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Department of Human Evolution - Max Planck Institute for Evolutionary Anthropology

Daniel S. Adler and Guy Bar-Oz. Seasonal Patterns of Prey Acquisition During the Middle and Upper Palaeolithic of the Southern Caucasus

Zeresenay Alemseged & Rene Bobe. Diet in the earliest hominins: a paleoenvironmental perspective

Hervé Bocherens. Neanderthal Dietary Habits: Review of the Isotopic Evidence

A. Brooks. The Importance of Fish in Human Diets -- What is the Early Evidence?

Steven E. Churchill and Jill A. Rhodes. The Evolution of the Human Capacity for “Killing at a Distance”: The Human Fossil Evidence for the Evolution of Projectile Weaponry

Sabine Gaudzinski-Windheuser. Middle Palaeolithic Subsistence in a Diachronous Perspective
John F. Hoffecker. Neanderthal and Modern Human Diet in Eastern Europe
Gottfried Hohmann, Sylvia Ortmann Diet composition, digestive strategies and food acquisition in nonhuman primates: A review of recent studies and their possible implications for the evolution of human diet

Martin Jones. Plant Evidence for Subsistence in the Middle and Upper Palaeolithic of Europe

Richard G. Klein. Animal Remains from Archaeological Sites Suggest that Later Stone Age People Foraged more Effectively than their Middle Stone Age Predecessors

Staffan Lindeberg. Modern Human Physiology with Respect to Evolutionary Adaptations That Relate to Diet in the Past

Peter Lucas. The Relationship Between the Human Dentition and Diet During the Paleolithic

K. MacDonald, W. Roebroeks, A. Verpoorte. An Energetics Perspective on the Neandertal Record

Natalie D. Munro. Epipaleolithic Subsistence Intensification in Southwest Asia: The Faunal Evidence

M. P. Richards. Isotopic evidence for European Upper Palaeolithic human diets

Margaret J. Schoeninger. Hair δ 13 C values reflect aspects of primate ecology

John Shea. The Ecological Impact of Projectile Weaponry in Late Pleistocene Human Evolution

J. Josh Snodgrass, William R. Leonard, and Marcia L. Robertson. Brain Size Change as Related to Diet in the Earliest Hominids

Matt Sponheimer. Biogeochemical Evidence for the Ecology of Australopiths and Early Homo

Teresa E. Steele. Late Pleistocene Subsistence Strategies and Resource Intensification in Africa

Mary C. Stiner. Trends in the Economic and Demographic Resilience of Paleolithic Forager Societies in the Mediterranean Basin

Paola Villa. Lower and Middle Paleolithic Faunal Exploitation in Europe


Seasonal Patterns of Prey Acquisition During the Middle and Upper Palaeolithic of the Southern Caucasus

Daniel S. Adler1 and Guy Bar-Oz2

1 Department of Anthropology, University of Connecticut U-2176, Storrs, CT 06269, USA.
2 Zinman Institute of Archaeology, University of Haifa, Mount Carmel, 31905 Haifa, Israel.

Zooarchaeological and taphonomic analyses provide essential background information for the discussion of Late Middle Palaeolithic (LMP) and Early Upper Palaeolithic (EUP) patterns of mobility, land-use, and hunting within the southern Caucasus, a topographically and environmentally diverse region circumscribed by a variety of natural barriers to human mobility. Recent research documented the targeting of prime-age adult Caucasian tur (Capra caucasica) by LMP populations and the organization of hunting activities according to this species’ specific migratory behaviors, which made them locally abundant on a seasonal basis (Bar-Oz and Adler 2005; Adler et al. 2006). Our analyses suggest that Neanderthals and modern humans were equally capable of learning the feeding, mating, migratory, and flight behaviors of Caucasian tur, as well as acquiring and exploiting key biogeographical information pertaining to species habitat preference and the seasonal distribution of resources. In these respects there appear to have been few alterations in medium/large game hunting practices between the LMP and EUP, with ungulate species abundance in the entire stratigraphic sequence of Ortvale Klde reflecting seasonal fluctuations in food supply rather than specialization, or differences in hunting ability or technology (see also Stiner 2002; Grayson and Delpech 2003). Consequently, we argue that such hunting behaviors are poor proxies for behavioral modernity and the Middle—Upper Palaeolithic Transition.

