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Paleobiogeography of Miocene to Pliocene Equinae of North America: a Phylogenetic

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Paleobiogeography of Miocene to Pliocene Equinae of North America: a Phylogenetic Paleobiogeography of Miocene to Pliocene Equinae of North America: A Phylogenetic Biogeographic and Niche Modeling Approach A thesis presented to the faculty of the College of Arts and Sciences of Ohio University In partial fulfillment of the requirements for the degree Master of Science Kaitlin Clare Maguire June 2008 2 This thesis titled Paleobiogeography of Miocene to Pliocene Equinae of North America: A Phylogenetic Biogeographic and Niche Modeling Approach by KAITLIN CLARE MAGUIRE has been approved for the Department of Geological Sciences and the College of Arts and Sciences by Alycia L. Stigall Assistant Professor of Geological Sciences Benjamin M. Ogles Dean, College of Arts and Sciences 3 ABSTRACT MAGUIRE, KAITLIN CLARE, M.S., June 2008, Geological Sciences Paleobiogeography of Miocene to Pliocene Equinae of North America: A Phylogenetic Biogeographic and Niche Modeling Approach (195 pp.) Director of Thesis: Alycia L. Stigall The biogeography and evolution of the subfamily Equinae is examined using two separate but related analyses, phylogenetic biogeography and ecological niche modeling. The evolution of Equinae is a classic example of an adaptive radiation during a time of environmental change. Both analyses employed here examine the biogeography of the equine species to interpret how environmental and historical variables led to the rise and fall of this clade. Results determine climate change is the primary factor driving the radiation of Equinae and geodispersal is the dominant mode of speciation between regions of North America. A case study in the Great Plains indicates distributional patterns are more patchy during the middle Miocene when speciation rates are high than in the late Miocene, when the clade is in decline. Statistical results and distributional patterns show equine species tracked their preferred habitat throughout North America as climate changed in the Miocene. Approved: _____________________________________________________________ Alycia L. Stigall Assistant Professor of Geological Sciences 4 ACKNOWLEDGMENTS The completion of this thesis would not have been possible without the help and guidance of several people. I would like to thank the following people for providing data: B.H. Passey, J.D. Damuth, J.R. Thomasson and a special thanks to G.J. Retallack. I would also like to thank C.M. Janis and Y. Wang for pointing me in the right direction for data, R.C. Hulbert and J. Alroy for clarification with phylogenetic relationships in the clade, and R. Purdy for access to the collections at NMNH. A big thanks to Chris Dobel for assisting me with data collection. This research was funded by an Ohio University Geological Sciences Graduate Alumni Research Grant, GSA Grants-in-Aid Award, and OHIO Center for Ecology and Evolutionary Studies Fellowship. I would especially like to thank my advisor, Alycia Stigall for all of her help, guidance and support that has not only made me a better student, but a better professional. Thank you also to my committee members, Dan Hembree, Pat O’Connor and Keith Milam for reviewing parts of this thesis intended for publication and for all of their help and support along the way. 5 TABLE OF CONTENTS Page Abstract............................................................................................................................... 3 Acknowledgments............................................................................................................... 4 List of Tables ...................................................................................................................... 8 List of Figures..................................................................................................................... 9 Chapter 1: Introduction..................................................................................................... 10 References..................................................................................................................... 14 Chapter 2: Paleobiogeography of Miocene Equinae of North America: A phylogenetic biogeographic analysis of the relative roles of climate, vicariance, and dispersal ........... 16 Abstract......................................................................................................................... 16 Introduction................................................................................................................... 17 Geologic and Paleoclimatic Framework....................................................................... 21 Evolutionary Framework .............................................................................................. 23 Materials and Methods.................................................................................................. 26 Taxa and geographic regions ................................................................................... 26 Analytical Biogeographic Method ............................................................................ 28 Results........................................................................................................................... 30 Speciation Patterns ................................................................................................... 30 Biogeographic Area Analysis ................................................................................... 31 Discussion..................................................................................................................... 33 Distributional Patterns ............................................................................................. 33 6 Vicariance Patterns .................................................................................................. 37 Geodispersal Patterns............................................................................................... 40 Comparison and Synthesis........................................................................................ 42 Conclusions................................................................................................................... 43 References..................................................................................................................... 45 CHAPTER 3: Distribution of fossil horses in the Great Plains during the Miocene and Pliocene: An ecological niche modeling approach........................................................... 52 Abstract......................................................................................................................... 52 Introduction................................................................................................................... 53 Methods ........................................................................................................................ 57 Geographic and Stratigraphic Intervals................................................................... 57 Species Occurrence Information............................................................................... 63 Environmental Data.................................................................................................. 64 Creation of Environmental Layers............................................................................ 72 Distribution Modeling............................................................................................... 73 Biogeographic Analyses............................................................................................ 77 Results and Discussion ................................................................................................. 81 Habitat Fragmentation ............................................................................................. 81 Habitat Tracking....................................................................................................... 83 Range Size vs. Survival ............................................................................................. 89 Regional Trends ........................................................................................................ 92 Conclusions................................................................................................................... 95 7 References..................................................................................................................... 97 Chapter 4: Conclusion..................................................................................................... 106 References................................................................................................................... 109 APPENDIX A: Vicariance and geodispersal matrix ...................................................... 110 APPENDIX B: Published references for geographic location data................................ 112 APPENDIX C: Species occurrence data for the great plains ......................................... 124 APPENDIX D: Original environmental data.................................................................. 143 APPENDIX E: Predicted species distribution maps....................................................... 186 8 LIST OF TABLES Page Table 1.Species modeled in the middle and late time slices............................................. 58 Table 2.Environmental data refereneces by variable........................................................ 59 Table 3.Environmental data for each grid box in the middle time slice........................... 69 Table 4.Environmental data for each grid
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