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.     

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.     

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.     

Is Pile Seeding Wyoming Big Sagebrush (Artemisia tridentata subsp. wyomingensis) an Effective Alternative to Broadcast Seeding?

Sagebrush plays an important role in the ecological functions of sagebrush steppe plant communities and is a necessary component of habitat for a variety of wildlife including greater sage-grouse (Centrocercus urophasianus). At lower elevations, increased fire frequency associated with exotic annual grass invasion has heightened the need for effective sagebrush restoration strategies, but existing techniques have been largely ineffective. Our objective was to evaluate “pile seeding” (placing mature seed heads on the ground) of Wyoming big sagebrush (Artemisia tridentata subsp. wyomingensis) as an alternative to broadcast seeding. We used a randomized block design (n = 5) replicated in 2 yr at two contrasting ecological sites in southeastern Oregon. Treatments applied to 100 × 1.5 m plots included 1) pile seeding (four mature seed heads · pile^--1 × 10 piles · plot^-1), 2) broadcast seeding (0.5 kg pure live seed [PLS] · ha^-1), and 3) natural recovery (i.e., nonseeded). Planting occurred in fall 2008 and 2009, and plots were monitored for seedling establishment for three or two growing seasons postplanting. Seedling density was estimated at the plot scale within a 50-cm radius of each seed head pile (“island scale”). In the year following planting, sagebrush seedling density at the plot scale was up to 60-fold higher (P ≤ 0.05) in pile-seeded plots compared to natural recovery and broadcast plots. Seedling mortality was high (up to 98% reduction in density) for pile-seeded plots between the first and second growing seasons postplanting and differences between broadcast and pile-seeded plots dissipated by 2–3 yr postplanting. Although pile-seeding had higher initial density than broadcast seeding, neither technique had sufficient multiyear survival to suggest restoration efficacy at the plot scale. Seedling density at the island scale suggests that pile-seeding may be useful for establishing sagebrush islands, depending on year conditions. Research is needed to determine strategies capable of increasing long-term sagebrush seedling survival.     

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.     

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.     

Factors Affecting Bromus Tectorum Seed Bank Carryover in Western Utah

Cheatgrass (Bromus tectorum L.) is a winter annual weed that presents a serious obstacle to rangeland restoration in the Intermountain West. The objective of this study was to evaluate factors regulating the size and persistence of cheatgrass carryover seed banks on semiarid sites in western Utah. We prevented current-year seed production in each of four habitats, then tallied emerging seedlings over the next 44 yr. Twoyr. Two iterations of the study were conducted during consecutive years. One year before initiation of each iteration, we estimated seed rain at each site. Above-average precipitation in 1998–1999 resulted in relatively high seed rain (13942 seeds·m⁻²) for the first iteration, whereas seed rain for the second iteration averaged only 3567·m⁻² because of drought conditions in 1999–2000. Mean total number of seedlings emerging from carryover seeds for the first and second iterations were 1304 and 270 seedlings·m⁻². Seedling emergence from carryover seed was positively correlated with production-year seed rain (R² = 0.69). The fraction of seed rain that carried over tended to be lower when precipitation the year following production favored fall emergence of the transient seed bank. First-year emergence of carryover seeds averaged 96% of total emergence, whereas third-year emergence averaged < 1% and was zero for six of eight cases. Carryover seeds persisted somewhat longer at the xeric black greasewood site than at more upland sites. Our study shows that cheatgrass seeds rarely persist beyond the second carryover year even on semiarid sites. Emergence from the carryover seed bank can be predicted from site attributes and precipitation patterns in previous years.     

Hydrologic Impacts of Mechanical Seeding Treatments on Sagebrush Rangelands

In and around the Great Basin, United States, restoration of shrub steppe vegetation is needed where rangelands are transitioning to annual grasslands. Mechanical seedbed preparation can aid native species recovery by reducing annual grass competition. This study was designed to investigate the nature and persistence of hydrologic and erosion impacts caused by different mechanical rangeland seeding treatments and to identify interactions between such impacts and related soil and vegetation properties. A cheatgrass (Bromus tectorum L.)–dominated site was burned and seeded with native grasses and shrubs in the fall of the year. An Amazon-drill and a disk-chain seeder were used to provide varying levels of surface soil disturbance. An undisturbed broadcast seeding was used as a control. Simulated rainfall was applied to 6 large (32.5-m²) plots per treatment over 3 growing seasons at a rate of 63.5 mm · h^-1. Rainfall was applied for 60 minutes under dry antecedent moisture conditions and for 30 minutes, 24 hours later under wet antecedent moisture conditions. The disk-chain created the largest reduction in infiltration and increase in sediment yield, which lasted for 3 growing seasons posttreatment. The Amazon-drill had a lesser impact, which was insignificant after the second growing season posttreatment. Surface soil properties showed little correlation with treatment-induced hydrologic and erosion impacts. Hydrologic recovery was strongly correlated with litter dynamics. The seeding treatments were unsuccessful at establishing seeded plant species, and the site once again became dominated by cheatgrass. A continuous upward trend in biomass production and surface litter cover was observed for all treatments between the beginning and end of the study because of cheatgrass invasion. Although the initial goal of using mechanical seeding treatments to enhance recovery of native grass species failed, cheatgrass production provided sufficient biomass to rapidly replenish surface litter cover necessary for rapid hydrologic stability of the site.     