This is not surprising if we accept that Neanderthals and modern humans were the top predators in whatever environments they occupied, and given this position in the prey-predator hierarchy that each population exploited many resources in a similar manner. However, following the principle of competitive exclusion (Begon, Harper, and Townsend 1996), we contend that when these two populations encountered one another within a given environment there was little chance for coexistence; two populations in direct competition for identical limited resources cannot remain sympatric. Further advantage was to be gained by modern human populations exploiting a wider array of resources (Stiner, Munro, and Surovell 2000; Hockett and Haws 2005), however such data are currently lacking in the southern Caucasus.

From a purely ecological perspective it would appear that the transition from the LMP to the EUP in the southern Caucasus represents the replacement of one top predator (Neanderthals) by another (modern humans), with little discernable shift in the predator-prey hierarchy or the medium/large game species exploited. In terms of resource and niche preference we suggest that Neanderthals and modern humans were sympatric to the point of exclusion.

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Diet in the earliest hominins: a paleoenvironmental perspective

Zeresenay Alemseged1 & Rene Bobe2

1 Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany

2 Department of Anthropology, State University of New York at Buffalo, USA

Understanding the diet of a species is critical to our comprehension of its paleobiology as well as the selection forces that were responsible for shaping its morphology and behavior. It is known to all that the fossil evidence for the earliest members of our family is scarce. As a result we have only limited knowledge of their paleoecology as the dietary and paleoenvironmental reconstructions of these species are fragmentary. Yet, considerable efforts have been put into addressing problems pertaining to early hominin diet based on dental gross morphology, dental microstructure, dental topography, biomechanics, and isotopic evidence, with encouraging results. On the other hand, the paleoenvironmental context of these species has been addressed using associated fauna and in some cases sedimentary and isotopic evidence. However, these paleoenvironmental data are rather static, site-based reconstructions that are highly sensitive to taphonomic and other biasing factors. In most cases early hominin species are attributed to a woodland/forested mosaic environment without specific implication for their probable preferences. In this paper we review both lines of evidence (dietary and paleoenvironmental) and show that hominin environmental preferences can be addressed using associated bovid taxa. This is done using data that come from the Omo Shungura Formation, which is well constrained chronologically and taphonomically. Results of dietary reconstructions obtained using diverse approaches can then be evaluated in light of hominin habitat preference.

Neanderthal Dietary Habits: Review of the Isotopic Evidence

Hervé Bocherens 1,2

 1 Institut des Sciences de l'Evolution, Université Montpellier 2, France

2 Institute für Ur- und Frühgeschichte und Archäologie des Mittelalters, Universität Tübingen, Germany

Carbon and nitrogen isotopic abundances of fossil bone collagen reflect those of the average diet, and they can be preserved for tens of thousands of years under favorable conditions. Contrarily to Charentian specimens dating from 150,000 to 120,000 years, other European Neanderthal bones ranging in age from 100,000 to 28,000 have yielded reliable collagen. For this well-preserved collagen, isotopic signatures offer the possibility to reconstruct the dietary habits of Neanderthals. The degree of interpretation of the isotopic results depends on the palaeoecological context, especially on the knowledge of the available food resources and their isotopic signatures. Animal bones associated with the studied human remains provide the most reliable source for such information. In addition, isotopic data can be retrieved from nearby sites of similar age when they lack in the hominid site. However, the precision of the interpretation decreases when distance and age difference between hominids and fauna increases.

his presentation will illustrate how such isotopic investigations have impacted our understanding of Neanderthals' dietary habits. A critical review of the available data will be presented, with a discussion of some methodological points, such as preservation assessment and quantification of consumed proteins resources. Comparisons of prey selection patterns based on isotopic results will be made between Neanderthals and animal predators, such as hyaenas. It will be attempted to evaluate the variation of Neanderthal subsistence strategies through time and space. Finally, the possible implications of dietary strategies on the final demise of Neanderthals will be discussed in view of the isotopic results obtained on late European Neanderthals.

