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The official electronic file of this thesis or dissertation is maintained by the University

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Ecogeographic Variation in Neandertal Dietary Habits:

Evidence from Microwear Texture Analysis

A Dissertation Presented


Sireen El Zaatari


The Graduate School

in Patrial fulfillment of the Requirements for the Degree of Doctor of Philosophy in Anthropology Physical Anthropology Stony Brook University May 2007 Copyright by Sireen El Zaatari May 2007 Stony Brook University The Graduate School Sireen El Zaatari We, the dissertation committee for the above candidate for the Doctor of Philosophy degree, hereby recommend acceptance of this dissertation.

Fredrick E. Grine, Chairman Dissertation Advisor Department of Anthropology William Jungers Chairperson of Defense Department of Anatomical Sciences John Shea Member Department of Anthropology Peter S. Ungar, Professor Outside Member University of Arkansas Department of Anthropology Jean-Jacques Hublin, Professor Outside Member Max-Planck Institute for Evolutionary Anthropology Department of Human Evolution This dissertation is accepted by the Graduate School Lawrence Martin Dean of the Graduate School ii


of the Dissertation

Ecogeographic Variation in Neandertal Dietary Habits:

Evidence from Microwear Texture Analysis by Sireen El Zaatari Doctor of Philosophy in Anthropology Physical Anthropology Stony Brook University 2007 For over 100,000 years, Neandertals inhabited a variety of ecological zones across western Eurasia, between glacial and interglacial conditions. To elucidate the still poorly understood effects of climatic change and variability, and possible competition on the Neandertal subsistence patterns, this study employed dental microwear texture analysis to reconstruct the dietary habits of 54 Neandertal, Pre-Neandertal, and early Upper Paleolithic (EUP) modern human specimens from 28 sites in western Eurasia. Microwear signatures of seven modern hunter-gatherer groups (n = 155) of known and diverse diets were analyzed for comparative purposes.

Microwear signatures of Neandertals and Pre-Neandertals are correlated with variation in vegetation-cover, such that individuals from cold-steppe/tundra vegetation had less complex microwear surfaces than those from forested environments. The microwear pattern of the EUP fossils did not differ significantly from those of the Pre

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the former had a more varied diet. However, significant differences in microwear signatures were found between the southern European Neandertals and the EUP fossils.

In accord with the stable isotope results, microwear analysis classifies Neandertals as toplevel carnivores. However, dental microwear analysis detected some subtle dietary differences. Thus, the microwear signatures of Neandertals and Pre-Neandertals from steppe/tundra vegetation are similar to meat-eating Fuegians from comparable habitats, whereas those of Neandertals and pre-Neandertals from forested environments resemble the Chumash, who inhabited a Mediterranean-like environment. Neandertals from the deciduous forests of southern Europe have a microwear signature that falls within the ranges of Australian and African aborigines with mixed diets. EUP fossils have microwear signatures that resemble those of both the modern Chumash and Fuegians.

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I would like to thank my advisor, Prof. Fredrick Grine for his guidance, motivation and support through all these years, especially the last one. I would also like to thank the rest of my committee members, Profs. William Jungers, John Shea, Peter Ungar, and Jean-Jacques Hublin for their advice and valuable comments that helped shape this project.

This dissertation would not have been possible without access to museum collections. I gratefully acknowledge the following museum curators: Almudena Arellano and Pierre Elie Moullé (Musée de Préhistoire Régionale, Menton, France), James Brink and Zoe Henderson (National Museum Bloemfontein, Bloemfontein, South Africa), Angiolo Del Lucchase (Museo Preistorico dei Balzi Rossi, Ventimiglia, Italy), Vincenzo Formicola (Dpt. di Etologia, Ecologia ed Evoluzione Università di Pisa, Pisa, Italy), Almut Hoffmann (Museum für Vor- und Frühgeschichte, Berlin, Germany), Rob Kruszynski, Chris Stringer and Louise Humphrey (London-NHM), Giorgio Manzi (Dipartimento di Biologia Animale e dell'Uomo, Universita' di Roma "La Sapienza", Rome, Italy), Alan Morris (Cape Town University, Cape Town, South Africa), David Morris (McGregor Museum, Kimberley, South Africa), Ken Mowbray (New YorkAMNH), Ildiko Pap (Hungarian Natural History Museum, Budapest, Hungary), Catherine Schwab (Musée des Antiquités Nationales, St. Germain-en-Laye, France), Maria Teschler-Nicola (Naturhistorisches Museum, Vienna, Austria), and Reinhard Ziegler (Staatliches Museum für Naturkunde, Stuttgart, Germany). I also thank Alejandro Pérez-Pérez for giving me access to his collection of dental molds and Yoel Rak for making molds of the Levantine specimens. I am grateful for Rob Scott for providing technical assistance.

