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The Evolution of Occlusal Enamel Complexity in Middle THE EVOLUTION OF OCCLUSAL ENAMEL COMPLEXITY IN MIDDLE MIOCENE TO RECENT EQUIDS (MAMMALIA: PERISSODACTYLA) OF NORTH AMERICA by NICHOLAS ANTHONY FAMOSO A THESIS Presented to the Department of Geological Sciences and the Graduate School of the University of Oregon in partial fulfillment of the requirements for the degree of Master of Science June 2013 THESIS APPROVAL PAGE Student: Nicholas Anthony Famoso Title: The Evolution of Occlusal Enamel Complexity in Middle Miocene to Recent Equids (Mammalia: Perissodactyla) of North America This thesis has been accepted and approved in partial fulfillment of the requirements for the Master of Science degree in the Department of Geological Sciences by: Dr. Edward Davis Chair Dr. Qusheng Jin Member Dr. Stephen Frost Outside Member and Kimberly Andrews Espy Vice President for Research & Innovation/Dean of the Graduate School Original approval signatures are on file with the University of Oregon Graduate School. Degree awarded June 2013 ii © 2013 Nicholas Anthony Famoso iii THESIS ABSTRACT Nicholas Anthony Famoso Master of Science Department of Geological Sciences June 2013 Title: The Evolution of Occlusal Enamel Complexity in Middle Miocene to Recent Equids (Mammalia: Perissodactyla) of North America Four groups of equids, “Anchitheriinae,” Merychippine-grade Equinae, Hipparionini, and Equini, coexisted in the middle Miocene, and only the Equini remains after 16 million years of evolution and extinction. Each group is distinct in its occlusal enamel pattern. These patterns have been compared qualitatively but rarely quantitatively. The processes controlling the evolution of these occlusal patterns have not been thoroughly investigated with respect to phylogeny, tooth position, and climate through geologic time. I investigated two methods of quantitative analysis, Occlusal Enamel Index for shape and fractal dimensionality for complexity. I used analyses of variance and an analysis of co-variance to test hypotheses of process. Results suggest that enamel shape was controlled by phylogeny, tooth position, and climate. The lower taxonomic levels are shown to have a strong effect on complexity, suggesting behavior is driving complexity rather than overarching phylogenetic constraint. iv CURRICULUM VITAE NAME OF AUTHOR: Nicholas Anthony Famoso GRADUATE AND UNDERGRADUATE SCHOOLS ATTENDED University of Oregon, Eugene, OR South Dakota School of Mines and Technology, Rapid City, SD DEGREES AWARDED: Master of Science, Geological Sciences, 2013, University of Oregon Bachelor of Science, Geology, 2009, South Dakota School of Mines and Technology AREAS OF SPECIAL INTEREST: Vertebrate Paleontology Biostratigraphy Conservation Paleobiology Phylogeny of the Equidae PROFESSIONAL EXPERIENCE: Graduate Teaching Fellow, Geological Sciences, Eugene, OR, 2011 to present Baldwin Museum Curation Research Fellow, Museum of Natural and Cultural History, Eugene, OR, 2011 - 2013 Assistant Paleontology Lab Manager, Badlands National Park, Interior, SD, 2012 Highway Salvage Preparator, University of Nebraska State Museum, Lincoln, NE, 2009 - 2011 Curation Assistant, South Dakota State Archaeological Research Center, Rapid City, SD, 2007 - 2009 Museum/Curation Assistant, Museum of Geology, Rapid City, SD, 2005 - 2009 Student Assistant, Ashfall Fossil Beds State Historical Park, Royal, NE, 2008 v David B. Jones Intern, Ashfall Fossil Beds State Historical Park, Royal, NE, 2007 GRANTS, AWARDS, AND HONORS: Honorable Mention, National Science Foundation, Graduate Research Fellowship Program, 2013 Johnston Scholarship, University of Oregon, Department of Geological Sciences, 2012 Clarence and Lucille Dunbar Scholarship, University of Oregon, College of Arts and Sciences, 2012 Richard K. Bambach Award, Paleontological Society, 2012 Graduate Student Research Grant, Geological Society of America, 2012 Magna cum Laude, South Dakota School of Mines and Technology, 2009 Dean’s List, South Dakota School of Mines and Technology, 2006 - 2009 Society of Vertebrate Paleontology Student Member Travel Grant, Jackson School of Geosciences, 2008 PUBLICATIONS: Famoso N. A., Feranec, R. S., Davis, E. B., in revision. The relationship between lophodonty, hypsodonty, body mass and diet in extinct and extant ungulates. Palaeogeography, Palaeoclimatology, Palaeoecology. Famoso, N.A., Pagnac, D. C., 2011. A Comparison of the Clarendonian Equid Assemblages from the Mission Pit, South Dakota and Ashfall Fossil Beds, Nebraska. Transactions of the Nebraska Academy of Sciences and Affiliated Societies 32, 98-107. vi ACKNOWLEDGMENTS I would like to thank many people for their assistance, support, and cultivation of this project. This was a project that I became interested in while in my undergraduate at the South Dakota School of