Transition From Sagebrush Steppe to Annual Grass (Bromus tectorum): Influence on Belowground Carbon and Nitrogen

Vegetation changes associated with climate shifts and anthropogenic disturbance have major impacts on biogeochemical cycling. Much of the interior western United States currently is dominated by sagebrush (Artemisia tridentata Nutt.) ecosystems. At low to intermediate elevations, sagebrush ecosystems increasingly are influenced by cheatgrass (Bromus tectorum L.) invasion. Little currently is known about the distribution of belowground organic carbon (OC) on these changing landscapes, how annual grass invasion affects OC pools, or the role that nitrogen (N) plays in carbon (C) retention. As part of a Joint Fire Sciences-funded project called the Sagebrush Treatment Evaluation Project (SageSTEP), we quantified the depth distribution of soil OC and N at seven sites experiencing cheatgrass invasion. We sampled plots that retained sagebrush, but represented a continuum of cheatgrass invasion into the understory. Eighty-four soil cores were taken using a mechanically driven diamond-tipped core drill to a depth of 90 cm, or until bedrock or a restrictive layer was encountered. Samples were taken in 15-cm increments, and soil, rocks, and roots were analyzed for OC and total N. We determined that cheatgrass influences the vertical distribution of OC and N within the soil profile and might result in decreased soil OC content below 60  cm. We also found that OC and total N associated with coarse fragments accounted for at least 10% of belowground pools. This emphasizes the need for researchers to quantify nutrients in deep soil horizons and coarse fragments.     

Prescribed Fire,Soil, and Plants: Burn Effects and Interactions in the Central Great Basin

Pinyon and juniper expansion into sagebrush ecosystems results in decreased cover and biomass of perennial grasses and forbs. We examine the effectiveness of spring prescribed fire on restoration of sagebrush ecosystems by documenting burn effects on soil nutrients, herbaceous aboveground biomass, and tissue nutrient concentrations. This study was conducted in a central Nevada woodland and included control and burn treatment plots sampled before and after a prescribed fire. Six native understory plant species (Crepis acuminata, Eriogonum umbellatum, Eriogonum elatum, Poa secunda secunda, Festuca idahoensis, and Lupinus argenteus) important for native sagebrush obligate foragers were chosen to represent the understory plant community. L. argenteus is also important for system nutrient cycling and nitrogen fixation. Plants were collected from three microsites (under tree canopy, under shrub canopy, and interspace) common in transitional woodlands during peak growth the summer before a spring prescribed burn and each of two summers following the burn. Soils were collected from corresponding locations at two depth intervals (0–8 and 8–52 cm) to determine the relationships between soil and plant nutrients following fire. Microsite affected soil nutrients but did not influence plant tissue concentrations with the exception of F. idahoensis. Burning resulted in increases in soil surface NH+4, NO₃⁻, inorganic N, Ca²⁺, Mn²⁺, and Zn²⁺. Increases in NO₃⁻, inorganic N, and Zn²⁺ were also observed in deeper horizons. Burning did not affect aboveground plant biomass or nutrient concentrations in the first year with the exception of F. idahoensis, which had increased tissue P. By the second year, all species had statistically significant responses to burning. The most common response was for increased aboveground plant weight and tissue N concentrations. Plant response to burning appeared to be related to the burn treatment and the soil variables surface K⁺, NO₃⁻, and inorganic N.     

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 &lsqb;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.     

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.     

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.     

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.     

