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Reproductive Life Histories of Mammoths by Joseph Jameel El Adli

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Reproductive Life Histories of Mammoths by Joseph Jameel El Adli Reproductive Life Histories of Mammoths by Joseph Jameel El Adli A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy (Earth and Environmental Sciences) in the University of Michigan 2018 Doctoral Committee: Professor Daniel C. Fisher, Chair Professor Catherine Badgley Professor Emeritus Philip D. Gingerich Professor Kyger C Lohmann Associate Professor Jeffery A. Wilson Joseph J. El Adli [email protected] ORCID iD: 0000-0002-3989-5409 © Joseph J. El Adli 2018 DEDICATION To all those I’ve met along the way. ii ACKNOWLEDGEMENTS Looking back on the five years of work that went into assembling this body of research, it is impossible (and would quite frankly be disingenuous) for me to treat this as anything but the result of a collective effort. Whether through discussions of methods and results, direct contributions to projects, or just sheer moral support, I have completed this document carried on the backs of many individuals whom I would like to thank here. My gratitude for your presence in my life is greater than you can know. First and foremost, I would like to thank my advisor, Dan Fisher, for giving me the chance to work on such fantastic material and teaching me new, creative methods of understanding past life. Dan’s guidance, insights, support, and discussions have immensely improved how I think about conducting scientific research and how I communicate my work to others. I am indebted to Dan for the many opportunities that he has provided for me to work in the field and travel to other countries to present my work. Our adventures in Siberia will always be some of my fondest memories from graduate school. I would like to thank Catherine Badgley, Philip Gingerich, Kacey Lohmann, and Jeff Wilson for sitting on my dissertation committee and providing constructive feedback on my research. Their time and efforts have greatly improved the experimental design of my research, which was crucial to the completion of this body of work. I also am appreciative of my labmate, Mike Cherney, and of Adam Rountrey who provided iii discussions of many aspects of proboscidean research, as well as training in various techniques used within this study. My research also benefitted from many discussions with Tom Baumiller, Alessio Capobianco, Matt Friedman, John Fronimos, Pam Horsley, Katie Loughney, Kelly Matsunaga, Kyle Meyer, Dan Miller, Kris Rhodes, Kierstin Rosenbach, James Saulsbury, Tara Smiley, Selena Smith, Meg Veitch, Ian Winkelstern, and Miriam Zeldich. The specimens used within this study were provided by a variety of sources, which all warrant recognition. John Harris, Aisling Farrell, and Shelley Cox (La Brea Tar Pits and Museum) provided access to and allowed removal of the tusk sample from Zed, which provided the data for Chapter 2. Rafael Labarca (Universidad Austral de Chile) collected and sent the material required to complete Chapter 3. Bernard Buigues (International Mammoth Committee), Semyon Grigoriev (Mammoth Museum, Institute of Applied Ecology of the North, North-East Federal University of Yakutsk), Alexei Tikhonov (Zoological Institute, Russian Academy of Sciences), and Sergey Vartanyan (Northeast Interdisciplinary Research Institute, Far East Branch, Russian Academy of Sciences) were responsible for the procurement of many of the specimens used in Chapters 4 and 5. The isotopic work done within this study could not have been completed without the help of Lora Wingate (University of Michigan Stable Isotope Lab) and Kacey Lohmann. Access to the University of Michigan Dental School MicroCT Core facility was provided by Michelle Lynch. I owe enormous gratitude to Matt Friedman, Selena Smith, and Brian Ellis for allowing me to use the Nikon X TH 225 ST industrial microCT in the Computed Tomography in Earth & Environmental Sciences laboratory. I am further iv thankful to Matt and Selena for allowing me to use their allotted scan times for such extended periods of time. I am also greatly indebted to Kelly Matsunaga for training me on how to use and maintain the microCT scanner and for spending countless night and weekend hours repairing the machine for me. I also certainly could not have completed my research without the help of several undergraduate students, such as Jess Hicks, Ethan Shirley, Kat Lewandowski, Nichole Lorhke, and Lang DeLancey. I would like to thank Scott Beld for his assistance with thin section preparation and production of photomicrographs of specimens. Linda Garcia, Anne Hudon, and Cindy Stauch were instrumental in helping me navigate the administrative aspects of the graduate program. I am grateful for the friendship and comradery that I have found during my time at the University of Michigan. It would take too many pages to express my thanks for every person that has enriched my life since arriving in this program, so I am only able to list their names and hope that they understand how much they mean to me: Alyssa Abbey, Will Bender, Laura Bilenker, Austin and Camilla Bowles, Tristan Childress, Anaïs Ferot and Thomas Giachetti, “Man Sam” Haugland, Tom Hudgins, Jaayke Knipping and Adrian Fiege, Erin Lynch, Ross Maguire and Kate Volk, Kyle Meyer, Tiffany Napier and Jimmy Holub, Sammie Sammie NemNem and Sean Wilson, Yi Niu, Xiaofei Pu, Mark Robbins, Liz Tanis, Meghan Taylor, and Ian Winkelstern. I would like to thank my parents for their constant support and help. Without them I would not have been able to pursue a career in science. Most importantly, I would like to thank my wife, Pam Horsley, for allowing me to drag her halfway across the country (and out of her homeland) so that I can pursue a doctoral degree. Thank you for your patience, care, and understanding through these past five years. I know that the best is v still to come for us. Finally, I would like to thank Karley for always being a fat little dog with occasionally good intentions. vi TABLE OF CONTENTS DEDICATION .................................................................................................................. ii ACKNOWLEDGEMENTS ............................................................................................... iii LIST OF TABLES ........................................................................................................... x LIST OF FIGURES ......................................................................................................... xii ABSTRACT .................................................................................................................. xiv CHAPTER 1: Introduction ............................................................................................. 1 THE END ..................................................................................................................... 1 THE PROBOSCIDEA .................................................................................................. 2 The mammoths ........................................................................................................ 2 TUSKS ......................................................................................................................... 4 CHAPTER OVERVIEWS ............................................................................................. 5 General overview ..................................................................................................... 5 Chapter 2 ................................................................................................................. 6 Chapter 3 ................................................................................................................. 7 Chapter 4 ................................................................................................................. 7 Chapter 5 ................................................................................................................. 8 Chapter 6 ................................................................................................................. 9 REFERENCES .......................................................................................................... 10 CHAPTER 2: Last years of life and season of death of a Columbian mammoth from Rancho La Brea .................................................................................................. 12 ABSTRACT ............................................................................................................... 12 INTRODUCTION ....................................................................................................... 13 MATERIALS AND METHODS ................................................................................... 14 RESULTS .................................................................................................................. 17 External features of the tusk .................................................................................. 17 Stage of dental development and sex .................................................................... 20 Computed tomographic features ............................................................................ 23 Second-order increment profile .............................................................................. 25 vii Isotope data ........................................................................................................... 27 DISCUSSION ............................................................................................................ 28 CONCLUSION ..........................................................................................................
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