RED DEER: THEIR ECOLOGY AND HOW THEY WERE HUNTED BY LATE PLEISTOCENE HOMINIDS IN WESTERN EUROPE A DISSERTATION SUBMITTED TO THE DEPARTMENT OF ANTHROPOLOGICAL SCIENCES AND THE COMMITTEE ON GRADUATE STUDIES OF STANFORD UNIVERSITY IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY Teresa Eleanor Steele August 2002 © Copyright by Teresa Eleanor Steele 2002 All Rights Reserved ii Abstract Fossil hominid morphology, archaeology, and genetics indicate that in Europe 30,000-40,000 years ago, anatomically modern humans and their Upper Paleolithic industries replaced Neandertals and their Middle Paleolithic tools. Neandertals had thrived for hundreds of thousands of years, so why were they replaced? One possibility is that modern humans were able to extract more resources from the environment. This dissertation tests this explanation by assessing variation present in ancient hunting practices and investigating the relationship between Late Pleistocene hominids, tool industries, and hunting. I examined the hunting of one species, red deer (Cervus elaphus), through time and across spaceusing prey age-at-death as an indicator of hunting strategy. In the process, I evaluated the ability of the Quadratic Crown Height Method to accurately assign age-at-death; compared how well histograms, boxplots, and triangular graphs reconstruct mortality profiles from fossil assemblages; and developed a novel method for statistically comparing samples on triangular graphs. My results show that Neandertals and modern humans did not differ significantly in their ability to hunt prime-age red deer. None of the mortality distributions from the archaeological samples resemble the distribution constructed from elk killed by wolves in Yellowstone National Park, Wyoming. Like other carnivores, wolves usually take young, old, and infirm prey. Nevertheless, the samples included in this study show a shift in prey age-at-death during the Middle Paleolithic approximately 50 kya. Young adult prey are more abundant in recent assemblages than in more ancient assemblages. Over 25 archaeological samples from western Europe contribute to these conclusions, making this iv dissertation the most comprehensive study of Pleistocene hunting to date. More well- dated samples are needed, however, to confirm these results. Because red deer skeletal and tooth size fluctuated across my samples, I investigated the relationship between climate and C. elaphus size to determine if body size could indicate paleoclimates. In modern North American specimens, distal metatarsal breadth has a good relationship with climate, and tooth breadth has a similar but weaker relationship. The modern European data do not relate clearly to climate. Fossil red deer are larger during glacials than interglacials, but additional data are needed to better define patterns. v to my parents Gary and Nancy Steele and my grandmother Helen Steele vi Acknowledgements Many people contributed to this dissertation in innumerable ways, and I am grateful to all of them. First and foremost, I would like to thank my primary advisor, Richard Klein. I am very appreciative of his generosity with his time, advice, data, and references, to name a few of his contributions. Without his support, this project would not have been possible. John Rick and Elizabeth Hadly both brought unique perspectives to my research, enriching it greatly. Their encouragement and enthusiasm were important for the completion of this project. William Durham, Robert Franciscus, Jean-Jacques Hublin, and Joanna Mountain provided valuable discussions and training during my graduate career. My research required the support of many museums in Europe and the United States. I am grateful to all of the people at these institutions who not only provided access to collections under their care but also valuable discussions about the material. In particular, I would like to thank these individuals: Françoise Delpech, Jean-Philippe Rigaud, and Dominique Armand (l’Institut de Préhistoire et Geologie du Quaternaire, Université Bordeaux I, Talence); Henry de Lumley (l’Institut de Paléontologie Humaine, Paris); Patricia Valensi and Annie Echassoux (Laboratoire Départemental de Préhistoire du Lazaret, Nice); Anne-Marie Moigne and José Moutoussamy (Centre Européen de Recherches Préhistoriques de Tautavel, Tautavel); Roland Nespoulet and Laurent Chiotti (Musée de l'Abri Pataud, Les Eyzies de Tayac); Patrick Auguste (Laboratoire Préhistoire et Quaternaire, Université des Sciences et Technologies de Lille, Villeneuve d'Ascq); Andy Currant (Deptartment of Palaeontology, British Natural History Museum, London); vii Paula Jenkins (Mammal Group, Department of Zoology, British Natural History Museum, London); Elaine Turner (Römisch-Germanisches