I investigated how shrub-induced spatial heterogeneity influenced and was manifested by a representative ground-story plant species at a sagebrush-steppe site in southwestern Wyoming. The dispersion of cheatgrass (Bromus tectorum) reflected differences between undershrub (higher plant densities) and interspace (between shrubs, supporting lower densities) microenvironments, hence the population ecology of this annual species served as a biological probe of shrub-associated patch structure in this community. Since cheatgrass is an annual, factors affecting the seed portion of its life cycle were of special interest. First, attributes of the above- and below-ground seed pool were characterized. The environment-wide seed depositional pattern was assessed using seed traps of several designs, and the legacy of seed incorporation into the soil was examined by separating seeds from soil samples. For both components of the seed pool, annuals' seeds predominated. Seeds at the surface were subject to substantial redistribution, moving readilythrough interspace, and their deposition was related to both the interaction of wind and shrub canopies and the presence of litter. More annuals' seeds were encountered in undershrub than in interspace soils; seeds of cheatgrass were restricted to the soil surface. Second, a manipulative experiment tested effects of granivoryfherbivory and presence/absence of a replicate shrub's canopy upon success of cheatgrass plants arising from known numbers of seeds introduced into undershrub versus interspace microenvironments. While biomass of plants in treatments accessible to herbivores was less than that of protected plants, consumers did not affect plant densities, and herbivore effects were not microenvironment-specific . Shrub canopy removal had no effect on plant success, and, contrary to expectations based on the dispersion of indigenous plants, interspace plants fared better than undershrub counterparts. Finally, demographic fates of individually marked seeds were observed, to disentangle effects of microenvironment from effects of microenvironment-specific surface types on determining safe sites. Littered microsites were strongly associated with undershrub microenvironments, and on these surfaces, cheatgrass seeds were less likely to move and to suffer depredation, and more likely to become favorably positioned for subsequent germination and establishment, than on bare ground surfaces typifying interspace. \

Changes in shrubsteppe passerine bird habitat associations in response to disturbance were investigated at multiple temporal and spatial scales. Spatial measures incorporated the effects of area at different ecological scales (nest site, territory, and landscape) to include ecologically meaningful extents. Temporal measures included seasonal and annual effects, and were designed to detect lagged responses should they occur. Local-to-landscape scale effects of mechanical restoration treatments on local extirpation and abundances of nine species indicated most were insensitive to changes in habitat quality, while abundance models showed only broad declines. Changing the availability of nesting habitat on both the attractiveness and quality of an area at multiple extents confirmed the need for long-term study effects due to lagged responses in expressed preference and changes to nesting habitat quality. Time since treatment affected nest success in two of the four species, yet the changes in habitat quality did not forecast changes in habitat preference as expected. Non-adaptive mismatches seemingly occurred as habitat preferences indicated treatments may create benign-appearing 'sink' habitat for species that remained in the area. The umbrella species concept is misapplied at this scale: each species' response was consistent, but responses varied in scale, timing, and direction among species. Patterns of nest density and nest site descriptions demonstrated population-level movement in response to treatments, suggesting half the focal species moved nest sites to remaining habitat areas. Larger scale responsive movements were observed in the remaining species, both out of and into the nest plot. Descriptions of nesting habitat characteristics for the focal species tested if the selected nesting habitat was consistent between pre- and post-treatment, and determined which habitat characteristics, including distance to disturbance, were related to nest success. Descriptions of nesting habitat characteristics support previous work in terms of structural characteristics. Habitat selection was consistent even when the available habitat was not, implying these species choose sites and are not merely settling randomly. However, selected nesting habitat was not strongly tied to nest success at local scales and nest success was negatively related to landscape qualities that treatments were designed to enhance.

Thesis (M.S.)--Oregon State University, 2009. / Printout. Includes bibliographical references (leaves 90-100). Also available on the World Wide Web.

Thesis (Ph. D.)--University of Wyoming, 2007. / Title from PDF title page (viewed on July 15, 2008). Includes bibliographical references.

