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Archosaur Hip Joint Anatomy and Its Significance in Body Size and Locomotor Evolution

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Archosaur Hip Joint Anatomy and Its Significance in Body Size and Locomotor Evolution ARCHOSAUR HIP JOINT ANATOMY AND ITS SIGNIFICANCE IN BODY SIZE AND LOCOMOTOR EVOLUTION HENRY P. TSAI JULY 2015 APPROVAL PAGE The undersigned, appointed by the dean of the Graduate School, have examined the dissertation entitled ARCHOSAUR HIP JOINT ANATOMY AND ITS SIGNIFICANCE IN BODY SIZE AND LOCOMOTOR EVOLUTION Presented by Henry Tsai, a candidate for the degree of doctor of philosophy, and hereby certify that, in their opinion, it is worthy of acceptance. Professor Casey Holliday Professor Carol Ward Professor Kevin Middleton Professor John Hutchinson Professor Libby Cowgill ACKNOWLEDGEMENTS I would like to acknowledge numerous individuals in aiding the completion of this project. I would like to thank my doctoral thesis committee: Casey Holliday, Carol Ward, Kevin Middleton, John Hutchinson, and Libby Cowgill, for their insightful comments and, as well as numerous suggestions throughout the course of this project. For access to specimens at their respective institution, I would like to thank Bill Mueller and Sankar Chatterjee (Museum of Texas Tech University), Gretchen Gürtler (Mesalands Community College's Dinosaur Museum), Alex Downs (Ruth Hall Museum of Paleontology), William Parker (Paleontological Collection at the Petrified Forest National Park), Robert McCord (Arizona Museum of Natural History), David and Janet Gillette (Museum of Northern Arizona), Kevin Padian (University of California Museum of Paleontology), Joseph Sertich and Logan Ivy (Denver Museum of Nature and Science), Peter Makovicky, William Simpson, and Alan Resetar (Field Museum of Natural History), Scott Williams (Burpee Museum of Natural History), Ruth Elsey (Rockefeller Wildlife Refuge), Yen-nien Cheng (National Museum of Natural Science, Taiwan), Randall Irmis and Carolyn Levitt (Natural History Museum of Utah), Daniel Chure (Dinosaur National Monument), Steve Sroka (Utah Field House of Natural History), Brooks Britt and Rodney Scheetz (Brigham Young University Museum of Paleontology), Luis Chiappe, Maureen Walsh, Gregory Pauly, Neftali Camacho, Kenneth Campbell, and Kimball Garrett (Los Angeles County Museum of Natural History), Oliver Rauhut (Paläontologisches Museum München), Rainer Schoch (Staatliches Museum für Naturkunde Stuttgart), Daniela Schwarz-Wings (Museum für Naturkunde Berlin), Matthew Brown and J. Christopher Sagebiel (Texas Memorial Museum), Daniel ii Brinkman (Yale Peabody Museum), Jessica Cundiff (Museum of Comparative Zoology, Harvard University), Matthew Lamanna and Amy Henrici (Carnegie Museum of Natural History), Diego Pol, José Luis Carballido and Eduardo Ruigómez (Museo Paleontológico Egidio Feruglio), Julia Desojo and Stella Alvarez (Museo Argentino de Ciencias Naturales). Travel funds for museum and institutional visits were provided by the Jurassic Foundation Research Grant, the Doris O. and Samuel P. Welles Research Fund, as well as the University of Missouri Life Science Travel Grant. Additionally, I would like to thank the following individuals for providing digital- and physical specimens used in this project. I thank Heinrich Mallison for providing CT- scan data of Plateosaurus, Bill Parker for providing laser scan data for Coelophysis, Sterling Nesbitt for providing the skeletal cast of Asilisaurus, John Hutchinson for sharing 3D scan data for numerous theropods, Ruth Elsey for providing the ontogenetic sample of Alligator, and Alan Resetar for allowing me the opportunity to dissect a Sphenodon specimen in the Field Museum’s herpetological collection. Discussions with the following individuals contributed to ideas and directions of this project: Ashley Hammond, Kaleb Sellers, Sarah Werning, Rachel Menegaz, Tomasz Owerkowicz, Victoria Ngo, Adam Summers, and Heinrich Mallison. I would like to thank Jimmy C. Lattimer (MU Veterinary Medical Diagnostic Laboratory) and Ashley Szczodroski (MU Veteran’s Hospital Biomolecular Imaging Center) for providing computed tomography services for wet specimens, as well as Kevin Middleton and Jill Harper-Judd for providing extensive help on the phylogenetic comparative analyses part of this project. I thank my cohort of MU Integrative Anatomy students for their constant moral support, as iii well as Johnny Yang, Barbara Williams, and Noelle Nelson for graciously providing me with boarding accommodations and work space during my collection visits. Lastly, I would like to thank my family for continued moral support during my tenure as a graduate student. In particular, I would like to recognize my parents, Steve and Debby Tsai, as well as my grandfather Tony Wu, for fostering my interest in biology and vertebrate paleontology ever since my kindergarten years, and continue to support it as my chosen field of study. This work would not have been possible without their unwavering encouragements. iv Table of Contents LIST OF FIGURES AND ILLUSTRATIONS.................................................................. vi LIST OF TABLES ........................................................................................................... viii ABSTRACT ........................................................................................................................ 