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The Importance of Fish in Human Diets -- What is the Early Evidence?

A. Brooks

George Washington University/ Smithsonian, Washington D.C., USA

Fish can be an important element in human diets, especially since they often contain long-chain fatty acids, particularly docasohexanoic acid, not widely present in pre-agricultural terrestrial food sources. Since long-chain lipids are a major constituent of brain tissue, these long-chain fatty acids in human diets are thought to improve brain growth. In addition, concentrations of fat-soluble vitamins in fish liver and other organs can compensate for low levels of vitamins A and D. Yet the archaeological evidence for the origins of fishing suggest a fairly late onset of use of this resource by humans. This paper will review the evidence from the African Middle Stone Age sites at Katanda and Aduma, with some comments on the evidence for fishing at other Middle Stone Age and Middle Palaeolithic sites. Problems of documenting this practice archaeologically will also be addressed, along with the implications of fishing for human subsistence and settlement patterning.

The Evolution of the Human Capacity for “Killing at a Distance”: The Human Fossil Evidence for the Evolution of Projectile Weaponry

Steven E. Churchill and Jill A. Rhodes

Department of Biological Anthropology and Anatomy, Box 90383, Duke University, Durham, NC USA

 When in evolutionary history did long-range projectile weapons become an important component of hunting tool kits? Recent analyses of Middle Stone Age and Middle Paleolithic points suggest that true long-range projectile weaponry – most likely in the form of spearthrower-delivered darts - evolved in Africa sometime between 90-70 kya, and was part of the tool kit of modern humans who expanded out of Africa after this time. This possibility has important implications for our understanding of behavior change during the MSA, the evolution of modern human predatory behavior and subsistence strategies, and the nature of the competitive interactions that occurred between modern humans and the archaic humans they encountered on their diaspora from Africa

The origins of projectile weaponry can be addressed, in part, through analyses of the skeletal remains of the prehistoric humans who made and used them, since habitual behavior patterns – especially those involving skeletally stressful actions like forceful throwing – can be imprinted on the skeleton through both genetic and epigenetic pathways. Previous studies of humeral diaphyseal cross-sectional geometry in Neandertals and early modern Europeans have concluded that habitual, forceful throwing is reflected in the fossil record only after 20 kya – well after modern humans had invaded Europe. Other aspects of the skeleton that might contain a signature of habitual throwing have not yet been examined. Recent studies in the field of sports medicine indicate that throwing athletes have increased humeral torsion angles in their throwing arms, and a greater degree of bilateral asymmetry in torsion angles than do non-throwers. Other skeletal evidence may be found in the forearm and shoulder. Bilaterally asymmetric hypertrophy of the ulnar supinator crest has been associated with throwing behavior in Prehistoric hunter-gatherer societies of South Asia. Additionally, the morphology of the scapular glenoid fossa may reflect adaptations to stresses generated during forceful throwing. A reduced glenoid index, such as that previously identified in Mousterian-associated fossil humans (Neandertals and possibly Levantine early modern humans), suggests a shoulder joint poorly-adapted to withstand dorsally- and ventrally-directed forces at the shoulder that occur during throwing.

Here we examine the magnitude of, and asymmetry in, humeral torsion angles, supinator crest morphology and glenoid fossa breadth in Middle and Upper Paleolithic fossil hominins and comparative samples of North American recent humans relative to the question of the origin of projectile weaponry. Although the results are not clear-cut, the overall pattern of osteological indicators is consistent with the claim that projectile weapons arose in the African later MSA and moved into Europe in the hands of modern humans.  

 Middle Palaeolithic Subsistence in a Diachronous Perspective

Sabine Gaudzinski-Windheuser 1, 2

1 Römisch-Germanisches Zentralmuseum Mainz, Palaeolithic Research Unit
2 Johannes Gutenberg-Universität Mainz

Current models concerning behavioural implications derived from Middle and Early Upper Pleistocene archaeological data call strongly for a determined hunting perspective. The current state of knowledge implies that the European Middle Palaeolithic saw a variety of exploitation tactics which were imbedded in broader settlement systems. In a diachronous perspective this evidence will be evaluated against the Middle and Upper Pleistocene faunal record from Lower and Upper Palaeolithic European archaeological sites in order to assess whether Neanderthals` anatomical uniqueness is reflected in their subsistence behaviour.