I extend my thanks to all the professors at both the Anthropology and Anatomy Departments at Stony Brook University, especially Prof. Elizabeth Stone for her continuous support. I also thank Jean Moreau for making me feel that I have family when I was so far away from my family. Thank you to my fellow graduate students at the Anthropology Department and to my friends in the USA especially Rania Roushdy, Tereza Capelos, Melixa Abad-Izquierdo, Stephen Patnode, Lina Mahmoud, Helen Malko, Ghufran Ahmad, Zaid El Rawi, Saad Al-Rubayaa, Danielle Royer, Anthony Olejniczak, Tanya Smith, Mark Coleman, Amanuel Beyin, David Fernandez, Mathew Banks, and Heather Smith, and all my friends back home in Lebanon.

I owe special thanks to Christos Sourdis for making the past eight years of my life so interesting. Finally, I offer profound thanks to my family, my grandmother, aunts, uncles, cousins and, of course, my parents and brother for their endless love, motivation, and encouragement.

This research was supported by the National Science Foundation, the Leakey Foundation, the Ruggles-Gates Fund (Royal Anthropological Institute of Great Britain and Ireland), Interdepartmental Doctoral Program in Anthropological Sciences Research

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Temporal and geographical spread of the Neandertals Neandertals were very successful throughout western Eurasia for more than a hundred thousand years before they suddenly disappeared around 28 kyr (Zilhão 2001).

Over this period of time, the Neandertals had to cope with sometimes harsh and severely changing environments over oxygen isotope stages (OIS) 6 through 3 (van Andel and Tzedakis 1996). Nevertheless, at the beginning of OIS 2, they were effectively replaced by more modern human populations. Documenting the dietary habits of the Neandertals through time and across the various glacial and interglacial periods is the key to understanding their capabilities to adapt and survive for such a lengthy period. It has been argued that modern humans were behaviorally superior to the Neandertals and outcompeted them for resources. A study of the Neandertal dietary variability over time is essential to determine whether there was any evidence of dietary shifts that could be related to competition at the time of the advent of modern humans in Eurasia.

Brief introduction to the climatic changes during the Ice Ages in Europe Neandertals lived during the alternating glacial/interglacial cycles of the Middle and Late Pleistocene. A clear picture of the climate and environment of western Eurasia

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Shackleton 1969; Shackleton 1975; Shackleton 1977; Shackleton 1987), the land based ice cores records (1992; Johensen et al. 1972; 1993; Jouzel et al. 1987; 1989), and long pollen sequences from a number of European and Middle Eastern sites (Behre and van der Plicht 1992; Cheddadi and Rossignol-Strick 1995; de Beaulieu and Reille 1984;

1992a; 1992b; Follieri et al. 1998; Guiot et al. 1989; Mangerud et al. 1981; Pérez-Obiol and Juliá 1994; Pons and Reille 1988; Reille and de Beaulieu 1990; Rossignol-Strick and Planchais 1989; Tzedakis 1993; Tzedakis 1994; Tzedakis and Bennett 1995; van Zeist and Bottema 1991; Watts 1985; Wijmstra 1969; Woillard 1978a; Woillard and Mook 1982). Climatic reconstructions show that during any single OIS, western Eurasia was divisible into several climatic zones and that the climate of any one area varied between glacial and interglacial times. The fossil and archaeological records indicate that the Neandertals were able to survive in most of these western Eurasian Ice Age environments. Their remains are associated with warm forested environments as well as more cold and open ones. Changes in vegetation cover in response to climatic changes must have certainly affected food availability. During glacial times, plant and animal species in the areas mostly affected by the cold (i.e., northern Europe) became extinct.

This is not the case in the warmer southern/Mediterranean part of the continent, which is characterized by continuity in the plant and animal records (Tzedakis and Bennett 1995).

