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Dispersal Effects on Species Distribution and Diversity Across Multiple Scales in the Southern Appalachian Mixed Mesophytic Flora

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DISPERSAL EFFECTS ON SPECIES DISTRIBUTION AND DIVERSITY ACROSS MULTIPLE SCALES IN THE SOUTHERN APPALACHIAN MIXED MESOPHYTIC FLORA Samantha M. Tessel 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 Ecology in the Curriculum for the Environment and Ecology. Chapel Hill 2017 Approved by: Peter S. White Robert K. Peet Alan S. Weakley Allen H. Hurlbert Dean L. Urban ©2017 Samantha M. Tessel ALL RIGHTS RESERVED ii ABSTRACT Samantha M. Tessel: Dispersal effects on species distribution and diversity across multiple scales in the southern Appalachian mixed mesophytic flora (Under the direction of Peter S. White) Seed and spore dispersal play important roles in the spatial distribution of plant species and communities. Though dispersal processes are often thought to be more important at larger spatial scales, the distribution patterns of species and plant communities even at small scales can be determined, at least in part, by dispersal. I studied the influence of dispersal in southern Appalachian mixed mesophytic forests by categorizing species by dispersal morphology and by using spatial pattern and habitat connectivity as predictors of species distribution and community composition. All vascular plant species were recorded at three nested sample scales (10000, 1000, and 100 m2), on plots with varying levels of habitat connectivity across the Great Smoky Mountains National Park. Models predicting species distributions generally had higher predictive power when incorporating spatial pattern and connectivity, particularly at small scales. Despite wide variation in performance, models of locally dispersing species (species without adaptations to dispersal by wind or vertebrates) were most frequently improved by the addition of spatial predictors. Patterns in plant communities were also compared among dispersal categories, and though it was shown that species without dispersal adaptations were less likely to co-occur, this was more likely to be caused by differences in frequency across dispersal categories than by dispersal limitation per se. Spatial pattern, distance, and connectivity were significant predictors of non-random patterns of species turnover at all scales and were stronger among dispersal- iii limited species groups. Species with limited dispersal were also less frequent at all three sampling scales than those with adaptations for vertebrate dispersal and had smaller geographic ranges than either wind- or vertebrate-dispersed species. Species with no dispersal adaptations were overrepresented among southern Appalachian endemics and lineages endemic to North America, whereas species dispersed by vertebrates were overrepresented among Tertiary disjunct lineages, and species dispersed by spores were cosmopolitan. Relationships among dispersal and biogeographic affinity reflect the evolutionary history of the mixed mesophytic flora owing to adaptation of dispersal mechanisms to regional environmental conditions, the relative ages of lineages, and the effect of dispersal mechanism on the distributions of plants across time and space. iv To my family, for fostering in me a love of plants, to Lucy Braun, who continues to inspire me despite the fact that she died long before I was born, and to Yontof, who always tried to steal my sedges from underneath the microscope. v ACKNOWLEDGEMENTS I could not have completed this dissertation without the acedemic and moral support of many people. First, I owe my thanks to Peter, who has always been willing to help in any way he can, and is always a source of stimulating ideas and ecological and botanical tangents. This dissertation would not have been the same without the botanical, ecological, statistical, and theoretical expertise learned from interactions with my other committee members, Bob, Alan, Dean, and Allen. My time doing field work in the Smokies was wonderful, and I am grateful to all those that helped me in the field, in the science center, and in the cabin: Megan McClung, Julie Tuttle, Tom Colson, Tom Remaley, Bill Mize, and Jackie White. For both moral support and botanical fun, I am indebted to Derick Poindexter, Pat Gensel, and Carol Ann McCormick in addition to Megan and Julie. And as always, I am grateful for the love and support from my family, my friends, and my cat Yontof. Last but not least, for being an unending source of inspiration and wonder, thanks to all plants (especially the ones with umbrella-shaped and ternately compound leaves). vi TABLE OF CONTENTS LIST OF TABLES ........................................................................................................................ xii LIST OF FIGURES ..................................................................................................................... xiii CHAPTER 1. SYSTEMS INTRODUCTION: THE HISTORY AND COMPOSITION OF THE MIXED MESOPHYTIC FORESTS OF THE SOUTHERN APPALACHIAN MOUNTAINS ............................................................................. 1 The legacy of E. Lucy Braun....................................................................................................... 2 Mixed mesophytic forest definition......................................................................................... 3 Braun's assumptions ................................................................................................................ 4 Climax communities ................................................................................................................ 5 Tertiary geoflora ...................................................................................................................... 7 Modern understanding of vegetation history .............................................................................. 8 Dispersal and migration ............................................................................................................ 13 Post-glacial migration ............................................................................................................ 13 The distribution of mixed mesophytic forest species in the southern Appalachians ............ 15 Inferring dispersal in natural communities ............................................................................ 16 Sampling mixed mesophytic forests ...................................................................................... 16 Analysis of scales of dispersal importance in mixed mesophytic forests ............................. 18 References ................................................................................................................................. 23 CHAPTER 2. EVALUATING THE EFFECTS OF DISPERSAL PROCESSES ON PLANT SPECIES DISTRIBUTIONS AND DIVERSITY ACROSS MULTIPLE SCALES: A REVIEW AND SYNTHESIS ............................................................. 29 Introduction ............................................................................................................................... 29 Scales of dispersal processes ..................................................................................................... 30 vii Spatial pattern and dispersal .................................................................................................. 31 Space and time interaction..................................................................................................... 32 The local-regional dichotomy................................................................................................ 33 Characteristic scales of heterogeneity ................................................................................... 34 Neighborhood scale ........................................................................................................... 35 Habitat scale ....................................................................................................................... 36 Landscape scale ................................................................................................................. 37 Regional scale .................................................................................................................... 38 Biogeographic scale ........................................................................................................... 39 Relativity in space and time .................................................................................................. 41 Tools for assessing the importance of dispersal ........................................................................ 42 Approaches based on dispersal of individual focal species................................................... 46 Quantifying autocorrelation ............................................................................................... 46 Species distribution models ............................................................................................... 47 Landscape-scale occurrence and abundance ...................................................................... 49 Co-occurrence based approaches .......................................................................................... 50 Approaches addressing the effects of dispersal and colonization on α-diversity .................. 51 Approaches analyzing or explaining β-diversity ................................................................... 53 Distance approach .............................................................................................................
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