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Neanderthal and Modern Human Diet in Eastern Europe

John F. Hoffecker

Institute of Arctic and Alpine Research, University of Colorado at Boulder, Boulder, Colorado USA

The Late Pleistocene environments of Eastern Europe—especially the cool and dry environments of the central East European Plain—offer a unique setting for comparison of Neanderthal and modern human diet. There are some taphonomic factors specific to this setting, however, including those related to the scarcity of natural shelters and distribution of woody plants on the central plain, which complicate the analysis of Late Pleistocene human diet. Neanderthals occupied southern upland areas, and sometimes the central plain, probably sustaining themselves primarily on the hunting of large mammals, which often included steppe species such as Bison priscus and Saiga tatarica. By contrast, modern humans—who are present on the East European Plain as early as anywhere in Europe—broadened the diet to include small mammals, birds, and/or fish during the Middle Pleniglacial. Evidence of this shift, which seems to have been achieved through the design of novel food-getting technologies, is derived both from the faunal remains and stable isotope analysis of human bone. Modern humans occupied loess-steppe habitat on the central plain during the Late Pleniglacial, but probably consumed much of their food debris as fuel.

Diet composition, digestive strategies and food acquisition in nonhuman primates: A review of recent studies and their possible implications for the evolution of human diet

Gottfried Hohmann 1 & Sylvia Ortmann 2

1 Max-Planck-Institute for Evolutionary Anthropology, Dept. Primatology, Deutscher Platz 6,
D-04103 Leipzig

2 Institute for Zoo and Wildlife Research, Alfred-Kowalke-Str. 17 10315 Berlin

With a few exceptions, cercopithecine primates live on a diet that is dominated by plant items. In contrast to other vertebrates, primate omnivory is characterized by a wide spectrum of plant foods items and a narrow spectrum of animal food such as insects. The plant dominated diet composition is well reflected by anatomical, physiological, and behavioural features including (a) cheek pouches, (b) dental features facilitating mechanical access to physically protected items (e.g. strong fibres, hard fruit shells), (c) multi-chambered stomachs, (d) enlarged hindguts, (e) long retention times, (f) high tolerance towards toxic substances, and (g) diverse modes of food processing. Studies on hominid primates indicate extensive overlap with cercopithecines in terms of the nature of food sources. Plants provide the major nutritional source and insects are the most common supplement. Chimpanzees appear to deviate from the typically hominid life style as they occasionally hunt and consume the meat of other vertebrates. In spite of the similarity in dietary patterns between cercopithecines and hominids, anatomical and physiological adaptations commonly found in cercopithecines seem to be absent: Hominid primates lack cheek pouches, the gastro-intestinal tract is short and simple without significantly enlarged compartments, retention time is shorter than predicted by body mass, and the use of food processing techniques is rare and restricted to few items. This discrepancy between similarity of diet composition on one hand, and differences in digestive anatomy on the other is unexpected and requires explanation. Recent studies on bonobos and chimpanzees suggest that hominids use often the same food items as cercopithecines but in a different way, and that these differences in resource exploitation affect diet composition. In addition, new data demonstrate that carnivory is not confined to chimpanzees but applies equally to bonobos, and, perhaps to a lesser degree, also to orangutans. We argue that the observed differences in use of food sources and differences in meat consumption between cercopithecines and hominids reflect a shift in the mode of energy extraction from fermentation to non-fermentive nutrient absorption. It is tempting to speculate that the changes in the digestive strategy might be associated with increasing body mass and/or a more efficient mode of locomotion, but a better understanding of the evolution of the digestive system of hominid primates requires further research. However, the way bonobos and chimpanzees process and distribute food items offer new and interesting perspectives on the diet composition of human primates.

