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Accounting for Intraspecific Variation Transforms Our Understanding of Artiodactyl Social Evolution

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Accounting for Intraspecific Variation Transforms Our Understanding of Artiodactyl Social Evolution ACCOUNTING FOR INTRASPECIFIC VARIATION TRANSFORMS OUR UNDERSTANDING OF ARTIODACTYL SOCIAL EVOLUTION By Monica Irene Miles Loren D. Hayes Hope Klug Associate Professor of UC Foundation Associate Professor of Biology, Geology, and Environmental Science Biology, Geology, and Environmental Science (Chair) (Committee Member) Timothy Gaudin UC Foundation Professor of Biology, Geology, and Environmental Science (Committee Member) ACCOUNTING FOR INTRASPECIFIC VARIATION TRANSFORMS OUR UNDERSTANDING OF ARTIODACTYL SOCIAL EVOLUTION By Monica Irene Miles A Thesis Submitted to the Faculty of the University of Tennessee at Chattanooga in Partial Fulfillment of the Requirements of the Degree of Master of Science: Environmental Science The University of Tennessee at Chattanooga Chattanooga, Tennessee December 2018 ii Copyright © 2018 By Monica Irene Miles All Rights Reserved iii ABSTRACT A major goal in the study of mammalian social systems has been to explain evolutionary transitions in social traits. Recent comparative analyses have used phylogenetic reconstructions to determine the evolution of social traits but have omitted intraspecific variation in social organization (IVSO) and mating systems (IVMS). This study was designed to summarize the extent of IVSO and IVMS in Artiodactyla and Perissodactyla, and determine the ancestral social organization and mating system for Artiodactyla. Some 82% of artiodactyls showed IVSO, whereas 31% exhibited IVMS; 80% of perissodactyls had variable social organization and only one species showed IVMS. The ancestral population of Artiodactyla was predicted to have variable social organization (84%), rather than solitary or group-living. A clear ancestral mating system for Artiodactyla, however, could not be resolved. These results show that intraspecific variation is common in artiodactyls and perissodactyls, and suggest a variable ancestral social organization for Artiodactyla. iv ACKNOWLEDGEMENTS First and foremost, I would like to thank my incredibly dedicated advisor, Dr. Loren Hayes, for the unwavering support and guidance that he has provided throughout my graduate studies. I would also like to thank my committee members, Dr. Hope Klug and Dr. Tim Gaudin, for their invaluable insights. I am extremely thankful for my collaborators: Dr. Carsten Schradin, for his crucial advice during the conception and completion of this thesis; Dr. Marco Festa- Bianchet, for his knowledge on all things ungulate-related; and, Dr. Adrian Jaeggi, for his expertise and guidance with the phylogenetic work. Finally, I would like to give special thanks to my family, friends, and fellow graduate students who became the best support system one could ask for. v TABLE OF CONTENTS ABSTRACT ........................................................................................................................... iv ACKNOWLEDGEMENTS .................................................................................................... v LIST OF TABLES ................................................................................................................viii LIST OF FIGURES ................................................................................................................ix LIST OF ABBREVIATIONS ..................................................................................................x LIST OF TERMS ....................................................................................................................xi CHAPTER I. A REVIEW OF INTRASPECIFIC VARIATION IN ARTIODACTYLA AND PERISSODACTYLA SOCIAL SYSTEMS ................................................................1 Introduction ..................................................................................................................1 Social Systems .........................................................................................................1 Socioecological Theory ...........................................................................................3 Intraspecific Variation in Social Systems ...............................................................4 Artiodactyla and Perissodactyla ..............................................................................5 Thesis Objectives ....................................................................................................9 Artiodactyla Social Organization .................................................................................9 Bovidae .................................................................................................................13 Cervidae ................................................................................................................16 Suidae ....................................................................................................................19 Other Families ....................................................................................................... 21 Artiodactyla Mating Systems .....................................................................................24 Bovidae .................................................................................................................25 Cervidae ................................................................................................................28 Suidae ....................................................................................................................29 Other Families .......................................................................................................30 Summary of Artiodactyla Social Organization and Mating System ..........................31 Perissodactyla Social Organization ............................................................................32 Equidae .................................................................................................................33 Rhinocerotidae ......................................................................................................34 Tapiridae ...............................................................................................................34 vi Perissodactyla Mating System ...................................................................................35 Equidae .................................................................................................................35 Rhinocerotidae ......................................................................................................36 Summary of Perissodactyla Social Organization and Mating System .......................36 Discussion ..................................................................................................................36 II. ACCOUNTING FOR INTRASPECIFIC VARIATION TRANSFORMS UNDERSTANDING OF ARTIODACTYL SOCIAL EVOLUTION ......................39 Introduction ................................................................................................................39 Methods......................................................................................................................46 Social Organization and Mating System ..............................................................46 Ecology and Life History .....................................................................................48 Phylogeny ............................................................................................................48 Statistical Analysis ...............................................................................................49 Results ........................................................................................................................51 Occurrence of Intraspecific Variation in Social Organization .............................51 Phylogenetic Reconstruction of Ancestral Social Organization State and Factors Influencing IVSO .................................................................................51 Occurrence of Intraspecific Variation in Mating System ....................................58 Phylogenetic Reconstruction of Ancestral Mating System State and Factors Influencing IVMS .............................................................................................58 Discussion ..................................................................................................................61 Phylogenetic Reconstruction of Artiodactyla Social Organization and Mating System ..................................................................................................61 Ecological and Life History Predictors of IVSO and IVMS ...............................63 Limitations ...........................................................................................................65 Need for Long-term Studies and Manipulative Field Studies .............................66 Conservation Implications and Concluding Remarks ..........................................67 REFERENCES ......................................................................................................................69 APPENDIX A. SOCIAL ORGANIZATION DATA ..........................................................................85 B. MATING SYSTEM DATA .......................................................................................90 C. ECOLOGICAL AND LIFE HISTORY DATA ........................................................93 D. PHYLOGENETIC MODELS ....................................................................................98 VITA ....................................................................................................................................100
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