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Boyd Dissertation 2012212.Pdf Copyright by Clint Aaroen Boyd 2012 The Dissertation Committee for Clint Aaroen Boyd Certifies that this is the approved version of the following dissertation: TAXONOMIC REVISION OF LATEST CRETACEOUS NORTH AMERICAN BASAL NEORNITHISCHIAN TAXA AND A PHYLOGENETIC ANALYSIS OF BASAL ORNITHISCHIAN RELATIONSHIPS Committee: Julia A. Clarke, Supervisor Christopher J. Bell Timothy B. Rowe David C. Cannatella Peter J. Makovicky TAXONOMIC REVISION OF LATEST CRETACEOUS NORTH AMERICAN BASAL NEORNITHISCHIAN TAXA AND A PHYLOGENETIC ANALYSIS OF BASAL ORNITHISCHIAN RELATIONSHIPS by Clint Aaroen Boyd, B.S. Dissertation Presented to the Faculty of the Graduate School of The University of Texas at Austin in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy The University of Texas at Austin May 2012 Dedication To my grandfather, James Boyd, for teaching me how to hold history in my hands and be filled with its wonder. Acknowledgements Thank you first and foremost to my advisor, Julia Clarke, for giving me the opportunity to pursue a career in the field that I enjoy so much and for dedicating so much of her time and effort towards my professional development. The remaining members of my committee, C. Bell, D. Cannatella, P. Makovicky, and T. Rowe, provided me with valuable guidance during the course of my graduate studies that greatly improved the quality of my research. I would also like to thank the former members of my committee at North Carolina State University (J. Hibbard, M. Schweitzer, and B. Wiegmann) for their guidance and efforts on my behalf during my time at NCSU, which was very much appreciated. Access to collections and specimens was generously provided by C. Mehling and M. Norell (AMNH), X. Xing (IVPP), J. Horner and B. Baziak (MOR), V. Schneider and D. Russell (NCSM), K. Q. Gao (Peking University), S. Shelton and M. Greenwald (SDSM), J. Nelson (TLAM), M. Carrano and M. Brett-Surman (USNM), S. Sampson, M. Getty, and R. Irmis (UMNH), and D. Brinkman and C. Norris (YPM). P. Brinkman (NCSM) collaborated in the preparation of NCSM 15728 and M. Brown (VPL) provided instruction in the preparation of several new specimens of Scutellosaurus. D. Evans (ROM) provided information on the anatomy of ROM 804. D.A. Winkler provided anatomical information about the ‘Proctor Lake ornithopod.’ D. Varricchio provided additional information on the anatomy of MOR 1636a. C.M. Brown provided photographs of several specimens referred to Thescelosaurus, information regarding v orodromine material from the Dinosaur Park Formation, and details regarding the anatomy of Parksosaurus. P. M. Galton provided encouragement and enlightening conversations over the years regarding basal ornithischian taxa. T. P. Cleland provided select photographs of NCSM 15728 and USNM 7758. S. Nesbitt provided access to photographs of several basal ornithischian taxa from South Africa. E. L. Schmidt provided illustrations of NCSM 15728 and USNM 7758. I am indebted to S. Masters for bringing the basal neornithischian material from the Kaiparowits Formation of Utah to my attention and A. Titus of the Utah Bureau of Land Management for assistance in locating additional specimens referable to this new taxon. A. DeBee and S. Nesbitt assisted in identifying material from the holotype of Skaladromeus goldenii gen. et sp. nov. and A.D.B. Behlke assisted in constructing the name of this taxon. T. P. Cleland and N. Marrero assisted in the design of the ASCC program suite and E.B. Cleland, N. Marrero, and S.J. Nesbitt helped troubleshoot early versions of the ASCC program suite. A. Molineux provided generous access to the photographic equipment at the Nonvertebrate Paleontology Laboratory at the University of Texas at Austin. The Willi Hennig Society provided free access to the phylogenetic program TNT. The Polyglot Paleontologist website provided access to multiple translations of important scientific research papers on ornithischian dinosaurs. This research was funded via the American Museum of Natural History Collection Study Grant, an Ernest L. and Judith W. Lundelius Scholarship in Vertebrate Paleontology, a Francis L. Whitney Endowed Presidential Scholarship from the University of Texas, a Geological Society of America Graduate Student Research Grant, vi the National Science Foundation’s East Asia and Pacific Summer Institutes for U.S. Graduates Students program, the North Carolina Fossil Club, and financial support from the Jackson School of Geosciences at the University of Texas at Austin and the Department of Marine, Earth, and Atmospheric Sciences at North Carolina State University. A. D. B. Behlke, T. P. Cleland, A. DeBee, D. R. Eddy, T. Gates, J. L. Green, M. L. Householder, J. Hutchinson, E. A. Johnson, D. Ksepka, K. Lamm, S. Nesbitt, K. Pickett, D. Pol, R. Scheetz, N. A. Smith, and