Geospatial Variation of an Invasive Forest Disease and the Effects on Treeline Dynamics in the Rocky Mountains
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Geospatial Variation of an Invasive Forest Disease and the Effects on Treeline Dynamics in the Rocky Mountains Emily Katherine Smith-McKenna Dissertation submitted to the faculty of the Virginia Polytechnic Institute and State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy In Geospatial and Environmental Analysis Lynn M. Resler, Chair George P. Malanson Stephen P. Prisley Laurence W. Carstensen Jr. October 2, 2013 Blacksburg, Virginia Keywords: Treeline, Whitebark Pine, Pinus albicaulis, Blister Rust, Rocky Mountains, spatial pattern, GIS, GPS, DEM, Agent-Based Model Copyright 2013, Emily K. Smith-McKenna Geospatial Variation of an Invasive Forest Disease and the Effects on Treeline Dynamics in the Rocky Mountains Emily Katherine Smith-McKenna ABSTRACT Whitebark pine is an important keystone and foundation species in western North American mountain ranges, and facilitates tree island development in Rocky Mountain treelines. The manifestation of white pine blister rust in the cold and dry treelines of the Rockies, and the subsequent infection and mortality of whitebark pines raises questions as to how these extreme environments harbor the invasive disease, and what the consequences may be for treeline dynamics. This dissertation research comprises three studies that investigate abiotic factors influential for blister rust infection in treeline whitebark pines, how disease coupled with changing climate may affect whitebark pine treeline dynamics, and the connection between treeline spatial patterns and disease. The first study examined the spatial variation of blister rust infection in two whitebark pine treeline communities, and potential topographic correlates. Using geospatial and field approaches to generate high resolution terrain models of treeline landscapes, microtopography associated with solar radiation and moisture were found most influential to blister rust infection in treeline whitebark pines. Using field-based observations from sampled treeline communities, the second study developed an agent-based model to examine the effects of disease and climate on treeline pattern and process. Treeline dynamics were simulated under five hypothetical scenarios to assess changes in tree spatial patterns and populations. Blister rust-induced loss of whitebark pines resulted in a decline of facilitative processes, and an overall negative treeline response to disease—despite the beneficial effects of climate amelioration. The objective of the third study was to explore whether spatial patterns of tree proximity, size, and growth infer disease patterns. Comparing spatial patterns of tree characteristics between diseased and undiseased treeline communities, I found that trees growing near trees with larger stem diameters, and larger tree islands, tended to have more blister rust cankers, and displayed clustered spatial patterns. Undiseased treeline patterns revealed near neighbors smaller in stem diameter and tree island size, and were randomly dispersed. Blister rust diseased whitebark pines reveal spatial autocorrelation, despite the complex blister rust disease life cycle. Overall, findings from this dissertation reveal the implications of invasive disease on sensitive treeline ecotones dependent on a keystone species. Acknowledgements The journey toward the doctorate is a challenging endeavor. It is full of hurdles and setbacks, which are somehow compensated by the gratification of accomplishment to help make the push further. To the many people who provided mentorship and support to me through the process, and helped me make this push—thank you. My dissertation research was primarily supported by a National Science Foundation Geography and Spatial Sciences Program grant (808548) awarded to Dr. Lynn Resler, Dr. Diana Tomback, and Dr. George Malanson. I feel fortunate that I had the opportunity to work with these research experts in treeline ecosystems, in such beautiful and scenic mountain landscapes, and to have them as coauthors on papers—it has made me a better writer. I express much thanks to my committee members: to my advisor Lynn Resler, whose knowledge and experience guided my efforts throughout the PhD process; to George Malanson, his expertise in agent-based modeling and feedback processes provided an invaluable resource (“good stuff Maynard!”); and to Steve Prisley and Bill Carstensen, who helped me hash out ideas for the geospatial and statistical components of my research, and above all provided their support, encouragement, and trust in my accomplishments. Thank you. Sharing the trials and tribulations of being a graduate student with my fellow PhD graduate students, both in the VT Geography Department