Animal Remains from Archaeological Sites Suggest that Later Stone Age People Foraged more Effectively than their Middle Stone Age Predecessors

Richard G. Klein

Program in Human Biology, Stanford University, Stanford, CA 94305, USA

Artifacts suggest that Later Stone Age (LSA) people were behaviorally advanced over their Middle Stone Age (MSA) predecessors, and an advance in the ability to hunt and gather could especially explain how LSA people (or their Upper Paleolithic descendants) were able to spread rapidly throughout Eurasia beginning 50-45 ky ago. Animal remains provide the most direct evidence for MSA and LSA hunting-gathering, and so far, only the Western Cape Province of South Africa has provided large MSA samples that can be directly compared to local LSA samples. The most persuasive comparisons are between MSA samples dating from the Last Interglacial, mainly between about 125 and 100 ky ago, and LSA samples dating from the Present Interglacial, between 12 ky ago and the historic present. This is because the two interglacials were climatically similar, and any observed MSA/LSA faunal contrasts are thus likely to reflect a human behavioral difference as opposed to an environmental one. I have previously identified four contrasts that I summarize briefly here:

1) Present Interglacial LSA coastal sites ( Elands Bay Cave, Die Kelders Cave 1, Blombos Cave, Nelson Bay Cave, Klasies River Mouth Main Site, and others) contain many more bones of fish and of airborne birds than do Last Interglacial MSA sites ( Blombos Cave and the Klasies River Mouth Main Site). Only the LSA sites contain implements that recall ethnohistoric fishing and fowling gear, and artifactual observations thus reinforce the inference that only LSA people fished and fowled routinely. Greater LSA ability to catch fish and birds would probably have promoted larger LSA populations.

2) In Present Interglacial LSA sites ( Nelson Bay Cave, Byneskranskop Cave 1, and others), buffalo and wild pigs outnumber eland, roughly mirroring the dominance of buffalo and wild pigs in the historic environment. In contrast, in Last Interglacial MSA sites (Klasies River Mouth Main Site and Blombos Cave), eland outnumber buffalo and pigs, even though historic observations imply that eland were probably much rarer nearby. Eland continue to dominate in MSA sites that date from the early part of the Last Glaciation (Klasies River Mouth Main Site and Die Kelders Cave 1), which increases the probability that eland dominance reflects MSA behavior and not some undetected environmental factor. Since eland are much less dangerous to hunt than buffalo and wild pigs and since MSA sites lack secure evidence for projectile armatures, an MSA preference for eland may actually reflect MSA reluctance to attack species that were especially likely to injure hunters. LSA people almost certainly had projectiles, arguably including the bow and arrow from 20 ky ago, and they could thus have stalked buffalo and pigs at significantly reduced personal risk. If this inference is correct, and buffalo and pigs were more common than eland on the ground, LSA people would have obtained many more animals overall, even if their stalks were often unsuccessful. Again, a likely result would be larger LSA populations.

3) The tortoises, shellfish, or both in MSA sites (Klasies River Mouth Main Site, Blombos Cave, Die Kelders 1, Ysterfontein, Hoedjiespunt, Sea Harvest, and Boegoeberg 2) tend to be much larger than those in LSA sites (Nelson Bay Cave, Die Kelders Cave 1, Byneskranskop Cave 1, Kasteelberg A and B, Elands Bay Cave, and others) that were occupied under similar climatic conditions. Since tortoises and shellfish can be collected with limited technology and minimal risk, the smaller average tortoise and shellfish size in LSA sites probably reflects more intensive LSA collection that selectively removed the largest individuals first. The most plausible explanation for more intensive collection is that LSA collectors were more numerous, in keeping with their ability to fish, fowl, and hunt more effectively.

