AN ABSTRACT OF THE THESIS OF Joshua P. Averett for the degree of Master of Science in Rangeland Science presented on June 30, 2014. Title: Non-native and Native Plant Species Distributions and Variability along an Elevation Gradient in the Wallowa Mountains, Oregon. Abstract approved: ______________________________________________________ Timothy DelCurto, Michael M. Borman Invasion by non-native plants into natural areas is an important component of global change that threatens biodiversity and ecosystem structure and function. Mountains are currently among the least invaded ecosystems, however, these biodiversity hotspots are increasingly under threat of exotic plant invasion. Evaluation of plant species distribution patterns in mountain ecosystems can provide insight into dominant processes of plant invasion and inform management to reduce the spread of non-native plants in mountains. The objectives of this study were to: 1) evaluate the main drivers of plant invasions in the Wallowa Mountain Range of northeastern Oregon, and the extent patterns of native species distributions and their causes differed from non-native species along an elevation gradient; and 2) investigate inter- and intra-annual understory vascular plant species variability along an elevation gradient to inform long-term monitoring of plant invasion dynamics in the Wallowa Mountains. We sampled understory vascular plant communities in summer 2012 along three forest roads in the Wallowa Mountains. Transects (n=20) were evenly stratified by elevation (60 meters) along each road. Indicator species analysis (ISA) was used to identify habitat tendencies for non-native species. Non-metric multidimensional scaling (NMS) related community composition to environmental and species trait factors. Predictors of non-native and native species abundance and richness were evaluated using Non-parametric multiplicative regression (NPMR). Canopy openness and elevation niche widths were modeled for all common species and evaluated for evidence of high elevation or closed canopy specialization. Non-native species richness decreased continuously with increasing elevation. In contrast, native species richness displayed a unimodal distribution with maximum richness at the mid-elevations. According to NMS and NPMR, elevation, canopy openness, and disturbance were the strongest correlates of non-native species abundance. Non-native species were concentrated in low elevation bunchgrass, roadside, and open forest habitats with high canopy openness, and moderate to high disturbance intensity. Overlays of species trait factors onto ordinations revealed that the transition into the subalpine community and over-story canopy closure exceeding approximately 60 percent were potential barriers to non-native species establishment. Vegetation sampling was repeated three times (June, July, and August 2013) during the growing season in summer 2013 for a subset (n=10) of our transects. Blocked Multi Response Permutation Procedure (MRBP) was used to test the hypothesis of no change in species composition between years (summer 2012 and summer 2013) and between each sampling period (June, July, and August) within one growing season. NMS related community composition at different times to environmental and trait category factors. Blocked Indicator Species Analysis was used to identify species with tendencies towards specific sampling periods. Important predictors of vegetation change were identified and indicator species distributions were related to NMS axes using NPMR. Vegetation composition changed both between years and between each sampling period within one growing season. Species richness increased with sampling effort, dominant species were consistently found throughout all sampling periods and rare species showed the highest rate of turnover between sampling periods. The highest species richness and greatest number of indicator species were associated with the early summer (June) sampling period. Approximately 25% of all non-native species showed tendencies towards a specific sampling period where greater than half favored the June sampling period within lowland bunchgrass communities. Annual species including the most dominant non-native species within the lowland and montane zones showed the highest inter- and intra-annual variability compared to other trait categories. Total species richness was positively related to turnover and was the most important predictor of relative abundance change between all sampling periods. Species turnover was highest in species rich montane plots; however, trait abundance and richness did not statistically differ in montane plots. These data suggest that elevation, canopy, openness, and disturbance are important factors structuring non-native plant distributions in the Wallowa Mountains. Our findings that non-native species tended to be concentrated in low- elevation, early successional habitats coupled with a lack of high elevation or closed canopy specialization indicates that non-native plants that occurr at the highest elevations are likely generalist species that depend on disturbance (primarily open over-story canopy) for spread into higher elevation sites until they are eventually filtered out by harsh environmental conditions coincident with the subalpine transition zone. Collectively these results suggest that interactions between introduction pathways that favor human assisted dispersal into the low-elevations where ruderal species are favored and environmental filters (elevation, canopy openness, and disturbance) are the dominant factors structuring non-native plant distributions in the Wallowa Mountains. Our results also indicate that long-term monitoring of vegetation change in the Wallowa Mountains will be improved through periodic sampling during each growing season. Sampling once during the spring or early summer and then again coincident with peak biomass should improve estimates of species distributions and diversity patterns by capturing both vernal and late season species. Early season sampling may be particularly important for monitoring non-native plants in the Wallowa Mountains as most non-natives were concentrated in low elevation semi- arid bunchgrass communities where spring and early summer specialization is common. ©Copyright by Joshua P. Averett June 30, 2014 All Rights Reserved Non-native and Native Plant Species Distributions and Variability along an Elevation Gradient in the Wallowa Mountains, Oregon by Joshua P. Averett A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Master of Science Presented June 30, 2014 Commencement June 2015 Master of Science thesis of Joshua P. Averett presented on June 30, 2014 APPROVED: Co-Major Professor, representing Rangeland Sciences Co-Major Professor, representing Rangeland Sciences Head of the Department of Animal and Rangeland Sciences Dean of the Graduate School I understand that my thesis will become part of the permanent collection of Oregon State University libraries. My signature below authorizes release of my thesis to any reader upon request. Joshua P. Averett, Author ACKNOWLEDGEMENTS I am grateful to Tim DelCurto and Michael Borman for taking me on as their graduate student and for their guidance through my graduate school experience. I would like to express great thanks to Ricardo Mata-González and Bruce McCune for their excellent mentorship, technical support, and consistent optimism and encouragement throughout my research journey. Thank you to Julia Jones for the great advice and support that helped to steer me back on course whenever I veered. I also extend great thanks to Catherine Parks for her support with the conceptual design of this study and assistance with field sampling and Bridgett Naylor for her support and patience with data extraction and database management. This research would not have been possible without my outstanding field crew. A heart-felt thanks goes out to Kent Coe, Rachel Burton, Regina Thompson, Ryan Limb, and Samantha Pack. I would like to express gratitude to my family for all of their love and support through and leading up to this experience. Thanks to my mother Betty, father Jerry, and brother Jason. Lastly, and most importantly, I would like to thank my wife Adrienne. Without her love and encouragement to follow my passions, none of this would have been possible. CONTRIBUTION OF AUTHORS Dr. Bruce McCune contributed to the entire thesis through guidance and editing. Dr. McCune also provided technical support regarding analytical approaches and implementation of Non-metric multidimensional scaling, Non-parametric multiplicative regression, Blocked Multi-response Permutation Procedures, and Blocked Indicator Species Analysis using HyperNiche 2.0 and PC-Ord 6.0. Dr. Catherine Parks provided guidance regarding research questions, provided edits, and assisted in the development of the sampling protocol for both chapters. Bridgett Naylor contributed to chapters 2 and 3 through support of data extraction, data-base management, and assisted with the development of the sampling protocol. Dr. Ricardo Mata-González provided guidance and editing for the entire thesis. TABLE OF CONTENTS Chapter 1 Introduction .................................................................................................. 1 Objectives ...................................................................................................................................... 11 Chapter 2 Non-native and native plant distributions