and outside, was an important outlet and they made the experience so much more fun. In particular, Candice Luebbering, Arvind Bhuta, Melanie Stine, Jess Walker, Baojuan Zheng, and Tammy Parece—I appreciate your friendship and support as we worked towards the PhD. And to Jim Campbell, even though our office was at the far end of the building and end of the hallway, you always stopped by to visit us in the GEA office—thanks for lending your insight over the years. A special thanks to my fellow band members, music fed my soul—long live the Septapüs! And to Sinking Creek Pottery, working clay into art kept me balanced. Most importantly, I am so thankful for the immeasurable support from my family and friends. I especially owe a heartfelt thanks to my husband John McKenna, who has patiently endured the ups and downs of my journey toward the doctorate—I love you. iii Attribution My coauthors for chapters 3 and 4 consist of my advisory committee, co-principal investigator for the supporting research grant, and colleague. A brief description of their contributions towards each chapter is outlined below. Chapter 3: Topographic Influences on the Distribution of White Pine Blister Rust in Pinus albicaulis Treeline Communities Emily K. Smith-McKenna, Lynn M. Resler, Diana F. Tomback, Huaiye Zhang, and George P. Malanson Chapter 4: Feedbacks, disease, and climate influences on whitebark pine treeline dynamics: An agent-based treeline model. Emily K. Smith-McKenna, George P. Malanson, Lynn M. Resler, Laurence W. Carstensen, Stephen P. Prisley, and Diana F. Tomback Dr. Lynn M. Resler, Ph.D., is an Associate Professor in the Department of Geography at Virginia Tech. Dr. Resler was a co-author on chapters 3 and 4, my advisor, principal investigator of the National Science Foundation grant that supported this research, and contributed intellectual and editorial comments. Dr. Diana F. Tomback, Ph.D., is a Professor in the Department of Integrative Biology at the University of Colorado Denver. Dr. Tomback was a co-author on chapters 3 and 4, co-principal investigator of the National Science Foundation grant that supported this research, and contributed intellectual and editorial comments. Dr. Huaiye Zhang, Ph.D., earned his doctorate at the Department of Statistics, Virginia Tech. Dr. Zhang was a co-author on chapter 3, and was a consultant at the Virginia Tech’s Laboratory for Interdisciplinary Statistical Analysis where he contributed his statistical expertise to the statistical model. Dr. George P. Malanson, Ph.D., is the Coleman-Miller Professor in the Department of Geographical and Sustainability Sciences at the University of Iowa. Dr. Malanson was a co- author on chapters 3 and 4, co-principal investigator of the National Science Foundation grant that supported this research, and contributed intellectual and editorial comments and his expertise to the agent-based model in Chapter 4. Dr. Laurence W. Carstensen, Ph.D., is Professor and Head of the Department of Geography at Virginia Tech. Dr. Carstensen was a co-author on chapter 4, and contributed intellectual and editorial comments. Dr. Stephen P. Prisley, Ph.D., is an Associate Professor in the Department of Forest Resources and Environmental Conservation at Virginia Tech. Dr. Prisley was a co-author on chapter 4, and contributed intellectual and editorial comments. iv Table of Contents List of Figures .............................................................................................................................. viii List of Tables ................................................................................................................................. xi Chapter 1: Introduction .................................................................................................................. 1 Significance ................................................................................................................................. 3 Objectives .................................................................................................................................... 5 References ................................................................................................................................... 7 Chapter 2: Literature Review ....................................................................................................... 12 The Alpine Treeline Ecotone And The Role Of Whitebark Pine In Treeline Formation ......... 12 Treeline Whitebark Pine and Blister Rust ................................................................................. 14 Treeline Processes In Whitebark Pine Ecosystems ................................................................... 16 Indirect/Direct Effects ..........................................................................................................