Vegetation,Hydrologic, and Erosion Responses of Sagebrush Steppe 9 Yr Following Mechanical Tree Removal

Land managers across the western United States are faced with selecting and applying tree-removal treatments on pinyon (Pinus spp.) and juniper (Juniperus spp.) woodland-encroached sagebrush (Artemisia spp.) rangelands, but current understanding of long-term vegetation and hydrological responses of sagebrush sites to tree removal is inadequate for guiding management. This study applied a suite of vegetation and soil measures (0.5 ? 990 m²), small-plot rainfall simulations (0.5 m²), and overland flow experiments (9 m²) to quantify the effects of mechanical tree removal (tree cutting and mastication) on vegetation, runoff, and erosion at two mid- to late-succession woodland-encroached sagebrush sites in the Great Basin, United States, 9 yr after treatment. Low amounts of hillslope-scale shrub (3 ? 15%) and grass (7 ? 12%) canopy cover and extensive intercanopy (area between tree canopies) bare ground (69 ? 88% bare, 75% of area) in untreated areas at both sites facilitated high levels of runoff and sediment from high-intensity (102 mm ? h^? 1, 45 min) rainfall simulations in interspaces (~ 45 mm runoff, 59 ? 381 g ? m^? 2 sediment) between trees and shrubs and from concentrated overland flow experiments (15, 30, and 45 L ? min^? 1, 8 min each) in the intercanopy (371 ? 501 L runoff, 2 342 ? 3 015 g sediment). Tree cutting increased hillslope-scale density of sagebrush by 5% and perennial grass cover by twofold at one site while tree cutting and mastication increased hillslope-scale sagebrush density by 36% and 16%, respectively, and perennial grass cover by threefold at a second more-degraded (initially more sparsely vegetated) site over nine growing seasons. Cover of cheatgrass (Bromus tectorum L.) was < 1% at the sites pretreatment and 1 ? 7% 9 yr after treatment. Bare ground remained high across both sites 9 yr after tree removal and was reduced by treatments solely at the more degraded site. Increases in hillslope-scale vegetation following tree removal had limited impact on runoff and erosion for rainfall simulations and concentrated flow experiments at both sites due to persistent high bare ground. The one exception was reduced runoff and erosion within the cut treatments for intercanopy plots with cut-downed-trees. The cut-downed-trees provided ample litter cover and tree debris at the ground surface to reduce the amount and erosive energy of concentrated overland flow. Trends in hillslope-scale vegetation responses to tree removal in this study demonstrate the effectiveness of mechanical treatments to reestablish sagebrush steppe vegetation without increasing cheatgrass for mid- to late-succession woodland-encroached sites along the warm-dry to cool-moist soil temperature ? moisture threshold in the Great Basin. Our results indicate improved hydrologic function through sagebrush steppe vegetation recruitment after mechanical tree removal on mid- to late-succession woodlands can require more than 9 yr. We anticipate intercanopy runoff and erosion rates will decrease over time at both sites as shrub and grass cover continue to increase, but follow-up tree removal will be needed to prevent pinyon and juniper recolonization. The low intercanopy runoff and erosion measured underneath isolated cut-downed-trees in this study clearly demonstrate that tree debris following mechanical treatments can effectively limit microsite-scale runoff and erosion over time where tree debris settles in good contact with the soil surface.     

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.     

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.     

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.     

Herbicide-Assisted Restoration of Great Basin Sagebrush Steppe Infested With Medusahead and Downy Brome

Downy brome or cheatgrass (Bromus tectorum) and medusahead (Taeniatherum caput-medusae) are the most problematic invasive annual grasses in rangelands of the western United States, including sagebrush communities that provide habitat to sage grouse. Rehabilitation of infested sites requires effective weed control strategies combined with seeding of native plants or desirable competitive species. In this study, we evaluated the effect of three fall-applied pre-emergence herbicides (imazapic, rimsulfuron, and chlorsulfuron + sulfometuron), and one spring-applied postemergence herbicide (glyphosate) on the control of downy brome and medusahead and the response of seeded perennial species and resident vegetation in two sagebrush communities in northeastern California. All pre-emergence treatments gave > 93% control of both invasive species at both sites in the first year. Glyphosate was less consistent, giving > 94% control at one site and only 61% control of both species at the other site. Imazapic was the only herbicide to maintain good control (78–88%) of both species 2 yr after treatment. No herbicide caused detectible long-term damage to either perennial grasses or annual forbs, and imazapic treatment resulted in an increase in resident native forb cover 3 yr after treatment. Broadcast seeding with or without soil incorporation did not result in successful establishment of perennial species, probably due to below-average precipitation in the year of seeding. These results indicate that several chemical options can give short-term control of downy brome and medusahead. Over the course of the study, imazapic provided the best management of both invasive annual grasses while increasing native forb cover.