Dental variation in African apes with implications for understanding patterns of variation in species of fossil apes by Varsha Catherine Pilbrow A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy Department of Anthropology New York University January 2003 _________________________ Terry Harrison © Varsha Pilbrow All Rights Reserved, 2003 ACKNOWLEDGEMENTS For providing support and encouragement at various stages of this project I owe thanks to several individuals and research and funding agencies. First, I must acknowledge the financial support I received. The Margaret and Herman Sokol Predoctoral Travel Award, the L. S. B. Leakey Foundation, the Wenner-Gren Foundation for Anthropological Research and the National Science Foundation provided funding for the data collection phase of this project. Without their help this project might still have got done, but differently, someplace else, and by someone else. I must also thank the Graduate School of Arts and Science at New York University for supporting me as a teaching and research assistant while I was completing my graduate education. The New York Consortium of Evolutionary Primatology (NYCEP) provided support for travel to the anthropology meetings, for which I am grateful. For his superb mentoring qualities and his unflinching support throughout my time as a graduate student, I thank my Ph.D. supervisor, Terry Harrison. Terry encouraged me to try again when my grant proposals got rejected, and helped by proofing and editing the numerous versions. The successful completion of this project is largely due to his tenacity. I also thank Cliff Jolly for his longstanding association with my project. Cliff, like Terry, has been on my committee since the start and was always obliging with his advice, letters of recommendation when needed, and general words of encouragement. I also acknowledge the writings, the iv lectures and the stimulating conversations I’ve had with Cliff and Terry. I feel fortunate to have had the opportunity to study with them. I thank Pam Crabtree for her helpful comments on the final draft of the thesis. Pam has been my source of enlightenment and courage for several years. Her strong commitment to both her professional and personal life and her deft handling of both without compromising either provided me with a beacon of hope. I benefited immensely from the input of Tony diFiore and Vince Stefan. Tony (NYU) and Vince (Lehman College) helped me analyze my data and had several useful suggestions that helped strengthen my project. I thank them for their patience in reading the thesis in its various forms and for their critical comments. I am grateful for the congenial atmosphere provided by the “Ninth Floor Research Group”. I should particularly like to thank Leahanne Sarlo for saving my mail when I was away. For providing a relatively non-threatening environment for voicing my faltering opinions I thank NYCEP, and the organizer of the student research colloquia, Eric Delson. I would like to thank the following individuals and institutions for providing access to hominoid dental material: Salvador Moyà Solà (Institute de Paleontologia, “Miguel Crusafont”), Jacques Cuisin, Frances Renoult and Brigitte Senut (Muséum National d’Histoire Naturelle), Maria Rutzmoser (Museum of Comparative Zoology), Paula Jenkins and Peter Andrews (British Museum of Natural History), Malcolm Harman (Powell-Cotton Museum), Nathalie Memoire v (Muséum d’Histoire Naturelle, Bordeaux), Robert Martin (Anthropologisches Institüt der Universität Zürich-Irchel), O. Röhrer-Ertl (Zoologische und Anthropologisches Staatssammlung, Munich), Prof. Rabeder (University of Vienna), Franz Stojaspal (Geologisches Bundesanstalt), Güdrün Höck (Naturhistorisches Museum), Laszlo Kordos (Geological Institute of Hungary), P. J. Heizmann (Staatlisches Museum für Naturkunde), Jens Franzen (Senckenberg Museum), A. Liebau (Karl-Eberhardt Universität ), Wim Van Neer (Royal Museum of Central Africa, Tervuren), Meave Leakey (Kenya National Museum), Richard Thorington, Jr. and Linda Gordon (Smithsonian Institution’s United States National Museum), Bill Stanley (Field Museum of Natural History), David Pilbeam, Larry Flynn and John Barry (Peabody Museum), Manfred Ade (Museum für Naturkunde der Humboldt Universität, Berlin). I should thank all these individuals additionally for their kind hospitality. Knowing that I was traveling with young children they did their utmost to find me suitable living quarters and make my stay comfortable. While traveling through the museums I was fortunate to make the acquaintance of several colleagues. For stimulating conversations, idle chatter, and help in innumerable ways I thank: Meike Köhler, Martin Pickford, Andrew Cabrinovic, Richard