4) Finally, the ages of fur seals in LSA sites ( Elands Bay Cave, Kasteelberg A and B, Die Kelders 1, Nelson Bay Cave, and others) suggest that the people focused their coastal visits on the August-to-October interval when nine-ten-month-old seals could be harvested on the shore and when resources inland were probably at their nadir. The ages of MSA fur seals suggest that MSA people remained at the coast more or less throughout the year, even when resources were probably more abundant inland. This difference is the most weakly substantiated of the four listed here, since only the Klasies River Mouth Main site has provided a large enough MSA seal sample for numerically significant comparison to the LSA samples. If fresh MSA samples confirm a likely MSA/LSA contrast in seasonal mobility, a possible explanation is that MSA people could not transport water efficiently. So far, only LSA sites have provided secure evidence for water containers, in the form of ostrich eggshell canteens.

In sum, Western Cape faunal data suggest that LSA people were more effective hunter-gatherers than their MSA predecessors, but a compelling case will require observations from other regions. Fresh observations will also be necessary to determine whether the advance occurred abruptly in the earliest LSA, putatively dated between 50 and 40 ky ago, or perhaps more gradually within the LSA. This is because the Western Cape MSA samples all antedate 60 ky ago, while the LSA samples all postdate 20 ky ago. The gap may never be filled in locally, since it probably reflects adverse regional climatic conditions (extreme aridity) in the middle of the Last Glaciation that sharply reduced the archeological visibility of human populations. The case for an LSA foraging advance thus remains tentative, but pending new evidence, it still seems reasonable to hypothesize that MSA faunal remains, like MSA artifacts imply less than fully modern behavior.
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Modern Human Physiology with Respect to Evolutionary Adaptations That Relate to Diet in the Past

Staffan Lindeberg

Dept of Clinical Sciences, Lund University, Sweden

In medical science, pre-agricultural diets are increasingly acknowledged as templates for healthy diets, partly because contemporary populations with ”ancient” dietary habits have shown very low age-adjusted rates of cardiovascular and other nutrition-related diseases compared to people in developed countries. After the appearance of fully modern humans, the human genome has apparently changed very little, considering the close resemblance in biochemistry between different ethnic groups. For the average westerner, more than 70 % of the energy intake is provided by foods that were practically unavailable during human evolution, i.e. cereals, legumes, dairy products, refined sugar, fats and oils. It is very unlikely that human physiology is fully adapted for these recently introduced staple foods. While such evolutionary reasoning is plausible and straightforward, it is much more difficult to reverse the argument in order to see if human physiology can tell us something about genetic adaptations to food habits in the past.

Humans have the anatomy and physiology of omnivores, and we seem to thrive on vastly different proportions of animal and plant foods, as shown by such extremes as meat-eating Eskimos and high-carb-eating Pacific islanders. Vitamin C cannot be synthesised by humans, but offal from wild game can easily provide the required amounts. Vitamin B 12, another essential nutrient, is absent in plant foods and must be supplied from meat, fish, shellfish or insects, but the required amounts are apparently small. Contemporary vegetarians are at risk of additional deficiencies, such as iron, zinc and calcium, but this is largely due to a high intake of phytate from cereals and legumes, which were obviously not staple foods during human evolution. Wheat and other grains are also low in lysine and threonine, but there is no evidence that a mixture of leafy vegetables, fruits and nuts is inadequate with regard to essential amino acids. The notion that ”animal protein” is unhealthy is largely based on studies of casein, the dominant protein in milk. The recent scientific confusion over desirable proportions of dietary fat, carbohydrate and protein clearly shows the methodological constraints of epidemiology and molecular biology. Hence, the jury is still out on the optimal balance, if there is one, between animal and plant foods. There is no evidence, however, that we are well adapted to a high intake of seeds.

Since iodized salt and dairy products were not available to pre-agricultural humans, only those with high regular access to fish or shellfish would be expected to have reached the currently recommended intake of iodine. There is insufficient data to suggest that humans, by way of natural selection, would have become completely dependent on marine food sources. Therefore, it is highly possible that human requirements for iodine are currently increased by some dietary factors. These theoretically include goitrogens in certain roots, vegetables, beans and seeds. The notion that humans are designed to consume large amounts of long-chain omega-3 fatty acids, irrespective of the intake of omega-6 and omega-9 fatty acids, still awaits solid evidence.

