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Vertical Climbing Adaptations in the Anthropoid Ankle and Midfoot: Implications for Locomotion in Miocene Catarrhines and Plio-Pleistocene Hominins

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Vertical Climbing Adaptations in the Anthropoid Ankle and Midfoot: Implications for Locomotion in Miocene Catarrhines and Plio-Pleistocene Hominins VERTICAL CLIMBING ADAPTATIONS IN THE ANTHROPOID ANKLE AND MIDFOOT: IMPLICATIONS FOR LOCOMOTION IN MIOCENE CATARRHINES AND PLIO-PLEISTOCENE HOMININS. by Jeremy M. DeSilva A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy (Anthropology) in The University of Michigan 2008 Doctoral Committee: Associate Professor Laura M. MacLatchy, Chair Professor Daniel C. Fisher Professor John C. Mitani Professor Milford H. Wolpoff Research Scientist William J. Sanders © Jeremy M. DeSilva All rights reserved 2008 Acknowledgements This research would not have happened without the generous funding of the Leakey Foundation. Thank you to Paddy Moore for her assistance during this process. Part of this research was also funded by the National Science Foundation dissertation improvement grant BCS 0751010. Thank you to the following for insightful conversations about hominin ankles and feet: Bruce Latimer, Owen Lovejoy, Will Harcourt-Smith, Jeff Meldrum, Roshna Wunderlich, Thank you Susy Cote for all of those great conversations about Miocene catarrhines! I am grateful to Karen Isler for sharing unpublished data and for discussing vertical climbing with me. Jack Stern, Susan Larson, and Brigitte Demes generously allowed me to analyze unpublished EMG data of chimpanzees. Dissection of the male gorilla was made possible by Philip Myers, and Bradley Wathen, Thomas Meier, and Gail Rising were instrumental in allowing me to x-ray and dissect baboon and macaque specimens. I am grateful to Tomasz Baumiller for helping me to test the biomechanical properties of non-human primate ligaments and for insightful discussions about ligament structure and mechanics. Thanks to Dr. Harry Maisel for teaching me anatomy, and to both Dr. Maisel and Barbara Russo for permitting me to dissect human cadavers in their care. Study of living primates was made possible by the Toledo Zoo and its staff. I am so grateful to Dr. Ann Duncan at the ii Detroit Zoo for allowing me to study the apes and monkeys during their physical exams. It was an honor to watch you and your staff keeping the apes and monkeys healthy. I am extremely grateful for all of those who allowed me to study skeletal specimens in their care: Eileen Westwig at the American Museum of Natural History, Linda Gordon at the National Museum of Natural History, Kristof Zyskowski at the Yale Peabody Museum, Bill Stanley and Michi Schulenberg at the Field Museum, Owen Lovejoy at Kent State University, and Stephen Hinshaw at the University of Michigan Museum of Zoology. My sincere gratitude is extended especially to Judy Chupasko at the Harvard Museum of Comparative Zoology, and Lyman Jellema at the Cleveland Museum of Natural History for allowing multiple visits to study those collections! I am grateful to Yohannes Haile-Selassie, David Pilbeam, and Michele Morgan for letting me study casts of hominin and hominoid fossils in their care. Thank you as well to Carol Ward for sending me a cast of the A. anamensis distal tibia before I had a chance to see the original in Kenya. Thank you to Matt Tocheri, Jeff Meganck, and Aaron Wood for their assistance with the Next Engine 3D scanner. Many, many people made my work at Ngogo possible and enjoyable. Thanks to the Uganda Wildlife Authority, Dr. Jerry (Jeremiah Lwanga), David Watts, Kevin Potts, Godfrey, Alfred, and Adolf. John Mitani- it was an absolute pleasure to learn about these amazing chimpanzees first hand from you. Marissa Sobolewski, Brent Pav, and Lauren Sarringhaus- I cannot think of three other people that I would rather trudge through the papyrus swamp with. Studying fossils at the Uganda National Museum was made possible by the Uganda National Council for Science and Technology. Thank you to Dr. Rose Mwanja, and to Ezra Musiime for permission to study fossils in their care. To my iii friend Dr. Robert Kityo- it is always a pleasure to work with you, and I look forward to many future collaborations. To my good friends Moses Mafabi, Nelson Abiti, and Petra- thank you for always making me long to return to Uganda. My work in Tanzania was made possible by the Tanzanian Commission for Science and Technology. Thank you so very much to Amandas Kwekason and Dr. Paul Msemwa for their assistance in allowing me to study the OH 8 foot. Thanks to Terry Harrison for guidance as I prepared my trip to Dar es Salaam. Thank you to the Kenyan Ministry of Education, Science and Technology for permission to do research in Kenya. I am so grateful to Drs. Idle Omar Farah and Emma Mbua for permission to study hominins and hominoid fossils in their care. Meave Leakey and Lucy Mbogori assisted me while I was in the Museum, and