Harbord, Daphne Hills, Tania King, Robert Kruczynski, Van der Made, Mike Plavcan, John Harrison, Pam, Leonard Morris (who had great tips for taking good photos), Will Harcourt-Smith, Colin Groves, Leroi Geron, Michel Tranier, Pierre Mein, Alexandra Müller, Claudia Biber, Karin Isler, Rauscher, Doris Nagel, Pantak Wein, Michel Louette, Sven Lambrecht, Peter Smedt, vi Miranda, Annelies Louage (who worked out the coordinates for the hominoid localities), Francis Kirera, Michi Solomon, Terry MacFadden, Judy Chapasco, Irene Thomas, Andrea Mess and Sylke Fernhart. I thank my in-laws for their love and constant encouragement. John and Susan Pilbrow came over all the way from Australia to help with childcare. I am especially indebted to Susan. At three important junctures in my research (once while I was in Spain and twice in New York) Susan put aside all commitments and flew over to help. I could not have done this without her sacrifice. From a mother to mother I thank her. I thank my own parents, Austin and Vidya Fernandez. Thanks to their help and support I was able to continue with my grant writing even when pregnant and then with two young children. Their love has brought me this far and their glowing pride in me will sustain me always. I thank my beloved husband, Tim. Tim played the role of mother and father to our children, chauffeur to drive me from museum to museum, porter to carry my heavy equipment, statistician to help me figure out knotty statistical problems, computer whiz and photo technician, and all this while writing his own dissertation. For his love and commitment I thank Tim. Anupama and Maya provided much-needed distractions. Always bright- eyed and cheery, they turned my data collection into an extended family vacation – evenings at the playground and weekends for sightseeing. Two sentences from this vii period are forever etched in my memory: “Are we there yet?” and “Are you done yet”. I thank my darling daughters for putting up with their busy, working mother. I thank my brothers, Aloysius and Praful, and sisters, Smita, Kalpana and Ila for understanding that I chose to come to this country to chase a dream and for believing in me and urging me on. I should especially thank Ila for her help and tireless efforts during the final stages of my writing. I thank Diana Vanlancker Sidtis for giving me a chance to work in a stimulating research environment. Working with her gave me a fresh perspective for analyzing my data. And finally, I am grateful for the friendship of Deborah Rubien, her husband, Gregg and son, Harris. Deborah and her family provided friendship and encouragement, which was vital for me, particularly during the final period when I needed that extra push. For all her help I thank Deborah. So… I am merely a link in the chain – I could not have done it without them. viii PREFACE The Blind Men and the Elephant It was six men of Indostan To learning much inclined, Who went to see the Elephant (Though all of them were blind), That each by observation Might satisfy his mind The First approached the Elephant, And happening to fall Against his broad and sturdy side, At once began to bawl: “God bless me! but the Elephant Is very like a WALL!” The Second, feeling of the tusk, Cried, “Ho! what have we here So very round and smooth and sharp? To me ’tis mighty clear This wonder of an Elephant Is very like a SPEAR!” The Third approached the animal, And happening to take The squirming trunk within his hands, Thus boldly up and spake: “I see,” quoth he, “the Elephant Is very like a SNAKE!” The Fourth reached out an eager hand, And felt about the knee. “What most this wondrous beast is like Is mighty plain,” quoth he; “ ‘Tis clear enough the Elephant Is very like a TREE!” The Fifth, who chanced to touch the ear, ix Said: “E’en the blindest man Can tell what this resembles most; Deny the fact who can This marvel of an Elephant Is very like a FAN!” The Sixth no sooner had begun About the beast to grope, Than, seizing on the swinging tail That fell within his scope, “I see,” quoth he, “the Elephant Is very like a ROPE!” And so these men of Indostan Disputed loud and long, Each in his own opinion Exceeding stiff and strong, Though each was partly in the right, And all were in the wrong! John Godfrey Saxe (1816-1887) x ABSTRACT Studying patterns of diversity and demarcating species in the paleontological context is problematic because fossil remains are fragmentary and samples are typically limited. In contrast, extant species can be potentially diagnosed using several types of data. Yet the assumption is that fossil species are equivalent to modern species. In this thesis I argue that if fossil and modern species are equivalent concepts, then patterns of variation in modern species