Shifting the focus from general human characteristics to differences between ethnic groups, persistent lactase activity in adulthood is apparently not the only characteristic to have emerged under nutritional selection pressure. Other examples are a relative resistance against diseases of affluence in northern Europeans (the ’non-thrifty genotype’) and a relatively low prevalence of gluten intolerance in South East Europe.

 In conclusion, humans are well adapted to lean meat, fish, insects and highly diverse plant foods without being obviously dependent on any particular proportions of plants versus meat.

The Relationship Between the Human Dentition and Diet During the Paleolithic

Peter Lucas

Department of Anthropology, George Washington University, Washington DC, USA.

he evolution of diet is critical to a proper understanding of human evolution. Partly, this is because the acquisition and processing of food is always a primary focus in ecology, but it also reflects the fact that, compared to other large primates, the diet of modern humans is quite extraordinary. Modern humans (even those still reliant on hunting and gathering) eat discrete cooked meals of meat, grains and root vegetables, while other large primates consume snacks of raw fruits and leaves almost continually. This is a caricature of course – there is overlap – but even considered in detail, the contrast is very sharp. So when, why and where did such a drastic dietary transition take place and how can evidence for it be obtained? The principal morphological evidence must lie in the dentition because it acts directly on solid components of the diet. The key dietary factors influencing a decline in tooth size seen from the Paleolithic onwards, are the size of the mouthful, which must have increased with the advent of the meal, ingested food particle size, which probably decreased with the advent of stone tool use, and changes in the mechanical properties of food - particularly toughness, which probably decreased considerably after the discovery of cooking. Indirect evidence from genetic changes in modern human populations relevant to the oral processing of foods will be added to this analysis to produce a more general model that takes chemical reactions in the mouth into account.

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Plant Evidence for Subsistence in the Middle and Upper Palaeolithic of Europe

Martin Jones

Department of Archaeology, University of Cambridge, United Kingdom

Even among human diets heavily dependant on meat, plant foods will typically play a significant role, but it is not a role about which much is known in Palaeolithic communities. While this is in part due to the nature of the archaeological record in earlier periods, it is also consequent upon the scant use of archaeobotanical recovery techniques on Palaeolithic excavations. The situation is however, very gradually changing, and in this paper I shall review such direct archaeobotanical evidence as exists for the use of plant foods in Middle and Upper Palaeolithic sites in Europe and the Mediterranean region.

I shall preface my review with some general considerations about the significance of plant components to diets in the context of changing ecology and cognitive capacity. I shall follow the review with some further observations about implications for methodology within Palaeolithic fieldwork.

An Energetics Perspective on the Neandertal Record

Kathy MacDonald, Wil Roebroeks, & Alexander Verpoorte

Faculty of Archaeology,Leiden University, The Netherlands

Our aim in this paper is twofold: first to review current knowledge of the Neandertal archaeological record, providing a background to the detailed studies of fauna and particular aspects of technology presented by other participants. In the second part of the paper we (tentatively) explore an energetics perspective on the Neandertal record. Neandertals hunted large mammals in their prime in a wide range of environments, and may have exploited a smaller range of prey species than modern humans. Based on isotopic studies they can be characterized as ‘top carnivores’, eating limited amounts of plant foods or fish. In general, they used simple, low investment tools, including basic projectiles, with little change over time and space, and invested little in spatial structure. The Neandertal record (for diet, technology and use of space) contrasts strongly with the record of Upper Palaeolithic humans. This contrast has generally been interpreted in terms of a cognitive difference between the two species.

A number of recent studies of the fossil record have made progress in studying energy use and requirements. Studies of Neandertal skeletal morphology have come up with a range of estimates for Neandertal energy needs based on different data and calculation methods. These studies concur in suggesting that Neandertal BMR and hence energy requirements were considerably higher than those of modern humans, and also (to a lesser extent) higher than for Upper Palaeolithic humans. In this paper, we will argue that differences in energy requirements and position on the food web have important consequences for behaviour, and show that this perspective can yield interesting predictions that are testable in the archaeological record, focusing on the use of space as an example. This exercise suggests that differences in energy requirements between Neandertals and AMH could provide an alternative explanation for differences in the archaeological record of the Middle and Upper Palaeolithic.