1 CHAPTER 1: Introduction: The evolving view of saurischian locomotion ....................... 2 CHAPTER 2: ................................................................................................................... 15 INTRODUCTION ......................................................................................................... 15 MATERIALS AND METHODS .................................................................................. 19 RESULTS...................................................................................................................... 24 DISCUSSION ............................................................................................................... 36 CONCLUSION ............................................................................................................. 52 CHAPTER 3: Articular soft tissue anatomy of the archosaur hip joint: Structural homology and functional implications .............................................................................. 79 INTRODUCTION ......................................................................................................... 79 MATERIALS AND METHODS .................................................................................. 82 RESULTS...................................................................................................................... 86 DISCUSSION ............................................................................................................. 114 CONCLUSION ........................................................................................................... 137 CHAPTER 4: Convergence and disparity in saurischian dinosaur hip joints associated with gigantism ................................................................................................................. 159 INTRODUCTION ....................................................................................................... 159 MATERIALS AND METHODS ................................................................................ 162 RESULTS.................................................................................................................... 171 DISCUSSION ............................................................................................................. 175 CONCLUSION ........................................................................................................... 187 CHAPTER 5: Summary .................................................................................................. 212 LITERATURES CITED ................................................................................................. 215 VITA ............................................................................................................................... 239 v LIST OF FIGURES AND ILLUSTRATIONS Figure 1-1. Simplified phylogeny of Saurischia and its successive diapsid outgroups. Figure 2-1. Evolutionary relationships of the tetrapod epiphyseal region and their anatomical nomenclature. Figure 2-2. Orthogonal reference planes used to describe anatomical structures on the proximal femur of extant archosaurs. Figure 2-3. Schematic representation of hip joint tissues homology in Diapsida. Figure 2-4. Gross morphology of lepidosaur acetabular soft tissues. Figure 2-5. Microstructure of lizard supraacetabular tissues. Figure 2-6. Gross morphology of the crocodylian acetabulum and microstructure of the acetabular labrum. Figure 2-7. Morphology of the Alligator antitrochanter region. Figure 2-8. Gross morphology and microstructure of avian acetabular soft tissues. Figure 2-9. Gross morphology and microstructure of the lizard proximal femoral epiphyseal region. Figure 2-10. Gross morphology and microstructure of the crocodylian proximal femoral epiphyseal region. Figure 2-11. Gross morphology and microstructure of the avian proximal femoral epiphyseal region. Figure 2-12. Irregular rugosities on the growth plate surface (green arrow) are the osteological correlates of thick hyaline cartilage. Figure 2-13. Evolutionary history of the hip joint characters in sauropsids, with emphasis on archosaurs. Figure 3-1. Orthogonal reference planes used to describe articular structures on the proximal femur of Tyrannosaurus, and exemplary theropod. Tissues nomenclature and color schemes are labeled according
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