Thesis (Ph. D.)--Oregon State University, 2011. / Printout. Includes bibliographical references (leaves 123-135). Also available on the World Wide Web.

Planting native species after a major disturbance is a critical tool land managers use to stabilize soils, restore ecosystem processes, and prevent weed invasion. However, within the sagebrush steppe and other arid and semi-arid environments the percentage of sown seeds that produce an adult plant is remarkably low. Applying fertilizers at the time of planting may improve native plant establishment by increasing the ability of the seedlings to cope with environmental stresses. However, traditional fertilizer applications are often economically infeasible and may be counterproductive by encouraging weed invasion. Seed coating technology allows for the efficient application of fertilizers within the microsite of the seeded species. The objective of our research was to determine the optimal rate of fertilizer to apply to the seed to improve seedling emergence and plant growth. We applied a phosphorus (P) rich fertilizer (0.13 g P g-1) to bluebunch wheatgrass (Pseudoroegneria spicata (Pursh) Á. Löve) seeds in a rotary coater at rates ranging from 0 to 50 g of fertilizer 100 g-1 seed. Three separate studies were conducted to test germination, biomass, relative growth rate, and tissue nutrient uptake. Study one showed decreasing root and shoot biomass and increasing time to 50% germination as fertilizer rates increased. Study two showed no difference in relative growth rate between the controls and fertilizer treatments. Study three showed no difference in root and shoot biomass or nutrient concentration between treatments except in the lowest fertilizer treatment (10 g fertilizer 100 g-1 seed), which was significantly lower in root and shoot biomass than all other treatments but had higher P tissue concentrations than all other treatments. Collectively these results showed no evidence that a P fertilizer coating could aid in bluebunch wheatgrass seedling establishment. Because bluebunch wheatgrass and similar late-seral plants have evolved with low nutrient requirements they may not be physiologically capable of handling increased nutrient supply, which may explain the results of our studies. Continued studies and fieldwork need to be performed to evaluate the potential of fertilizer seed coatings in restoration efforts.

Understanding the relative importance of density-dependent and density-independent factors in driving population dynamics is one of the oldest challenges in ecology, and may play a critical role in predicting the effects of climate change on populations. We used long-term observational data to describe patterns in plant population regulation for 57 forb and grass species from three different ecosystems (arid desert grassland, semiarid sagebrush steppe, and subhumid mixed-grass prairie). Using a hierarchical partitioning approach, we (i) quantified the relative influence of conspecific density, heterospecific composition, and climate on temporal variation in population growth rates, and (ii) asked how the relative importance of these drivers depends on site aridity, species growth form and life expectancy, and abundance and spatial patterns. The data from one of the sites in this analysis are presented in one of the chapters of this thesis. We found that density-dependence had the strongest effect on species. Climate often had a significant effect, but its strength depended on growth form. Community composition rarely explained significant variation in growth rates. The relative importance of density, composition, and climate did not vary among sites, but was related to species' life histories: compared to forbs, grasses were more sensitive to climate drivers. Abundance and spatial clustering were negatively correlated with the importance of density dependence, suggesting that local rarity is a consequence of self-limitation. Our results show that interspecific interactions play a weaker role than intraspecific interactions and climate variability in regulating plant populations. Forecasting the impacts of climate change on populations may require understanding how changes in climate variables will affect the strength of density-dependence, especially for rare species.

