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Morphological and Biomechanical Predictions of Cranial Kinesis in Reptile Evolution

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Morphological and Biomechanical Predictions of Cranial Kinesis in Reptile Evolution Morphological and Biomechanical Predictions of Cranial Kinesis in Reptile Evolution IAN N. COST July 2019 APPROVAL PAGE The undersigned, appointed by the dean of the Graduate School, have examined the dissertation entitled Morphological and Biomechanical Predictions of Cranial Kinesis in Reptile Evolution Presented by Ian N. Cost, 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 Kristina Aldridge _______________________________________ Professor Kevin Middleton _______________________________________ Professor Manuel Leal ACKNOWLEDGMENTS I am grateful for the assistance of numerous individuals and institutions for aiding in the completion of this project. I will begin by expressing my sincerest thanks to my doctoral dissertation committee who helped to develop and guide this project: Casey Holliday, Kevin Middleton, Kristina Aldridge, and Manuel Leal. Many institutions and individuals helped in procuring specimens and data for this project. I am extremely grateful to the following individuals at loaning and scanning institutions I worked with: Peter Larson (Black Hills Institute), Art Anderson (Virtual Surfaces), Larry Witmer (Ohio University), Ben Marks (Field Museum of Natural History), Zachary Morris and Jeremiah Trimble (Harvard Center for Nanoscale Systems and Museum of Comparative Zoology), Constance Woodman (Hill Country Aviaries), Mark Robbins (University of Kansas Biodiversity Institute), James Schiffbauer and Tara Selly (University of Missouri X-ray Microanalysis Core), Abby Rainwater (MU Raptor Rehabilitation Program), Matt Colbert, Jesse Maisano, and Timothy Rowe (University of Texas High-Resolution X-ray CT Facility), and the Truman VA Biomolecular Imaging Center. My special thanks go out to M. Scott Echols who helped forge a number of aviary and veterinary connections at Tracy Aviary, Hill Country Aviaries and with Constance Woodman, Mark Nielsen, and Brian Speer. I also thank M. Scott Echols for his direct contributions throughout the length of this project. Many people took the time to ask questions and discuss this project as it developed and as results began to manifest. Without these conversations and interactions with colleagues at conferences, over email, and on a daily basis ii some of the ideas in this project may have hobbled along or never fully taken flight. These individuals span from undergraduate students to emeritus faculty. I specifically thank Kelsie Abrams, Alida Bailleul, Jen Bright, Julieta Carril, Don Cerio, Jasmine Croghan, Julian Davis, Federico Degrange, Catherine Early, Daniel Field, Stephen Gatesy, Paul Gignac, Nat Graham (and everyone else at Science on Tap), Brandon Hedrick, Anna Nele Herdina, Anthony Herrel, Tobin Hieronymus, Dominique Homberger, Marc Jones, Mackenzie Kirchner-Smith, Daniel Ksepka, Sharon Kuo, Melissa Macias, Armita Manafzadeh, Faye McGechie, Ralph Molnar, JP Nassif, Aaron Olsen, Benjamin Perlman, Kristen Prufrock, Emily Rayfield, Ryan Ridgely, Rachel Rozin, Kaleb Sellers, Anmol Sethi, Cailtyn Smith, Eric Snively, Anthony Spates, Spiro Sullivan, Diego Sustaita, Ian Trevethan, Morgan Turner, Conner Verhulst, Alec Wilken, Larry Witmer, and Natalie Wright. Many of the above mentioned people have become good friends over the years and I value their involvement in this project and everyday life jointly. I thank my closest friends, who have checked in on not just the status of the project, but the status of life. They kept me balanced throughout the process. For that I sincerely thank Nina Haro and Joanna Kerby (for asking questions about teaching), Jeff Johnson (for never forgetting to do something to make me laugh on my birthday), Anthony Luna (for all sorts of things, but mostly making me jealous because he saw some rare bird while I was typing away), Frances Owen, Clint Helms, iii Jess Casey, and Brian Zinke (for continuing to work outdoors like we all did at Fort Hays and sharing photos of their work). I also must thank S. Christopher Bennett for taking a chance on me as a master’s student, beginning this whole journey and Greg Farley for encouraging my interests in birds. I also thank all of the students I ever taught, undergraduate students and middle school alike. I thank my family for always supporting me and urging me to push myself to get what I want from life and never to settle. My mother Diana for having the energy of five people and tiring me out over the phone talking about my project and life and my brother Tyler for updating me constantly about music, hockey, and video games over the years, giving us something different to talk about. My second set of parents, Tammie and Brian for always being interested in whatever I was doing during this project because they were just naturally curious about some of the things I was doing, have my heartfelt thanks. And finally, and never least, my wife Jennifer who suffered through my desire to quit a full time job to go back to school, became a reluctant amateur bird watcher, moved to western Kansas, and listens to me talk about parrots and computer modeling on a more than regular basis and does it all with a smile, even if she just wants to read a book and have a quiet moment without birds being mentioned. iv TABLE OF CONTENTS Acknowledgments ................................................................................................. ii List of Figures and Illustrations ............................................................................ vi List of Tables ........................................................................................................ ix Abstract ................................................................................................................ x Chapter 1: Introduction: Avian Evolution and Feeding .......................................... 1 Chapter 2: Visualization of Complex Muscle Function of Gape, Ontogeny, and Evolution in Three Dimensions ........................................................................... 16 Introduction ......................................................................................................... 16 Methods .............................................................................................................. 19 Results and Discussion ...................................................................................... 24 Conclusion .......................................................................................................... 33 Chapter 3: Palatal biomechanics and its significance for cranial kinesis in Tyrannosaurus rex .............................................................................................. 43 Introduction ......................................................................................................... 43 Methods .............................................................................................................. 46 Results .............................................................................................................. 54 Discussion .......................................................................................................... 59 Conclusion .......................................................................................................... 67 Chapter 4: Analysis of the Biomechanical Environment of the Feeding Apparatus of Parrots Using Finite Element Models, Statistics, and Bone Mechanics .......... 83 Introduction ......................................................................................................... 83 Methods .............................................................................................................. 88 Results .............................................................................................................. 96 Discussion ........................................................................................................ 114 Conclusion ........................................................................................................ 123 Chapter 5: Summary ........................................................................................ 159 Literature Cited ................................................................................................. 164 Vita ................................................................................................................... 189 v LIST OF FIGURES AND ILLUSTRATIONS Figure 1-1. Phylogenetic tree of Archosauria and Lepidosaur sister group ....... 14 Figure 1-2. Diagrammatic representation of cranial kinesis with an emphasis on the contraction of m. pterygoideus et quadrati (mPPT) ...................................... 15 Figure 2-1. Phylogenetic tree of archosaur taxa in this study ............................ 35 Figure 2-2. Muscles pictured in situ in diffusible iodine contrast-enhanced computed tomography (DiceCT) ......................................................................... 36 Figure 2-3. Illustrated description of ternary diagram construction .................... 37 Figure 2-4. Muscle maps of models used to analyze muscle resultants in this study ................................................................................................................... 38 Figure 2-5. Changes in muscle orientations during a gape cycle in A. mississippiensis (MUVCAL008 and MUVCAL031) ............................................
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