TAXONOMY OF THE STERKFONTEIN FOSSIL CERCOPITHECINAE: THE PAPIONINI OF MEMBERS 2 AND 4 (GAUTENG, SOUTH AFRICA) Jason L. Heaton Submitted to the faculty of the University Graduate School in partial fulfillment of the requirements for the degree Doctor of Philosophy in the Department of Anthropology, Indiana University November 2006 Accepted by the Graduate Faculty, Indiana University, in partial fulfillment of the requirements for the degree of Doctor of Philosophy. Doctoral Committee ______________________________________ Travis R. Pickering, Ph.D. ______________________________________ Kevin D. Hunt, Ph.D. ______________________________________ Nicholas Toth, PhD. ______________________________________ Kathy D. Schick, PhD. October 16, 2006 © 2006 Jason L. Heaton ALL RIGHTS RESERVED To my wife, Alison “Wheresoever she was; there was Eden.” Mark Twain (1904) – The Diaries of Adam and Eve ACKNOWLEDGEMENTS This analysis originated on a trip to South Africa in 2000 and evolved into what you see before you. Along the way, its path was modified and forever altered by those individuals with whom I came into contact. Without those interactions, I must say this analysis would not be what it is today. First and foremost, I thank my committee, Travis Pickering, Kevin Hunt, Nick Toth and Kathy Schick. I was fortunate to have a group of individuals that truly provided the materials that I needed to succeed. Nick and Kathy are the reason that I went to Indiana to study paleoanthropology. I am grateful to Nick for writing a letter of recommendation for grad school, which undoubtedly facilitated my entrance into IU’s program. I enjoyed the many seminars that I took with Nick, Kathy and Kevin which shaped much of the way I view the prehistoric world, today. Also, I must thank Nick and Kathy for passing on the importance and value of experimental archaeology. Kevin was always great at allaying the uncertainties that most grad students encounter and was always encouraging. Kevin, Nick and Kathy were great supporters when I decided two years into the program that I wanted to dual-major in biological anthropology and archaeology (i.e. paleoanthropology). Each was assured of my success much before I was. Words cannot express the gratitude that I feel toward them. To each of them, I am forever thankful. v vi Over the six years that Travis and I have worked together, first as a grad student (JLH) and a post-doc (TRP), we have become close friends. It is not everyday that a grad student can truly call their chair a valued friend and a mentor. At times, it was difficult having a friend, as a chair, but no matter our disagreements, I knew that Travis truly had my best interest in mind. Travis is the sole reason that I ended up researching in South Africa; he was always great about including me in research projects and making me feel as an equal collaborator. Without him, the analysis produced herein would not have existed. Travis served both as facilitator and promoter for me during my initial trips to South Africa. It was his close relationship with others in South Africa that made my job much easier. For this, I am thankful. I am proud to call Travis a friend and a colleague, and I look forward to our continued collaboration with the Swartkrans Palaeoanthropological Research Project and beyond. I must also thank those individuals that provided access to the materials under their care. In my six years of research, dissertation and otherwise, the majority of my time has been spent in either Johannesburg at the Department of Anatomical Sciences (University of the Witwatersrand) or Pretoria at the Department of Vertebrate Palaeontology (Transvaal Museum). In the Anatomical Sciences at Wits, Ron J. Clarke, Bev Kramer, Phillip Tobias and Kevin Kuykendall were responsible for providing access to the Sterkfontein, Taung, Bolt’s Farm and Makapansgat Collections. Heather White was always accommodating and welcoming. I appreciate the personal touch that she brings to her work. She is a bright spot in the Department of Anatomical Sciences. Andre Keyser and Colin Menter provided access to the Drimolen primate sample and are continuing collaborators on its vii description. Andre and Colin were always willing to include me in any additional projects that they were involved in; however, in most cases, it involved animal feces of some sort. Some of my best memories from South Africa include our stint at Misgrot Cave in 2004. Spending a few days ankle deep in baboon dung, will teach you a lot about a person. I am also indebted to Colin for introducing me to Adobe Photoshop and Microsoft Access upon one of my first trips. The data collection process was greatly enhanced by the use of Access. Furthermore, the use of Photoshop to combine