Biological invasions are driving environmental state changes on a global scale. Exotic plant species must be successful at passing several abiotic and biotic filters to establish and disrupt the native plant community assembly. Understanding where exotic plants are on a regional scale and being able to characterize how exotic plants are generally interacting with their environment is crucial information for exotic species management (chapter 1). In the western United States human-related activities are augmenting the spread of exotic plant species by increasing the ignitions of wildfire. Wildfire can lead to nutrient pulses through the removal of intact native communities and returning some mineral content into the soil. Exotic plant species that have traits that efficiently acquire nutrients accompanied by rapid growth rates may outcompete native plants. In chapters 2, 3, and 4 experimental fires demonstrated that the direct effect of fire may not be as critical as the potential indirect effects of fire such as altering the behavior of consumers (chapter 2) and reducing competition (chapters 3 and 4). In the Mojave desert, rodent consumers can have strong top-down effects on plant community assembly through foraging selection preferences. Life history traits such as seedling and seed size can lead to differential herbivory and positively benefit some plant species while inhibiting others (chapter 1) which could indirectly alter plant-plant interactions. Plant competition is a biotic filter than can determine establishment success or failure. Species that with rapid growth rates and plastic growth responses are likely to be able to capitalize on fluctuations in available resources. In the Great Basin, forecasts in climate change models predict that precipitation timing will lead to heavier fall rains and more rain than snow in the winter. Water availability is one of the main limiting factors in semi-arid and arid ecosystems where native plants have adaptive traits to maximize resource use. The interaction of wildfire and changes in climate, specifically timing of precipitation is critical to understand to be able to predict and protect against increasing wildfire frequency and severity. In chapter three, the responses by a key exotic annual grass, Bromus tectorum, and keystone native perennial shrub Artemisia tridentata subsp. wyomingensis, were positive for increased early fall precipitation but much more pronounced for B. tectorum. Exotic annual plants are able to respond to changes in timing of fall precipitation and have extreme growth which leads to superior competitive abilities through interference and priority effects (chapter 4). Native plants can compete with exotics but the magnitude of the effects are diminished compared to the negative interaction from exotics. Together these findings demonstrate that across several regions exotic annual grasses are capable of passing through abiotic filters and disrupting biotic interactions of the native plant community. This is likely to lead to increased spread of exotic annual species and may indicate potential and availability of fine fuel production supporting increases in size and frequency of wildfires in the western United States.

Granivores are important components of sagebrush communities in western North America. These same regions are being altered by the invasion of the exotic annual Bromus tectorum (cheatgrass) that alters physical and biological dynamics in ways that appear to promote its persistence. This research directly relates to the restoration of B. tectorum-dominated systems in two inter-related ways. First, because these landscapes have large quantities of seeds applied during restoration, it is important to determine the major granivore communities in intact sagebrush communities and in nearby cheatgrass-dominated communities. Second, it is important to develop an understanding of patterns of seed harvest by granivores. In addition to the data chapters there are two review chapters; Chapter 1 highlights factors contributing to seed removal and Chapter 7 provides ecologically based techniques that could minimize the negative consequences of granivores during ecological restoration. Common groups of ants showed increased abundances; uncommon species and functional groups were generally negatively impacted by cheatgrass (Chapter 2). Conversely, rodents were negatively impacted by conversion to cheatgrass (Chapter 4). Ant seed removal was highly context-dependent (Chapter 3), depending on the background vegetation (large-scale among-patch effects), foraging distance from the nest mound (small-scale among-patch effects), and the presence of other seed species in mixture (within-patch effects). In addition, cheatgrass provided associational resistance to native seeds in mixture, meaning the presence of cheatgrass increased native seed survival. In Chapter 5 a novel statistical technique in the ecological sciences showed that rodents have marked preferences for some seeds over others and that more seeds were removed in sagebrush compared to cheatgrass-dominated sites, although associational effects among seed mixtures were not detected. In Chapter 6 we show that the amount of seed harvested depended on both intraspecific and interspecific seed density. B. tectorum seeds had associational susceptibility (increased harvest) in the presence of native seeds. Although the reciprocal effect may occur, we did not find statistical support for it. These sets of studies are not only of basic ecological interests, but are also important for developing management strategies for restoration of these degraded lands.

Following wildfires, plant materials are direct-seeded to limit erosion and annual weed invasion. Seedlings often fail to establish because selected plant materials are not always well adapted to local soil moisture and temperature conditions. In an effort to help improve plant materials selection and to evaluate sites potential revegetation, we have worked toward developing methodology to predict germination and root growth based on site specific soil moisture and temperature conditions. First, we characterized the seedbed environment of 24 sagebrush (Artemisia spp.) steppe sites throughout the Intermountain West to determine the wet-thermal time of five temperature ranges relevant to germination response and thermal-time model accuracy (Chapter 1). Second, we predicted potential germination for 31 plant materials at those same sites (Chapter 2). Third, in preparation to predict root growth at multiple sites, we characterized the drying patterns and the associated plant-available water for in the seedling root zone across nine woodland (Juniperus spp. and Piñus spp.) sites (Chapter 3). For all of these studies, we determined the effects of tree reduction and tree infilling phase at time of tree reduction. Our key findings are that seedbeds generally sum most wet-thermal time at temperature ranges where the germination rates fit thermal accumulation models quite well (R2 ≥ 0.7). The majority of plant materials summed enough wet-thermal time for a potential germination at most sites during the fall, early spring, and late spring. Soil drying primarily occurs from the soil surface downward. Drying rates and Plant available water associated with the first drying event increased with increasing soil depth. Root zone (1-30 cm) plant-available water increased before and decreased after the first spring drying event with increasing soil depth. Tree removal with increasing pretreatment tree infilling phase generally added progress toward germination, plant available water, and wet-thermal time in the seedbed and root zones of the sagebrush steppe in the Great Basin. Because soil moisture and temperature does not appear to be limiting for potential germination, combining germination and root growth models to create a more comprehensive model may allow for a more robust prediction for seedling survival. For either root growth or combined germination and root growth models, plant available water and wet-thermal time before the first spring drying period hold the most potential for successfully predicting seedling survival.