H. D. Sues provided thoughtful comments on previous drafts of this dissertation that greatly improved its quality. I thank my father, Gerald Kenneth, Boyd, for constantly reminding me that I can learn more by going out and experiencing the world around me than I can by reading about it, and for cultivating my love for the outdoors. My deepest thanks to my mother, Coralee Boyd, for always being the friend I needed at all the right moments. Thank you to my children, Alexis Marie Boyd and William Shaw Boyd, for brightening all of my days and constantly reminding me what is truly important in this world. Special thanks to Mindy Lynn Householder, my love, for being the steady hand I needed at the wheel when I found myself lost in the stormy seas of life. vii Preface This dissertation combines a variety of data and analytical methods to explore the taxonomy, systematics relationships, and biogeography of basal ornithischian dinosaurs (Archosauromorpha, Archosauria). Because the chapters following the introduction are formatted to be published as independent works, there is some repetition with respect to introductory statements and materials and methods sections. Although figures and tables are imbedded throughout the text of the dissertation, the reader is frequently referred to a wealth of information contained in the appendices that follow directly after the text of the final chapter. References included in each chapter and the appendices are combined into a single references section found at the end of the dissertation. All photographs were taken by the author unless otherwise noted. viii TAXONOMIC REVISION OF LATEST CRETACEOUS NORTH AMERICAN BASAL NEORNITHISCHIAN TAXA AND A PHYLOGENETIC ANALYSIS OF BASAL ORNITHISCHIAN RELATIONSHIPS Clint Aaroen Boyd, PhD The University of Texas at Austin, 2012 Supervisor: Julia A. Clarke The systematic relationships of basal ornithischian dinosaurs remain contentious, especially the position of basal neornithischians (i.e., ‘hypsilophodontids’). Prior analyses of basal ornithischian relationships have been hampered by the fact that the hypodigm material of many basal neornithischian taxa is fragmentary, denying access to character data crucial to resolving their relationships. The recent discovery of several new basal neornithischian taxa and the referral of more complete specimens to known taxa provide important new data pertinent to resolving these relationships. The results of this study supplement those recent advances by improving our understanding of the anatomy and systematic relationships of basal neornithischian taxa from the Late Cretaceous of North America. These new insights are accomplished through a taxonomic revision of the Maastrichtian taxa Bugenasaura and Thescelosaurus, a detailed anatomical description of the cranial anatomy of Thescelosaurus neglectus based on the referral of a specimen that includes a nearly complete skull (NCSM 15728), and description of a new basal ix neornithischian taxon from the Kaiparowits Formation (Campanian) of Utah. All of these new data are compiled into a dataset composed of 255 characters for 65 terminal taxa (all species exemplars) focused on assessing basal ornithischian relationships. The recovered strict consensus topology is the most highly resolved, stratigraphically congruent phylogenetic hypothesis of basal ornithischian relationships yet proposed. This analysis places all basal neornithischians except Hypsilophodon foxii outside of Cerapoda, substantially reducing the taxonomic contents of Ornithopoda. A new clade containing fourteen basal neornithischian taxa is recovered as the sister taxon to Cerapoda and includes all North American basal neornithischians from the Cretaceous. The historical biogeography of Ornithischia is also reconstructed using a method that incorporates time calibrated branch lengths that represent the implied missing fossil record of each taxon. The results of this analysis support two dispersals of neornithischian taxa into South America during the Cretaceous: one consisting of basal iguanodontians dispersing from Australia (possibly via Antarctica) and a second consisting of basal neornithischians dispersing from Asia through North America. x Table of Contents INTRODUCTION ...................................................................................................1 CHAPTER 1. Reassesing the Taxonomic Status of the Basal Neornithischian Taxa Thescelosaurus and Bugenasaura.........................................................9 INTRODUCTION ........................................................................................10 CLADISTIC METHODOLOGY..................................................................15 SYSTEMATIC PALEONTOLOGY
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