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The Function and Evolution of the Syncervical in Ceratopsian Dinosaurs with a Review of Cervical Fusion in Tetrapods

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The Function and Evolution of the Syncervical in Ceratopsian Dinosaurs with a Review of Cervical Fusion in Tetrapods The Function and Evolution of the Syncervical in Ceratopsian Dinosaurs with a Review of Cervical Fusion in Tetrapods by Collin S. VanBuren A thesis submitted in conformity with the requirements for the degree of Master’s of Science Ecology & Evolutionary Biology University of Toronto © Copyright by Collin S. VanBuren 2013 The Function and Evolution of the Syncervical in Ceratopsian Dinosaurs with a Review of Cervical Fusion in Tetrapods Collin S. VanBuren Master’s of Science Ecology & Evolutionary Biology University of Toronto 2013 Abstract Mobility of the vertebral column is important for many ecological aspects of vertebrates, especially in the cervical series, which connects the head to the main body. Thus, fusion within the cervical series is hypothesized to have ecological and behavioural implications. Fused, anterior cervical vertebrae have evolved independently over 20 times in ecologically disparate amniotes, most commonly in pelagic, ricochetal, and fossorial taxa, suggesting an adaptive function for the ‘syncervical.’ Fusion may help increase out-force during head-lift digging or prevent anteroposteriorly shortened vertebrae from mechanically failing during locomotion, but no hypothesis for syncervical function has been tested. The syncervical of neoceratopsian dinosaurs is hypothesized to support large heads or aid in intraspecific combat. Tests of correlated character evolution within a ceratopsian phylogeny falsify these hypotheses, as the syncervical evolves before large heads and cranial weaponry. Alternative functional hypotheses may involve ancestral burrowing behaviour or unique feeding ecology in early neoceratopsians. ii Acknowledgments I thank David C. Evans, Hernán López-Fernández, and Gerry De Iuliis for their supervision during the course of this work, and Mary Silcox and Deborah McLennan for serving on my examination committee (at 9am on a Friday, no less). This project was inspired initially by Darren Tanke and then progressed by Nicolás Campione and David Evans, which set the stage for my M.Sc. research. For access to specimens housed at their respective institutions and assistance in osteology and palaeontology collections, I thank Judy Galkin, Carl Mehling, Mark Norell, Eileen Westwig (American Museum of Natural History, New York, NY), Kieran Shepard (Canadian Museum of Nature, Ottawa, ON), Jack Horner (Museum of the Rockies, Bozeman, MT), Kevin Seymour (Royal Ontario Museum, Toronto, ON), Brandon Strilisky (Royal Tyrrell Museum of Paleontology, Drumheller, AB), Michael Brett-Surman, Darrin Lundy, John Ososky, and Charley Potter (National Museum of Natural History, Washington, D. C.). My supervisor, whose guidance helped develop my project and my future research direction in so many ways, graciously provided funding and support. Thanks for investing in me, David. For help with making my figures more artistic, putting up with me when I bombarded him with questions while he was busy working on his side-projects and thesis, and being a constant enabler of “Chocolate PM,” I thank my office-mate Caleb Brown. Because all work and no play creates an insane graduate student, I am grateful to Kirstin Brink, Sarah Steele, Sophia Lavergne, and Alix Cameron for all the brunches, cooking adventures, and glasses of wine during my twenty months in Toronto. I would also like to thank all of my lab mates, Kirstin Brink, Caleb Brown, Nicolás Campione, Thomas Cullen, and Derek Larson, as well as Kentaro Chiba, Lorna O’Brien, and the López-Fernández lab for their help and support. A special thanks to Nicolás Campione for helping with analyses, the many distracting discussions of science, being my TA ‘partner in crime’ for multiple courses, and overall serving as an unrecognized co-supervisor in so many iii respects. Finally, a big “Thank you” to all of my friends and family back home for believing in me and being there for me, even from a different country. iv Table of Contents ACKNOWLEDGMENTS.......................................................................................................................................III TABLE OF CONTENTS..........................................................................................................................................V LIST OF TABLES .................................................................................................................................................VII LIST OF FIGURES.............................................................................................................................................. VIII LIST OF APPENDICES...........................................................................................................................................X CHAPTER 1 CONVERGENCE AND FUNCTION OF THE TETRAPOD SYNCERVICAL............................1 1.1 ABSTRACT........................................................................................................................................................................1 1.2 INTRODUCTION...............................................................................................................................................................2 1.3 MATERIALS AND METHODS .........................................................................................................................................3 1.4 SYSTEMATIC REVIEW ....................................................................................................................................................5 1.4.1 Mammalia.................................................................................................................................................................5 1.4.2 Reptilia .................................................................................................................................................................... 14 1.5 GENERAL CHARACTERISTICS OF FUSED CERVICAL VERTEBRAE........................................................................18 1.6 WHAT IS THE FUNCTION OF THE SYNCERVICAL?...................................................................................................19 1.6.1 Function of the syncervical in fossorial taxa .......................................................................................... 20 1.6.2 Function of the syncervical in ricochetal taxa ....................................................................................... 21 1.6.3 Function of the syncervical in cetaceans.................................................................................................. 22 1.6.4 Function of the syncervical in hornbills.................................................................................................... 24 1.6.5 The mysterious syncervical of porcupines and pacaranas............................................................... 24 1.6.6 Interpreting the syncervical as an adaptation ...................................................................................... 25 1.7 IMPLICATIONS FOR FOSSIL TAXA ..............................................................................................................................26 1.8 CONCLUSIONS ..............................................................................................................................................................28 REFERENCES ....................................................................................................................................................... 30 CHAPTER 2 HEAD SIZE, WEAPONRY, AND CERVICAL ADAPTATION: TESTING CRANIOCERVICAL EVOLUTIONARY HYPOTHESES IN CERATOPSIA ................................................. 61 2.1 ABSTRACT.....................................................................................................................................................................61 2.2 INTRODUCTION............................................................................................................................................................61 2.3 MATERIALS AND METHODS ......................................................................................................................................64 2.3.1 Syncervical Specimens and Measurements ............................................................................................. 64 v 2.3.2 Construction of Phylogeny.............................................................................................................................. 65 2.3.3 Quantifying Relative Skull Size..................................................................................................................... 66 2.3.4 Phylogenetic Analyses....................................................................................................................................... 68 2.4 DESCRIPTION OF UNFUSED SYNCERVICALS ...........................................................................................................70 2.4.1 Atlas.......................................................................................................................................................................... 70 2.4.2 Unfused C3 ............................................................................................................................................................. 71 2.5 RESULTS........................................................................................................................................................................72 2.5.1 Quantifying Relative Skull Size....................................................................................................................
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