Zentralmuseum Mainz, Neuwied); Nicholas Conard, Hans-Peter Uerpmann, Beata Kozdeba, and Petra Kroneck (Abt. Ältere Urgeschichte und Quartärökologie and Institut für Ur- und Frühgeschichte, Universität Tübingen, Tübingen); Antonio Tagliacozzo, Ivana Fiore and M. A. Fugazzola (Museo Preistorico-Etnografico "L. Pigorini", Rome); Jesús Altuna and Koro Mariezkurrena (Departamento Prehistoria, Sociedad de Ciencias Aranzadi, San Sebastián/Donostia); Vicente Baldellou and Isidro Aguilera (Huesca Musuem, Huesca); Pilar Utrilla (Departamento de Ciencias de la Antigüedad, Universidad de Zaragoza, Zaragoza); Enrique Tessier (Museo Arqueológico de Asturias, Oviedo); Christopher Conroy and Barbara Stein (Museum of Vertebrate Zoology, University of California, Berkeley); Douglas Long (Dept. of Ornithology and Mammalogy, California Academy of Sciences, San Francisco); David Dyer (Philip L. Wright Zoological Museum, University of Montana, Missoula); Douglas Smith (National Park Service, Yellowstone National Park); Ken Hamlin (Montana Dept. Fish, Wildlife, and Parks, Bozeman); and Lynn Irby (Fish and Wildlife Management Program, Montana State University, Bozeman). Many additional colleagues provided valuable information that helped organize my data collection, and I am grateful to all of them for their assistance. In particular, I would like to thank Donald Grayson, Anne Pike-Tay, and Lawrence Straus. Adrian Lister, Marylène Patou-Mathis, Fernanda Blasco, and Rivka Rabinovich provided useful advice during my data collection. Federico Bonelli, William Harcourt-Smith, Brian and Lorna Kerr, Beata Kozdeba, Elaine Turner, and Robin Wyss kindly allowed me to stay in their homes during my data collection trip. Kathryn Cruz-Uribe, Richard Klein, Adrian viii Lister, Anne Pike-Tay (with kind permission from F. Bernaldo de Quiros and V. Cabrera), and Cornelia Wolf generously provided measurements of C. elaphus specimens. Richard Klein also permitted me to use many of his illustrations and his Kolmogorov-Smirnov program. The Louis S. B. Leakey Foundation, the Andrew W. Mellon Foundation, and the Stanford University Department of Anthropological Sciences generously provided financial support for this research. I am appreciative of my fellow graduate students, Elizabeth Burson, Matt Jobin, Silvia Kembel, Tim King, Richard Pocklington, Charles Roseman, and especially Deborah Stratmann, for making my time at Stanford a more enjoyable experience. I give my heartfelt thanks to Beth Alcalde, Dee Bardwick, Rose Bressman, Jannie Chan, Becky Post Gibbons, Amy Kashiwabara, Jen Lustig, Richard McElreath, Jen McGlone, Michael Osbourne, Hermun Puri, Tina Villanova, and Sarah Wyckoff for their support throughout the years. I also would like to acknowledge Michelle Lampl, Alan Mann, Janet Monge, and Ina Jane Wundram for encouraging my interest in paleoanthropology. I would like to extend my deepest gratitude to my family: my parents Nancy and Gary Steele, my brother Michael Steele, my grandmother Helen Steele, and my Aunt Mary McManus. They always have provided unwavering love and encouragement. Thank you for believing in me. Finally, I would like to express my deepest gratitude to Tim Weaver for everything from technical support to emotional support. I could not have done it without you. Thank you for being there for me from the very beginning. ix Table of Contents Abstract ........................................................................................................................iv Acknowledgements......................................................................................................vii Table of Contents ..........................................................................................................x List of Tables ...............................................................................................................xv List of Illustrations....................................................................................................xvii Chapter 1: Introduction, background, and research objectives..................................1 The Neandertals ..........................................................................................................2 Modern human origins ................................................................................................4 The evidence for replacement ..................................................................................6 Evidence against a complete replacement..............................................................11 The origins of modern behavior.............................................................................12 How did this replacement occur?...............................................................................14