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LI-DISSERTATION-2015.Pdf Copyright by Zhiheng Li 2015 The Dissertation Committee for Zhiheng Li Certifies that this is the approved version of the following dissertation: Evolution of the hyoid apparatus in Archosauria: implications for the origin of avian tongue function Committee: Julia A. Clarke, Supervisor Timothy B. Rowe Christopher J. Bell Richard A. Ketcham Tobias Riede Zhonghe Zhou Evolution of the hyoid apparatus in Archosauria: implications for the origin of avian tongue function by Zhiheng Li, B.S.; M.Natural Sci. 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 August 2015 Dedication To my parents and sister, for their unconditional love and full support over my life. Acknowledgements The study here cannot be possible without the contributions from so many people along the past six years of my life. I would particularly like to thank my advisor, Julia Clarke for her inertial suggestion to work on the dinosaur hyoid as a potential dissertation topic; she provided consistent support in terms of countless inspiration, critical thinking and discussion, improvement of English, and funding sources in traveling and exploring experiments over my doctoral training at UT-Austin. It is also a great pleasure to be embraced by her great passion in sciences and acute curiosity on avian anatomy and systematics. I would also like to thank all my committee (Timothy Rowe, Christopher Bell, Richard Ketcham, Tobias Reide, and Zhonghe Zhou) for their valuable advice and important discussion throughout the development of this project and for so many other aspects of my graduate life in both United States and China. I would like to thank Helen James, Christopher Milensky, Jeremy Jacobs, Zhonghe Zhou, Xing Xu, Xiaoting Zheng and Paul Sweet for their assistance and help in accessing museum specimens (Smithsonian Institution, IVPP, Shandong Tianyu Museum, and American Museum of Natural History). I also thank Matthew Colbert and Jessie Maisano for their great work for scanning tens of birds and alligators used in this project. I thank staff from both UT VP-Lab (Matthew Brown, Chris Sagebiel, and Kenneth Bader) and TNHC (Travis Laduc and Adam Cohen) for their assistance in using the facilities for living specimens transportation, temporary storage, and processing. UT Paleo-group in particularly Kenneth Bader, Xia Wang, Michelle Stoker, Katie Browne, James Proffitt, Rachel Simon, and Lauren English provided discussion and assistance in improvement of my English writing. I would also thank Sterling Nesbitt, Tomazs v Owerkowicz, Edward Theriot, Tobias Riede, and Ruth Elsey, who donated numerous fresh birds and Alligator specimens used for dissection and CT scans. Nathan Jeffery and Brian Metscher are thanked for their suggestions in the guidelines of enhance-contrast CT protocols provided in early stage of the project. Numerous financial support are essential for the completion of this research from Jackson school of Geosciences (Off-campus Research Fund, Analytic Fund, and Carsey Fellowship in Paleontology ‏), Lundelius Fund, Graduate school, University of Texas at Austin (GS Named Continuing Fellowship) and Smithsonian Institution (a Pre-doctoral Fellowship). Lastly I would express my thanks to all faculty, staff and students from UT- Austin who made the place a nice and comfortable community for my journey to the west toward acquiring my Ph.D. degree. vi Evolution of the hyoid apparatus in Archosauria: implications for the origin of avian tongue function Zhiheng Li, Ph.D. The University of Texas at Austin, 2015 Supervisor: Julia A. Clarke Abstract: While certain aspects of hyolingual function have been intensively studied in extant archosaurs, including both the crocodylian- and bird-lineages, major aspects remain unstudied within their closely related extinct taxa, e.g., basal archosaurs and non- avian dinosaurs. In archosaurs, the hyoid apparatus and the associated larynx control the movement of the tongue and the throat, further affected several essential functions including drinking, food intake and manipulation, sound making, as well as panting and breathing. In addition to feeding, the hyolingual apparatus has also been identified as closely involved in sound production in crocodylians. However, neither the hyoid morphology nor the tongue function has been systematically investigated in extinct archosaurs (e.g., non-avian dinosaurs or other basal birds) because of the assumed limited evidence in the fossil records. So far, little is known about the ability of basal archosaur or non-avian dinosaur to produce sound, what their tongue looked like and how they used it. Therefore, the major goal of the dissertation is to integrate anatomical and functional data on the hyolingual apparatus in Crocodylia and Aves, combined with the most comprehensive assessment of hyoid fossils from extinct archosaurs, finally reconstruct vii the evolution of hyoid morphology and tongue function in Archosauria. The recoveries of several key specimens from this study filled the morphologic gap in the hyoid evolution from ancestral archosaurs to living birds. New techniques applied as the contrast- enhanced Computed Tomography Imaging combined with a large number of dissections supplement prolific details for inferring the soft-tissues of hyolingual apparatus within archosaurs. The synthesis of data on both bony hyoid and associated muscles illustrate the major pattern of the hyolingual evolution in dinosaur and shed new light into the origin of bird tongue and novel function. viii Table of Contents Chapter 1: Introduction .........................................................................................1 Chapter 2: New insight into the anatomy of the hyolingual apparatus of Alligator mississippiensis and implications for reconstructing feeding in extinct archosaurs .......................................................................................................4 Abstract ...........................................................................................................4 Introduction .....................................................................................................5 Materials and Methods ....................................................................................8 Terminology and Novelty ...............................................................................9 Results ...........................................................................................................11 Discussion .....................................................................................................19 Conclusions ...................................................................................................28 Institutional abbreviations .............................................................................29 Supplemental document ................................................................................29 Acknowledgments.........................................................................................30 Tables ............................................................................................................31 Figures...........................................................................................................35 Chapter 3: Craniolingual morphology of waterfowl (Aves, Anseriformes) and its relationship with feeding mode revealed through enhanced-contrast X-ray computed tomography and 2D morphometrics.............................................46 Abstract .........................................................................................................46 Background ...................................................................................................47 Materials and Methods ..................................................................................50 Results ...........................................................................................................52 Discussion .....................................................................................................59 Conclusions ...................................................................................................61 Acknowledgments.........................................................................................63 Tables ............................................................................................................64 ix Figures...........................................................................................................65 Chapter 4: An investigation of the efficacy and mechanism of enhanced contrast X- ray Computed Tomography utilizing iodine for large specimens through experimental and simulation approaches ......................................................74 Abstract .........................................................................................................74 Background ...................................................................................................75 Results ...........................................................................................................78 Discussion .....................................................................................................86 Materials and Methods ..................................................................................94 List of abbreviations ...................................................................................102
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