Mines working with Dr. Darrin Pagnac. After discussions with Dr. Samantha Hopkins and my current advisor Dr. Edward Davis I was pointed in the direction of Dr. Robert Feranec whose help was invaluable to this study. I would like to specifically thank Dr. Feranec for providing part of my data set. I would also like to specifically thank R. G. Corner, G. Brown, R. Secord, R. Skolnick, S. Tucker, R. Otto, G. Boardman, P. Holroyd, J. Frankel, C. Sidor, B. Eng, J. and B. Orcutt, D. Pagnac, S. Shelton, J. and H. Carr, J. Samuals, C. Schierup, R. Evander, J. Galkin, J. Meng, S. K. Lyons, C. Ito, P. Wagner, and E. Scott for access to collections, hospitality, and important discussions. I would also like to thank Q. Jin and S. Frost, the members of my committee, for their feedback and suggestions on this project. The members of the UO Paleontology Group (past and present) have also contributed constructive feedback, advice, and hospitality related to this project. Funding for this project was provided by the University of Oregon Museum of Natural and Cultural History, South Dakota School of Mines and Technology Department of Geology and Geological Engineering, Paleontological Society Richard K. Bambach Award, and Geological Society of America Graduate Student Grant. Data collected by Dr. Feranec was funded by the Museum of Vertebrate Zoology, University of California Museum of Paleontology, University of California, Berkeley Department of Integrative Biology, the Society of Vertebrate Paleontology, and NSF DDIG 0308770. vii Personally I would like to thank my family in Oregon and New York for their continued support and hospitality. Thanks are extended to my non-paleontology graduate student friends, specifically H. Diettreich, J. K. Marks, S. Mordensky, R. Tuohy, and M. Haskin for keeping me grounded and not allowing me to get lost in my studies. And a special thank you to K. MacKenzie, my officemate. There are countless other people who I have not mentioned who have enriched not only my graduate experience, but this project. viii TABLE OF CONTENTS Chapter Page I. INTRODUCTION ................................................................................................... 1 Evolutionary Context ......................................................................................... 2 Measures of Complexity .................................................................................... 5 Geography .......................................................................................................... 8 Tooth Position .................................................................................................... 9 Questions............................................................................................................ 10 Hypotheses ......................................................................................................... 11 Predictions.......................................................................................................... 11 II. METHODS .............................................................................................................. 13 Institutional Abbreviations................................................................................. 14 Occlusal Enamel Index, OEI.............................................................................. 15 Fractal Dimensionality, D .................................................................................. 21 III. RESULTS ................................................................................................................ 23 Occlusal Enamel Index, OEI.............................................................................. 23 Tooth Position .............................................................................................. 23 Nested Multi-way Analysis of Variance ...................................................... 23 ANOVA 1: OEI vs. Region within Each NALMA ..................................... 24 ANOVA 2: OEI vs. Tribe ............................................................................ 26 ANOVA 3: OEI vs. NALMA within Each Region ..................................... 27 ix Chapter Page ANOVA 4: OEI vs. Tribe within Each NALMA ........................................ 28 ANOVA 5: OEI vs. Tribe within Each Region ........................................... 29 Fractal Dimensionality, D .................................................................................. 31 IV. DISCUSSION .......................................................................................................... 32 Occlusal Enamel Index, OEI.............................................................................. 32 Fractal Dimensionality, D .................................................................................
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