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Trait-Based Inference of Environmental Constraints on Lichen Epiphyte Communities at Multiple Spatial Scales

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Trait-Based Inference of Environmental Constraints on Lichen Epiphyte Communities at Multiple Spatial Scales TRAIT-BASED INFERENCE OF ENVIRONMENTAL CONSTRAINTS ON LICHEN EPIPHYTE COMMUNITIES AT MULTIPLE SPATIAL SCALES Jessica Coyle A dissertation submitted to the faculty at the University of North Carolina at Chapel Hill in partial fulfillment of the requirements for the degree of Doctor of Philosophy in the Department of Biology. Chapel Hill 2016 Approved by: Allen Hurlbert Charles Mitchell Robert Peet Joel Kingsolver François Lutzoni © 2016 Jessica Coyle ALL RIGHTS RESERVED ii ABSTRACT Jessica Coyle: Trait-based inference of environmental constraints on lichen epiphyte communities at multiple spatial scales (Under the direction of Allen Hurlbert) Ecological systems respond to processes that operate at a variety of spatial scales. Epiphytic lichens are a useful group for studying community response to cross-scale environmental variation because tree and forest architecture organize epiphyte communities into discrete hierarchical scales and because lichens’ limited homeostatic ability makes them potentially sensitive environmental indicators. Examining variation in the functional traits of organisms can elucidate the role of the environment in structuring communities because these traits provide a direct link between organismal fitness and environmental filters. For lichens, such trait-based approaches are not yet fully developed. This dissertation examines how lichen traits can provide insight into environmental constraints on communities across multiple spatial scales. The primary goals of this research were to 1) evaluate the extent to which larger-scale processes influence lichen epiphyte communities at smaller scales and 2) assess the utility of functional versus morphological trait- based approaches for understanding community assembly in lichen epiphytes. I utilized a national forest inventory (U.S. Forest Service) and conducted two field-based surveys in temperate deciduous forests across North Carolina (U.S.A.) to assess environmental factors affecting lichen assemblages at three focal scales: forest patches, tree trunks, and individual branches within tree canopies. To evaluate trait-based approaches, I quantified the response of functional and morphological traits to environmental variation at different scales and assessed iii whether observed relationships were consistent with hypothesized environmental constraints. Results highlighted a previously underappreciated role for larger-scale processes in determining the composition of lichen epiphyte assemblages at multiple scales, but also revealed high community variability at small scales, which was not well accounted for by linear environmental models. This suggests a potentially important role for stochasticity in lichen community formation. Analyses also indicated that future development of trait-based approaches to lichen community studies should focus on quantifiable and functionally interpretable traits rather than the categorical characters used for species identification. This research demonstrates that traits are a useful, but not powerful, tool for understanding processes shaping lichen assemblages. iv ACKNOWLEDGEMENTS I thank my advisor, Allen Hurlbert, for his consistently insightful advice and mentorship as I dove headlong into a study system new to both of us. Allen has had a profound impact on my writing and approach to research and mentoring. I also thank my thesis committee; François Lutzoni, Robert Peet, Charles Mitchell, and Joel Kingsolver for their enthusiasm and willingness to provide guidance on a topic outside of their primary areas of expertise as well as for helpful reviews of previous drafts of dissertation chapters. In addition, all members of my committee generously provided field equipment and laboratory facilities critical to the success of this research. The direction of my thesis and my thinking on traits and community assembly were initially developed and shaped by discussions with the Dimensions of Biodiversity Distributed Graduate Seminar team: Bianca Lopez, Kyle Palmquist, Fletcher Halliday, and Peter Wilfahrt. I am fortunate to have them as colleagues and friends. This research concluded in a timely manner only through the work of several field- and lab-assistants and I am grateful for their efforts: Mariah Goodman, Holly Pittard, Elias Athey, Jesse Fitzpatrick, Zach Mackenzie, Alex Gerber, Brian Sherman, Michael Auriemma, Sarah Krantz, Shelly Guo, Saloman Ariza, Ivy Somocurcio, Miguel Sandoval, and Bolu Aluko. This research also relied heavily on existing data on lichen distributions and climate and I want to acknowledge all those involved in collecting these data and making them publically accessible. In particular, thanks to Sarah Jovan for assistance locating data associated with the former Forest Health Monitoring Program. I would like to thank Water Dog Farms, Duke Forest, the U.S. Forest Service, North Carolina State Parks and the Triangle Land Conservancy for permission to conduct research on their lands as well as the NCU Herbarium for access to research facilities. This research was financially supported by the National Science Foundation, the Royster Society of Fellows, and Highlands Biological Station. Finally, I would like to express my deep gratitude to Joseph E. Pogue for endowing the fellowship that supported me throughout my graduate studies. vi TABLE OF CONTENTS LIST OF TABLES .......................................................................................................................... x LIST OF FIGURES ...................................................................................................................... xii INTRODUCTION ........................................................................................................................ 14 CHAPTER 1: ENVIRONMENTAL OPTIMALITY, NOT HETEROGENEITY, DRIVES REGIONAL AND LOCAL SPECIES RICHNESS IN LICHEN EPIPHYTES ........... 17 Introduction ............................................................................................................................... 17 Methods..................................................................................................................................... 20 Data ....................................................................................................................................... 20 Models................................................................................................................................... 22 Results ....................................................................................................................................... 25 Geographic distribution of lichen richness ........................................................................... 25 Variation partitioning ............................................................................................................ 25 Multi-model inference and parameter estimation ................................................................. 27 Discussion ................................................................................................................................. 28 Regional control of local communities ................................................................................. 28 Dominance of environmental optimality over heterogeneity across scales .......................... 30 Implications beyond lichen ecology ..................................................................................... 33 CHAPTER 2: SPATIAL VARIANCE IN DIVERSITY AND FUNCTIONAL STRUCTURE OF LICHEN EPIPHYTE ASSEMBLAGES IN DECIDUOUS FORESTS OF THE SOUTHEASTERN UNITED STATES ....................................................... 44 Introduction ............................................................................................................................... 44 Methods..................................................................................................................................... 46 Study location ....................................................................................................................... 46 Environmental data ............................................................................................................... 47 Lichen community assessment ............................................................................................. 48 Analysis of community variation across scales .................................................................... 50 Results ....................................................................................................................................... 52 Community variation across scales ....................................................................................... 52 Community response to environment ................................................................................... 53 Discussion ................................................................................................................................. 54 Epiphyte community assembly ............................................................................................. 54 Utility of taxon-free surveys of lichens ................................................................................ 56 Future directions ................................................................................................................... 58 CHAPTER 3: DETECTION OF ASSEMBLY PROCESSES IN OBLIGATE MUTUALIST COMMUNITIES USING MORPHOLOGICAL TRAITS .................................. 67 Introduction ..............................................................................................................................
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