In the canyon grasslands of Garden Creek Ranch Preserve in Idaho, where the threatened plant Silene spaldingii occurs and invasion by the exotic species Centaurea solstitialis and Bromus tectorum is proceeding rapidly, I examined environmental and community patterns of site invasion, and evaluated the apparent influence of invasion on Silene population vigor. In addition, two separate lightning fires at the preserve presented the opportunity to examine the short-term influence of late-season fire on this species and its associated bunchgrass plant community. I found that Silene-supporting sites most often invaded by exotics were on relatively gentle slopes that received more incident radiation. This pattern may relate, in part, to light requirements of Centaurea solstitialis. Invaded sites were also typically at higher elevations, which may indicate they were moister and therefore more productive. The plant communities in invaded Silene-supporting sites were similar to plant communities in uninvaded sites, although invaded sites tended to have greater legume and exotic annual grass cover. Exotic species invasion did not appear to influence negatively the vigor of Silene populations, as indicated by similar plant height and comparable levels of flowering, fruit and seed set in invaded and uninvaded populations. The similarity in Silene vigor between invaded and uninvaded sites may reflect a moderating influence of site productivity in invaded populations, or may indicate that mature Silene plants and the exotic species partition space or resources differently, potentially reducing competition between them. However, Silene recruitment may be limited by competition from weeds; my data did not allow a rigorous test of this possibility. Fire apparently decreased cover of Festuca idahoensis and increased cover of Lupinus sericeus in the first year after burning, while cover of Pseudoroegneria spicata, exotic grasses, and most other forb species did not differ between burned and unburned areas. Silene cover and abundance within populations were similar before and after fire. Burning did not appear to influence levels of flowering, change the number of flowers or capsules produced per stem, or alter the number of seeds per capsule. Burning decreased plant size slightly, and decreased the proportion of flowers that matured to seed-filled capsules. Silene and the plant communities that support this species appear well suited to late season fire, however the response to burning in other seasons or at higher frequencies remains unknown in this study area. / Graduation date: 2003

Juniper (Juniperus spp.) encroachment into sagebrush (Artemisia spp.) and bunchgrass communities has reduced understory plant cover and allowed juniper trees to dominate millions of hectares of semiarid rangelands. Trees are mechanically masticated or shredded to decrease wildfire potential and increase desirable understory plant cover. When trees are masticated after a major increase in tree population density and associated decrease in perennial understory cover, there is a risk that invasive annual grasses will dominate because they are highly responsive to the increased resource availability that commonly follows removal of the main resource user. To determine if tree mastication increases resource availability and subsequently favors invasive annual or perennial grasses, we compared soil temperature, water, and nutrient microenvironmental conditions and seedling establishment and growth. We used the major rangeland weed, cheatgrass (Bromus tectorum L.), to represent invasive annual grasses and Anatone bluebunch wheatgrass (Pseudoroegneria spicata (Pursh) A. Löve), a natural accession of native bluebunch wheatgrass, to represent the perennial grasses of the sagebrush-bunchgrass plant community. These comparisons were made between and within paired-adjacent masticated and untreated areas at three locations in Utah dominated by Utah juniper (Juniperus osteosperma (Torr.) Little). Juniper tree mastication generally increased resource availability with masticated areas having greater soil temperature, soil water availability, and soil N supply rates than untreated areas. Prior to juniper tree mastication litter mounds were not found to be resource islands probably because juniper trees themselves were using subcanopy soil water and nutrients. After juniper tree mastication and elimination of these predominant resource users, litter mounds served as resource islands with greater soil water availability and N supply rates than bare interspaces during the critical time for seedling establishment in spring. Plant growth followed in line with greater resource availability after tree mastication with masticated areas having more productive although fewer invasive-annual and perennial grass seedlings than untreated areas. These results suggest that increases in resource availability and warmer spring temperatures associated with mastication will not necessarily favor invasive annual over perennial grass seedling establishment. Resilience of the sagebrush-bunchgrass community to return to dominance after juniper control will likely be greatly influenced by how much of the sagebrush-bunchgrass community remains following tree control and the intensity of propagule pressure by invasive species. If only invasive annuals remain when the trees are treated then invasive annuals would be expected to dominate the post-treatment plant community especially with their ability to establish inside litter mounds unless they were also controlled and perennial grasses planted at the time of treatment.