Dusseldorp A view to a kill The sophistication of Neanderthal behavioural strategies have been the subject of debate from the moment of their recognition as a separate species of hominin G.L. Dusseldorp in 1856. This book presents a study on Neanderthal foraging prowess. Novel ethnographic and primatological insights, suggest that increasing dependence on high quality foods, such as meat, caused the brain to evolve to a large size and thus led to highly intelligent hominins. From this baseline, the author studies the Neanderthal archaeological record in order to gain insight into the “knowledge-intensity” of Neanderthal hunting behaviour. In this research, an optimal foraging perspective is applied to Pleistocene bone assemblages. According to this perspective, foraging success is an important factor in an individual’s evolutionary fitness. Therefore foraging is organised as efficiently as possible. The prey species that were selected and hunted by Neanderthals are analysed. The author investigates economic considerations that influenced Neanderthal prey choice. These considerations are based on estimates of the population densities of the available prey species and on estimates of the relative difficulty of hunting those species. The results demonstrate that A view to a kill when Neanderthals operated within poor environments, their prey choice was constrained: they were not able to hunt species living in large herds. In these environments, solitary species were the preferred prey. It is striking that Neanderthals successfully focussed on the largest and most dangerous species in poor environments. However, in richer environments, these constraints were lifted and species living in herds were successfully exploited. In order to assess the accuracy of this approach, bone assemblages formed by cave hyenas are also analysed. The combined results of the Neanderthal and hyena analyses show that an optimal foraging perspective provides a powerful tool to increase our understanding of Pleistocene ecology. The niches of two social carnivores of similar size, which were seemingly similar, are successfully distinguished. This result lends extra credence to the conclusions regarding Neanderthal foraging strategies. This book contributes to the debate surrounding Neanderthal competence and ability. It combines an up-to-date review of current knowledge on Neanderthal biology and archaeology, with novel approaches to the archaeological record. It is thus an important contribution to the current knowledge of this enigmatic species. ISBN 978-90-8890-020-4 A view to a kill Sidestone Press Sidestone ISBN: 978-90-8890-020-4 Investigating Middle Palaeolithic subsistence Bestelnummer: SSP39370001 using an Optimal Foraging perspective 69366508 9 789088 900204 A view to a kill Sidestone Press © 2009 G.L. Dusseldorp Published by Sidestone Press, Leiden www.sidestone.com Sidestone registration number: SSP39370001 ISBN 978-90-8890-020-4 Cover Illustration: Painting by Cornelis van Dalem: ‘Landschap met het begin van de Beschaving’ (c. 1565). Museum Boijmans Van Beuningen, Rotterdam. Cover design: K. Wentink, Sidestone Press Lay-out: P.C. van Woerdekom, Sidestone Press A view to a kill Investigating Middle Palaeolithic subsistence using an optimal foraging perspective Proefschrift Ter verkrijging van de graad van Doctor aan de Universiteit van Leiden op gezag van Rector Magnificus prof. mr. P.F. van der Heijden volgens het besluit van het College voor Promoties te verdedigen op donderdag 2 april 2009 klokke 13:45 uur door Gerrit Leendert Dusseldorp Geboren op 5 augustus 1979 te Eindhoven Promotiecommissie Promotor: Prof. dr. W. Roebroeks Co-promotor: Prof. dr. R.Corbey Referenten: Prof. dr. S. Gaudzinski Prof. dr. D. Guthrie Overige leden: Prof. dr. T. van Kolfschoten Prof. dr. A. Tuffreau Dr. A. Verpoorte Contents 1 Introduction 9 1.1 Research design 10 2 Neanderthal Biology 13 2.1 Introduction 13 2.2 Neanderthal evolution 13 2.3 Neanderthal distribution patterns 16 2.4 Neanderthal anatomy and adaptation 19 2.5 Neanderthal brains 21 2.6 Neanderthal dietary niche and its implications 23 2.7 Neanderthal lives 24 2.8 Concluding remarks 27 3 Neanderthal Archaeology 29 3.1 Introduction 29 3.2 Neanderthal mobility and the study of foraging behaviour 29 3.3 Neanderthal archaeozoology 32 3.3.1 Introduction 32 3.3.2 The hunting vs. scavenging debate 32 3.3.3 Specialised hunting of ungulates 33 3.3.4 Neanderthals and megafauna 35 3.3.5 Central places: Sites exhibiting the full suite of Neanderthal foraging strategies? 