95% CI: 11–21) and cheatgrass had 10% (95% CI: 5–15) fewer seedlings emerge per row in masticated than untreated areas (P < 0.001). However, bluebunch wheatgrass had 3.2 (95% CI: 2.0–5.2) times more tillers and 1.9 (95% CI: 1.6–2.2) times more aboveground biomass per row in masticated than untreated areas (P < 0.001). Similarly, cheatgrass had 2.3 (95% CI: 1.5–3.8) times more tillers, 2.0 (95% CI: 1.7–2.4) times more aboveground biomass, and 11.4 (95% CI: 6.3–20.7) times more spikelets per row in masticated than untreated areas (P < 0.001). This increased seedling growth in masticated areas was associated with increased inorganic nitrogen and soil water compared to untreated areas. Because mastication improves the growth of both cheatgrass and bluebunch wheatgrass seedlings, it could support dominance by either annual- or perennial-life forms. To avoid cheatgrass dominance where perennial understory cover is limited and cheatgrass propagule pressure is high, mastication should be accompanied by seeding desirable perennial species such as Anatone bluebunch wheatgrass.     

Defoliation Effects On Bromus Tectorum Seed Production: Implications For Grazing

Cheatgrass (Bromus tectorum L.) is an invasive annual grass that creates near-homogenous stands in areas throughout the Intermountain sagebrush steppe and challenges successful native plant restoration in these areas. A clipping experiment carried out at two cheatgrass-dominated sites in eastern Oregon (Lincoln Bench and Succor Creek) evaluated defoliation as a potential control method for cheatgrass and a seeding preparation method for native plant reseeding projects. Treatments involved clipping plants at two heights (tall = 7.6 cm, and short = 2.5 cm), two phenological stages (boot and purple), and two frequencies (once and twice), although purple-stage treatments were clipped only once. Treatments at each site were replicated in a randomized complete block design that included a control with no defoliation. End-of-season seed density (seeds · m⁻²) was estimated by sampling viable seeds from plants, litter, and soil of each treatment. Unclipped control plants produced an average of approximately 13 000 and 20 000 seeds · m⁻² at Lincoln Bench and Succor Creek, respectively. Plants clipped short at the boot stage and again 2 wk later had among the lowest mean seed densities at both sites, and were considered the most successful treatments (Lincoln Bench: F_6,45 = 47.07, P < 0.0001; Succor Creek: F_6,40 = 19.60, P < 0.0001). The 95% confidence intervals for seed densities were 123–324 seeds · m⁻² from the Lincoln Bench treatment, and 769–2 256 seeds · m⁻² from the Succor Creek treatment. Literature suggests a maximum acceptable cheatgrass seed density of approximately 330 seeds · m⁻² for successful native plant restoration through reseeding. Thus, although this study helped pinpoint optimal defoliation parameters for cheatgrass control, it also called into question the potential for livestock grazing to be an effective seed-bed preparation technique in native plant reseeding projects in cheatgrass-dominated areas.     

Differential Species Responses to Aspects of Resistance to Invasion in Two Columbia Plateau − Protected Areas

Protected-area sagebrush steppe ecosystems are few in number and increasingly important to the North American conservation network as sagebrush steppe faces growing threats from land use, climate change, and invasive species. We analyzed the distribution and abundance of native perennial and invasive annual plants to better understand patterns of plant invasion within two protected areas: John Day Fossil Beds National Monument (JODA), located in central Oregon, and Craters of the Moon National Monument and Preserve (CRMO), located in southeast Idaho. We used multivariate analysis to examine vegetation monitoring datasets and illuminate geographic variation in plant cover along gradients of well-known aspects of resistance to plant invasion (elevation, exposure [slope and aspect], precipitation and proximity to disturbance). Topographically mediated resistance to invasion appeared to manifest in the park with greater topographic variability (JODA), while increased elevation was more strongly associated with resistant sites in the park, which spanned a greater elevational gradient (CRMO). Factors that may mitigate moisture-mediated resistance also differed between sites. Slope and aspect were factors of apparent resistance for bunchgrass communities in JODA, while high crop year precipitation appeared to benefit medusahead (Taeniatherum caput-medusae [L.] Nevski) and the weedy native subshrub broom snakeweed (Gutierrezia sarothrae [Pursh] Britton & Rusby) over bunchgrasses and Wyoming big sagebrush (Artemisia tridentata Nutt. ssp. wyomingensis Beetle & Young). Increased elevation and distance to disturbed areas were the most important factors of resistance in forb-rich communities at CRMO, with the invasive annuals cheatgrass (Bromus tectorum L.), tumblemustard (Sisymbrium altissimum L.), and Descurainia spp. Webb & Bethel. invading in low elevations and in close proximity to roads or agricultural fields. Such complexity underscores the idiosyncratic nature of the manifestation of resistance and the need for place-based empirical studies to provide information for guiding protected-area management.