photos, standardize scales and in some cases, provide a solid background for my sometimes less than perfect photos resulted in a more superior thesis than would have otherwise been the case. For that, I thank Colin. My time in Pretoria was no less enjoyable or educational. At the Transvaal Museum, Francis Thackeray and Stephanie Potze aided kindly in access to the Kromdraai, Swartkrans, Sterkfontein, Schurweberg and Taung materials under their care. It was always a pleasure to work with Steph; one could not ask for a better person. I always enjoyed discussing palaeoanthropology, or life in general, while sitting on the steps of the TM. Teresa Kearney was also helpful in providing access to the modern comparative material and in some cases, the fossil collections. The library staff, most specifically Tersia Perregil, was always of great help in chasing down references, getting out-of-print articles and providing a means to photocopy any necessary materials. Thanks should also go to Eddie Reinecke, the IT Facilities manager at the TM, who was always great in getting internet access for me while I was at the museum and always a source of discussion about the latest gadgets for digital SLR’s. viii Additionally in 2004, I was able to spend a few days in Cape Town studying the Taung cercopithecoid collection. Graham Avery and Kerwin van Willingh of the South African Museum were most helpful in providing the necessary access and were gracious hosts. On a more personal, I must first thank Shuna Huffman for taking me into her home and allowing me to become part of her family. Ron Clarke was also gracious in allowing me to study the unpublished primate material from Sterkfontein. I am always impressed at Ron’s ‘eye for anatomy’ and enjoy the time that have we spent together. Not only is Ron a fountain of knowledge, but he is always a great person to be around. I have fond memories of the many dinners that I spent with Shuna, Ron and Kathy Kuman. Kathy is one of the sweetest people in this world, often unselfishly putting the needs of her students and/or colleagues above her own interests. Kathy also has a skill for what she does, and I enjoyed ‘digging’ termite mounds with her, Travis and the late Tom Loy. Ron, Kathy and Shuna have all made me feel like I have a home-away-from- home. The same can be said about the Brain family. On many occasions, the Brain’s – Bob, Laura, Mel and Duncan – welcomed me into their home providing me with an always appreciated home-cooked meal. Laura’s babotie was always outstanding. Over the last few years, I have enjoyed getting to know Bob, Laura and Mel, and I look forward to visiting them each time that I am in South Africa. Several friends, graduate students and colleagues have also supported my research in one way, or another, but a few deserve special acknowledgement. While an undergrad at South Alabama, I spent many hours discussing evolution with my friend, Edward Hed. Ed was one of the prime motivators of my going to graduate school and remains a true ix friend, today. A better friend could not be imagined. David Nelson, a professor at the University of South Alabama, changed my life when I took his course, Comparative Vertebrate Anatomy, and I begin to see the evidence for homology and common decent. The way I saw the world forever changed. Sheldon Gottlieb, also a professor at South, was a great promoter of evolution and was always willing to spark a lively debate on the intricacies of the science and religion debate. To each of them, I provide thanks. At Indiana, Dan Osborne was a great friend and has on many occasions served as a sounding board for my ideas. I always enjoyed the time that I spent with Dan and Britta Grieshaber while at IU. Charles Egeland has also been a source of entertainment and motivation over the years, as he is one of the most driven individuals that I know. Over the years, several people with whom I have had the pleasure of getting to know while in South Africa have provided hours of support and entertainment. They are: Morris Sutton, Dorian Staps, Kerstin Engel, Mathias Huck and Job Kibii, Vic Mackey and Cletus Van Damme. To each, I am thankful for the time that we spent together. This research was also funded by the Graduate and Professional Student Organization (GPSO) at Indiana University, the David Skomp Fellowship and Sigma Xi. Thanks are due to these agencies for providing support for this analysis. Ultimately, the greatest bulk of appreciation is due to my family. Growing up a poor kid in rural Alabama provides its own set of obstacles; however, my family gave me the strength and the intestinal fortitude to withstand any and all challenges. My father and step-mother, Cecil and Shirley Heaton, have supported my educational goals.