37 3.3.6 Broad spectrum revolution, division of labour 42 3.3.7 Summary and conclusion 45 3.4 Material culture 46 3.5 Other aspects of Neanderthal archaeology 51 3.6 Summary and Conclusion 52 4 Optimal foraging models and Neanderthal archaeology 55 4.1 Introduction 55 4.2 General assumptions and criticism 55 4.3 An example of an OFT model: The diet breadth model 58 4.4 Applications of diet breadth in the study of the Middle Palaeolithic 60 4.5 On reconstructing the model’s parameters 61 4.6 Possible confounding factors in the archaeological record 67 4.7 Modelling Neanderthal diet breadth 69 4.8 Modelling hyena diet breadth 71 4.9 Summary and conclusion 72 5 Biache-Saint-Vaast 75 5.1 Introduction 75 5.2 The site 75 5.3 Dating 76 5.4 Stratigraphy and archaeological horizons 77 5.5 The stone artefacts 79 5.6 The bone assemblage 81 5.7 The environment 90 5.8 Applying OFT to Biache-Saint-Vaast 92 5.9 Discussion 99 5.10 Conclusion 100 6 Taubach 103 6.1 Introduction 103 6.2 The site 104 6.3 Dating 104 6.4 The archaeological finds 108 6.5 The bone collection 109 6.6 The environment 113 6.7 Applying OFT to Taubach 118 6.8 Discussion 124 6.9 Conclusion 126 7 Hyena foraging 127 7.1 Introduction 127 7.2 Hyena ecology 128 7.3 Hyena sites 130 7.4 Expectations for the study of Pleistocene hyenas 132 7.5 Case-studies 132 7.5.1 Lunel-Viel 132 7.5.2 Camiac 137 7.6 Discussion 139 7.7 Conclusion 141 8 Discussion 145 8.1 Application of the diet breadth model to archaeological data 145 8.2 Reconstructing the model’s variables 146 8.3 Modelling Neanderthals 149 8.4 This study 150 8.5 Application of OFT to Biache-Saint-Vaast and Taubach 151 8.6 The analysis in context 156 9 Conclusion 159 10 References 161 Dutch Summary 189 Acknowledgements 197 Curriculum vitae 199 1 Introduction During human evolution our primate ancestors gradually switched from a diet based mainly on plant foods to one based to a significant degree on animal foods. This shift is thought to have been im- portant in the process of human evolution, some even regard it as critical (e.g. Milton 2003). Many important developments in human evolution, like increasing brain-size and changes in life histories, are assumed to have co-evolved with the increasing contribution of animal products to the diet (e.g. Aiello and Wheeler 1995, Isaac 1978, Kaplan et al. 2000, Kaplan and Robson 2002, Kuhn and Stiner 2006). Theories that make evolutionary links between factors in human evolution, like brain size with other factors like foraging niche, longevity or group size, have focused mainly on comparative studies of primates and ethnographically known hunter/gatherers (e.g. Dunbar 1992, Hawkes et al. 1998, Kaplan et al. 2000, Reader and Laland 2002). The general validity of these theories can be evaluated by testing them on other species in the hominin family. Neanderthals are especially inter- esting in this respect; we shared a common ancestor relatively recently and their general body plan was similar to that of modern humans. More importantly, they had large brains, in terms of both absolute and relative size (Wood and Collard 1999, 69). If we accept the increasing anatomical and genetic evidence that modern humans and Neanderthals are separate species (e.g. Green et al. 2008, Hofreiter et al. 2001, McDougall, Brown, and Fleagle 2005, Ovchinikov et al. 2000, Stringer 2003, White et al. 2003), Neanderthals present us with a case of parallel evolution of large brains, the fac- tor that is deemed to be most important in human evolution. This makes the study of the factors responsible for the similar Neanderthal evolutionary trajectory germane to our understanding of human evolutionary patterns in general. In order to assess the role of the foraging niche in the evolution of large brains and prolonged life-history trajectories, this thesis aims to assess the efficacy of Neanderthal foraging strategies, specifically Neanderthal meat procurement. It has been argued that hunting is a very knowledge- intensive activity practised by modern humans (e.g. Kaplan et al. 2000). It is also seen as the most “complex” activity that chimpanzees (Pan troglodytes) engage in (e.g. Boesch 2003). Therefore, study- ing the knowledge-intensity of the Neanderthal foraging niche and its evolutionary implications for the evolution of larger brains has bearing on the theories as proposed by Kaplan et al., among others since it may yield insight in the validity of their underlying assumptions. The importance of meat in the lives of our ancestors has been the subject of heated debate within Palaeolithic archaeology. First, the crucial contribution that gathering provides to the diets of many extant hunter/gatherers has been stressed (e.g. Dahlberg 1981). Second, the question of how meat was obtained has been debated heatedly (for a review see Domínguez-Rodrigo 2002). Hunting is seen as complicated behaviour and this, combined with meagre archaeological evidence has led some researchers to suggest that only anatomically modern humans (AMH) were able to subsist by regularly hunting large mammals (e.g. Binford 1984). Scientists studying Pleistocene ecology on the other hand, have stated that subsisting primarily by scavenging was not a viable option for hominins and hence predicted hunting as the basis of subsistence (Geist 1978, Tooby and DeVore 1987).