thank you to Rick Potts and Briana Pobiner for their help while I was in Nairobi. I would still be wandering the streets of Johannesburg (gulp) if it were not for Alison Groves, Maggie, and Adam. Thank you for letting me part of such a wonderful family while I was in South Africa. I am eternally grateful to Dr. Bernhard Zipfel for allowing me to study hominin bones in his care and for many stimulating conversations about hominin feet. I look forward to future collaborations with you! Thanks to Taflene Crawford for carting me around Johannesburg. Thanks to Christine Steininger for all of your help and Philip Tobias for directing me to Peter Christie’s dissertation. I am grateful to Matthew for getting me back and forth from Johannesburg to Pretoria. While in Pretoria, I am grateful to Francis Thackeray and Stephanie Potze for allowing me to study postcranial bones in their care. Thanks to the staff of the Helen Joseph Hospital in iv Pretoria for CT scanning hominin tali. Thanks to Jamie Clark for being a familiar face half-way around the world. I am so fortunate to have been part of an incredible cohort here at Michigan: Marissa, Brent, Lauren, Julie, Seki, Mary, and Elizabeth. Adam- thanks for always providing me with an example of how to get through Michigan fast and well. I am very excited to be heading east with you. Robin- thanks for understanding, truly understanding, as I hit dissertation roadblocks. Denise, I could not have asked for a better colleague and friend as a lab mate. Thanks for all you have done and I look forward to a continued friendship and professional collaborations as the years go on. I learned so much from my professors at Michigan: Drs. Frisancho, Strassmann, Brace, Zelditch, Caspari, Gingerich, Baumiller, Mukasa, Wilkinson, Wilson, Lohmann, Speth, Whallon, Irvine, Mannheim, and Pastner. I am especially grateful to Thore Bergman and Jacinta Beehner for giving me advice as the job process was unfolding, and for allowing me access to video data on cercopithecoids. Laurie Marx, you are a true gift to the department and to all of the anthropology graduate students- I’d be lost without you. Thank you. Thank you so very much to my committee: Laura MacLatchy, Dan Fisher, John Mitani, Bill Sanders, Milford Wolpoff. I have learned so much from you all and cannot ever express how grateful I am. There are not many students lucky enough to be able to say that their committee members are also their friends, and I have that great fortune. You have all provided for me a model for how to have a successful career as a scientist, a teacher, and a mentor. I look forward to a continued relationship with all of you. I could not have possibly asked for a better advisor than Laura MacLatchy. Laura, thank you so v much for taking a chance on me at BU back in 2003, and thank you for the opportunity to come to Michigan with you. You have challenged me, fostered my interest in functional morphology, and given me the model for how to be a true professional. I look forward to future collaborations with you! Kathleen: thanks for taking me in and being a Mom to me during those first few lonely months in Ann Arbor. Tim Kardatzke and Ken Pauley- thanks for always wanting to talk about science, Africa and human evolution- I cannot wait to be back in Massachusetts and pick-up where we left off. To Lucy Kirshner: none of this would have been possible without you. Thank you for believing in me, for first stimulating and then supporting my obsession with hominids, and for encouraging me to apply to grad school. My family has been incredibly supportive through this process- Mom, Dad, Rich, Deana, Mel, Patricia, Kittie, Rick, Lorrie, Mike. You have all been so wonderful to Erin and me. Especially to Alex, Lillian, Jake, Ella, and Anthony: thank you for providing me with the motivation to finish and come home. Charlie: thank you for taking me on long walks and allowing me to read large sections of my thesis to you. To the love of my life Erin: I cannot ever thank you enough for your support during the last five years. I could not have done this without you. You have made me laugh when none of this seemed funny (thank you…..thank you), you have made me interested in things other than ankles, and you have provided me with the motivation to finish my degree. I am so proud of what you have accomplished while you have been here and look forward to the rest of our life together back in Massachusetts. vi Table of Contents Acknowledgements………………………………………………………………………..ii List of Figures……………………………………………………………………….......viii List of Tables……………………………………………………………………………..xi Abstract………………………………………………………………..…………............xii Chapter 1. Introduction……………………………………………………………………..1 2. Kinematics of the talocrural joint during vertical climbing bouts in wild chimpanzees…………………………………………………………………….9 3. Vertical climbing adaptations in the anthropoid talocrural joint: Implications for early Miocene catarrhine locomotion……………...………...41 4.
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