Osteological, myological, and biomechanical investigations of the sauropod dinosaur Dreadnoughtus schrani and molecular paleontological investigation of the marine crocodile Thoracosaurus neocesariensis A Thesis Submitted to the Faculty of Drexel University by Kristyn K. Voegele in partial fulfillment of the requirements for the degree of Doctor of Philosophy August 2016 © Copyright 2016 Kristyn K. Voegele. All Rights Reserved. ii Dedication To those with dreams and those that help them achieve dreams iii Acknowledgements This project would not be possible without the support, assistance, and guidance of just about everyone I have had interacted with in the last five years. I am grateful for all of your time, effort, patience, and support. First, I would like to thank Drexel University, the College of Arts and Sciences, the Department of Biology, the Academy of Natural Sciences, the Department of Biodiversity, Earth, and Environmental Science, and all the faculty and staff that made my graduate education possible. At Drexel I have been fortunate to have the opportunity to learn from world-class scientists about research, teaching, and everything in between. I appreciate the resources and support provided to me over my graduate career to turn me into a capable scientist. I am also thankful to the organizations that have financially supported my research. Without the financial support from a National Science Foundation Graduate Research Fellowship (DGE Award #1002809) and a Paul Bond Scholarship from the Delaware Valley Paleontological Society much of my research would have been unaffordable. I am thankful to these institutions for helping me expand my area of focus and training in my field of study during my graduate career. I am very grateful to my advisor, Dr. Kenneth Lacovara, for accepting me into his program. The opportunities for diverse research using new technology as part of your program have opened many possibilities for me. Thank you for allowing me to join your research team and all experiences that went along. They have enriched my graduate experience and you have taught me skills I never knew would be useful in paleontology. Also, I am truly appreciative to Dr. James Spotila for "adopting" me as his graduate student for my last year of gradschool. Thank you for making this past year so much less stressful and for instantly supporting me as if I had always been your student. I am extremely grateful to the members of my dissertation committee for their time, advice, access to their laboratories and supplies, training, and overall support: Dr. Aleister Saunders, Dr. Spotila, Dr. Sorin Siegler, Dr. Matthew Lamanna, and Dr. Mary Schweitzer. Thank you for everything you have taught me. I am honored to have learned from each of you. I sincerely hope I can help future students reach their goals as you have helped me. I particularly want to thank Dr. Schweitzer for allowing me to join your lab for many months to conduct research and Dr. Siegler for the patience to teach me so much engineering. Thank you to all my labmates and fellow graduate students: Elena Schroeter, Paul Ullmann, Zachary Boles, Anna Jaworski, Rebecca Hoffman, Ashley Adams, Jason Schein, Emma Fowler, Aja Carter, Nathan Schiff, Athena Patel, Suraj Pandya, and George Keighton. I am grateful for all your help, assistance, and fun adventures we have had. Anna, we started and ended graduate school together. It has been a pleasure to have taken this road with you and always have someone who could immediately relate. Emma, your talents have been crucial to this project and I appreciate all the time you took to help me. Together we have learned so much about science and life. I also have to thank every engineer who has helped me learn your field: Dr. Jason Toy, Dr. Ramya Namani, Dr. Marissa Powers, Ben Senderling, Tyler Voegele, Jordan Senff, Eric Brace, Natalie Brace, iv David Mcdivitt, and Deeksha Seth. I could not have accomplished any of my biomechanics research without you. Your patience has been commendable. Several people have been instrumental to various portions of my dissertation. Thank you Wenxia Zheng, Dr. Allison Moyer, and Tyler Bridges for your help at North Carolina State University. Along with Paul Ullmann, Elena Schroeter, and Zack Boles you have helped me navigate and trouble shoot molecular paleontology. I have learned so much in this field from you and this research wouldn't have happened without you. Also, I deeply appreciate the friendship and hospitality of Cindy and Dr. Art Bogan. Thank you for opening your home and supporting me and my research endeavor. I also appreciate the assistance of Dr. Matthew Bonnan for the advice and insight into extant cartilage morphology. Thank you to Ruth Elsey and the Rockefeller Wildlife Refuge for a cadaver Alligator for my research. I also have to thank the hundreds of volunteers that excavated, prepared, moved, and 3D laser scanned the fossils used in my research. Your work was essential and I would still be working at it without you. I am extremely thankful to Dr. Joseph Hartman and Dr. Ron Nellermoe for my first opportunities in paleontology. These first tastes have kept me hooked on this science. I am immeasurably grateful for the opportunities your first belief in me have brought and for your continued support in me. Thank you for everything you have taught me about science and life. I am a better person and scientist because of you. Finally I need to thank my family for always seeing me as a paleontologist. My goals never seemed weird to you and you have always supported me. Thank you for all the extra effort over the years to help me follow my dream. I would not be where I am today without you. From you I have learned how to face opportunities and hardships and to keep going no matter what. You have gone out of your way to help me and it has not gone unnoticed. I am so grateful to have you with me on this journey. v Table of Contents LIST OF TABLES ............................................................................................................. ix LIST OF FIGURES .............................................................................................................x ABSTRACT ..................................................................................................................... xvi CHAPTER 1: OSTEOLOGICAL DESCRIPTION OF THE DORSAL VERTEBRAE OF THE GIANT TITANOSAURIAN SAUROPOD DREADNOUGHTUS SCHRANI ...........1 1.1 Abstract ....................................................................................................................1 1.2 Introduction ..............................................................................................................2 1.2.1 Terminology ....................................................................................................4 1.2.2 Anatomical Abbreviations ...............................................................................4 1.3 Description ...............................................................................................................5 1.3.1 Dorsal Vertebra 1156?-4 ................................................................................6 1.3.2 Dorsal Vertebra 1156?-5 ................................................................................9 1.3.3 Dorsal Vertebra 1156-6 ................................................................................10 1.3.4 Dorsal Vertebra 1156?-7 ..............................................................................12 1.3.5 Dorsal Vertebra 1156?-8 ..............................................................................13 1.3.6 Dorsal Vertebra 1156?-9 ..............................................................................15 1.3.7 Dorsal Vertebra 1156?-10 ............................................................................17 1.3.8 Dorsal Vertebra 1156-11 ..............................................................................18 1.4 Discussion ..............................................................................................................19 1.4.1 Comparisons .................................................................................................19 1.4.2 Assignment of predicted serial progression ..................................................22 1.5 Conclusion .............................................................................................................25 vi CHAPTER 2: APPENDICULAR MYOLOGICAL RECONSTRUCTION OF DREADNOUGHTUS SCHRANI ........................................................................................33 2.1 Abstract ..................................................................................................................33 2.2 Introduction ............................................................................................................34 2.3 Methods..................................................................................................................38 2.3.1 Material.........................................................................................................39 2.4 Results and Discussion ..........................................................................................39 2.4.1 Forelimb Musculature ..................................................................................39 2.4.1.1 Axial to Limb .......................................................................................39 2.4.1.2 Girdle to Distal .....................................................................................45