During interglacial periods, there was colonization by new species and re-colonization from refugia by older community members of plants and animals from the south. These colonization and re-colonization events resulted in the mixing of elements from steppe and forest, creating strange associations of animals in northern Europe and giving each

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1988). All interglacials, however, followed a broadly similar succession for forest regeneration. The Neandertals would have had to adapt to these changes to be able to survive. However, the role that these changes played in their diets is still unclear. Food availability strongly affects the social structure of a group, as well as population density, settlement patterns, and the territory that a group occupies. Thus, the effects of the changes in the environment on the Neandertal life might be addressed by examining dietary variation among Neandertal groups from different ecological zones.

Brief introduction to the previous studies on Neandertal diets The dietary habits of the Neandertals are still poorly understood. Reconstructions have been attempted through the analysis of associated Middle Paleolithic faunal assemblages (Auguste 1992; Binford 1988; Chase 1986; Chase 1989; Gaudzinski 1995;

Gaudzinski 1998; Gaudzinski and Roebroeks 2000; Hoffecker et al. 1991; Jaubert et al.

1990; Marean and Kim 1998; Mellars 1996; Munson and Marean 2003; Stiner 1994) and plant remains (Hardy 2004; Lev et al. 2005; Madella et al. 2002). However, the faunal assemblages studied have concentrated on the mode of exploitation (hunting versus scavenging) of the Neandertals. Very few studies have looked at plant exploitation during the Middle Paleolithic. Also, proving the dietary use of these plants has been shown to be very difficult if not almost impossible. Dietary reconstructions also have been attempted by analogy through the study of cranial morphology (Brace 1964; Coon 1962; Demes 1987; Franciscus and Trinkaus 1995; Howells 1975; Martin 1923; Puech

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Trinkaus 1992; Ungar et al. 1997; Wallace 1975), but linking morphological factors directly to dietary adaptations is fraught with difficulty. Studies of dental enamel hypoplasia have also been used to infer the amount of nutritional stress that some Neandertal groups faced (Cunha et al. 2004; Guatelli-Steinberg et al. 2004; Hutchinson et al. 1997; Molnar and Molnar 1985; Oglivie et al. 1989; Skinner 1996). All of these studies have treated the Neandertals as a single group, failing to consider any possible dietary differences resulting from different ecogeographic conditions. Stable isotopes in Neandertal bones have also been examined (Balter and Simon in press; 1999; 2001;

Bocherens and Drucker 2003; Bocherens et al. 2005; Bocherens et al. 1991; Fizet et al.

1995; Richards et al. 2000; Schoeninger 1982), but these analyses were based on samples from a limited number of sites. Finally, buccal microwear of various Neandertal teeth has been analyzed in an attempt to better understand their dietary habits (Lalueza-Fox and Pérez-Pérez 1993; Lalueza-Fox et al. 1993; Lalueza-Fox et al. 1996; Pérez-Pérez et al. 2003). However, the usefulness of this technique in dietary reconstructions is still uncertain.

Brief description of the potential of dental microwear analysis Numerous studies have demonstrated the potential of occlusal dental microwear to yield direct evidence of tooth use and diet in both extant as well as extinct species (Daegling and Grine 1999; El Zaatari et al. 2005; Grine 1986; 1987; Grine and Kay 1988;

Grine et al. 2006; Ryan 1981; Ryan and Johanson 1989; Scott et al. 2006; Scott et al.

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Teaford and Robinson 1989; Teaford and Walker 1984; Ungar 1994; 1996; Ungar et al.

2006; Walker 1976; Walker et al. 1994). Although variations in microwear patterns can be attributed to various factors [e.g., the characteristics of the abrasives ingested with food (Daegling and Grine 1999; Teaford and Glander 1991), enamel microstructure (Maas 1991), and properties of jaw biomechanics (Gordon 1982)], the mechanical properties of the diet itself have been considered to be the primary cause of these patterns. Studies have shown that specific diets and ingestive behaviors leave characteristic patterns of microscopic scratches and pits on extant primate teeth. Occlusal microwear analysis has been shown to be a useful tool for detecting slight differences in diets of closely related species (e.g., Teaford 1986; 1993). Thus, occlusal microwear has the capability to differentiate short-term variations in diet (Teaford and Oyen 1989a).

The established methods of occlusal dental microwear analysis have now been enhanced by the analysis of three-dimensional surface thus increasing accuracy, objectivity and repeatability of the measurements (Scott et al. 2006; Scott et al. 2005; Ungar et al. 2003).

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