Forecasting the effects of climate change on plant and animal populations is a high priority in ecology. We studied the effects of climate on plant populations through the use of observational and experimental data, as well as analytical models. Our research questions were: (1) Do the effects of interannual climate variation on the population growth rates of widespread species show a coherent pattern across gradients of mean annual climate? (2) How well can population models fit to observational data predict the response of populations to field experiments that manipulate climate? And (3) does niche overlap between competitors predict the magnitude of competition-mediated indirect effects in mechanistic resource competition models? To test the first question, we assessed how interannual variation in climate affected the abundance of big sagebrush (Artemisia tridentata) at 131 monitoring sites across its range. We found that years of above average temperature increased sagebrush abundance at cold sites, but decreased sagebrush abundance at hot sites. This pattern indicates that sagebrush distribution may be limited by hot and cold temperatures at the extremes of its distribution. We addressed iv our second research question by fitting statistical models to over 25 years of observational data on the performance of four dominant plant species in a sagebrush steppe community. We then experimentally manipulated soil moisture in this community and tested how well the statistical models fit to observational data could predict species’ responses to the experimental treatments. In two out of four species, we found that including climate effects in our models helped us predict the population-level responses to the experiment. Moreover, effects of historical soil moisture variation on vital rates were generally consistent with the effects of drought and irrigation treatments. Our results provide some evidence that observational data can be used to predict species’ responses to climate change in the future. We addressed our third question by simulating environmental change in analytical models of resource competition and quantifying the size of direct and competition-mediated indirect effects that resulted. We showed that the magnitude of indirect effects increased as the niche overlap between competitors increased.

In recent years, the importance of sagebrush to shrub-steppe ecosystems and associated plant and animal species has been recognized. The historical removal of herbaceous species by excessive and uncontrolled livestock grazing on many of our sagebrush ecosystems has resulted in a stagnant state where dense, competitive stands of sagebrush prevent herbaceous species from recovering. Most early research on sagebrush control was directed toward eradication to increase herbaceous forage for livestock production, rather than sagebrush thinning to improve shrub vigor and understory production for wildlife habitat and community diversity. Mechanical treatments have the ability to retain shrub and herbaceous components, while improving diversity within degraded sagebrush communities. This study evaluated the effects of 6 mechanical treatments and revegetation of a Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis) community in northern Utah that were treated in the fall of 2001 and spring of 2002 (aerator only). Disking and imprinting killed 98% of the sagebrush and significantly (p < 0.05) lowered cover and density of sagebrush more than any other treatment. Disking and imprinting was the only mechanical treatment to reduce cover and density of residual understory species, but also to successfully establish seeded grasses. One-way Ely chaining, 1-way and 2-way pipe harrowing, and aerating in the fall and spring reduced sagebrush cover from greater than 20% to less than 5% and reduced density by about half. Two years after mechanical treatment surviving sagebrush had greater leader and seed stalk growth than untreated sagebrush. Choice of a mechanical treatment to increase and diversify the perennial herbaceous component and retain the shrub component of sagebrush communities depends on the amount of residual herbaceous species, as well as economics. Chaining is potentially most economical for diversifying communities with a residual herbaceous perennial component. It is uncertain whether successful revegetation from disking and imprinting was a result of significant reduction in sagebrush, residual perennial herbaceous species, or both. Response of sagebrush communities with a very limited perennial herbaceous understory needs to be tested to determine how much and what kind of mechanical reduction in sagebrush is needed for successful revegetation.

Land-use legacies can affect landscapes for decades to millennia. A long history of shrub management exists in the sagebrush steppe of the Intermountain West where shrub-removal treatments, a type of managed disturbance, have been implemented for over 50 years to reduce sagebrush cover. The assumption behind managed disturbances is that they will increase forage for domestic livestock and improve wildlife habitat. However, the long-term effects of managed disturbance on plant community composition and diversity are not well understood. We investigated the legacy effects of three common types of managed disturbance (chemical, fire, and mechanical treatments) on plant community diversity and composition. We also examined sagebrush steppe resilience to managed disturbance. Based on management assumptions and resilience theory, we expected within-state phase shifts characterized by an initial reduction in biodiversity followed by a return to prior state conditions. We also expected changes in species proportions, characteristic of within-state shifts in state-and-transition models. We also expected an increase in non-native contribution to overall diversity. We found that plant communities experienced a fundamental shift in composition following disturbance, and responded in a flat linear fashion, giving no indication of return to prior community composition or diversity. As expected, we found post-disturbance increases in the number of non-native grass species present. However, native forb species made the largest contribution to altered diversity. Disturbance modified functional group composition, so contrary to our expectations, within-state changes did not occur as a result of disturbance. Our results indicated that sagebrush steppe plant communities are not resilient to chemical, fire, and mechanical treatments, and subsequent to managed disturbance, community composition tips over a threshold into an alternate stable state.