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Epipaleolithic Subsistence Intensification in Southwest Asia: The Faunal Evidence

Natalie D. Munro

Department of Anthropology, Unit 2176, 354 Mansfield Road, University of Connecticut, Storrs, USA

Subsistence intensification—the extraction of increased amounts of energy from a given area at the expense of foraging efficiency—figures prominently in discussions of the Epipaleolithic period (ca. 21,000 – 11,500 cal. BP). Despite their paramount status, intensification trends are rarely subjected to rigorous testing using robust archaeological data sets. This study aims to fill this gap by synthesizing relevant zooarchaeological data from Epipaleolithic sites in Southwest Asia. Intensification is examined not only at the level of the animal community, but also at the taxonomic, and individual carcass level. Multiple lines of faunal evidence including prey age and sex profiles, large and small game abundance, species diversity, and the fragmentation of animal bone support a gradual intensification trend across the Epipaleolithic period that culminates in the transition to agriculture. The pathway toward intensification was not always smooth—variation in Southwest Asian hunting practices indicates numerous stops and starts along the way. The intensification trend reported here corresponds with other long-term processes including increased site use intensity and growing human populations, but crosscuts major climatic events.

Isotopic evidence for European Upper Palaeolithic human diets

M.P. Richards

Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology

Isotope analysis (carbon and nitrogen) of bone collagen is a powerful tool for reconstructing past human and animal diets, and has been applied to a number of European Palaeolithic sites, especially in the middle Palaeolithic (Bocherens et al. 2001, Richards, Pettitt et al. 2000). In contrast to the, albeit limited, middle Palaeolithic isotope analyses of Neanderthals, the isotope values of Upper Palaeolithic humans have a greater diversity in isotopic values. While many Upper Palaeolithic individuals have isotopic values similar to Neanderthals, indicating a diet where the protein was mainly from terrestrial herbivores, others, especially those found close to large freshwater rivers as well as along coasts, have isotopic values indicating a significant consumption of freshwater/marine protein (Richards et al. 2001, Pettitt et al. 2003). In the later Upper Palaeolithic we see even more diversity in the human isotopic values, with many having isotopic values indicating a continued reliance on terrestrial heribivores (Drucker and Henry-Gambier, 2005, Richards, Jacobi et al. 2000) as their main protein source, however others have increased intake of aquatic resources (Richards et al. 2005). This talk will review the applications of isotope analysis to humans from the European Upper Palaeolithic, with a particular emphasis on the evidence for increased dietary diversity, likely indicating a broader spectrum diet, in this period.

Hair δ 13 C values reflect aspects of primate ecology

Margaret J. Schoeninger

University of California at San Diego, La Jolla, CA, USA

Hair samples from different C3-feeding primate species living in ecologically distinct areas of MesoAmerica, South America, Africa and Madagascar have δ 13 C values that vary with canopy cover. Within species variation in chimpanzees also varies with differential canopy cover across Africa. Mantled howler monkeys ( Ateles geoffroyi) from Costa Rica show seasonal differences between rainy and dry seasons at a low level of significance even though they show no significant differences between sexes, sampling year, or year of analysis.

Cebus capucinus (capuchins) and Ateles geoffroyi (spider monkeys) from La Selva, Costa Rica live in an area of tropical wet forest with continuous forest canopy and have identical average δ13C values. Alouatta palliata (howlers) from La Pacifica, Costa Rica, Brachyteles arachnoides (muriquis) from Fazenda Esmeralda, Brazil, and Galago zanzibaricus and Galago garnettii from Gedi, Kenya live in areas of tropical dry forest with broken forest canopy and are similar to each other but significantly different from capuchins and spider monkeys. The difference in average δ 13 C values is of the same magnitude and in the same direction as that in leaves from open canopies compared with closed canopies and is independent of specific primate diet. Lepilemur leucopus from Beza Mahafaly Special Reserve, Madagascar lives in a tropical dry forest with significant numbers of CAM plants. The species average δ 13 C is significantly less negative than the other species and the data vary in association with the percent of CAM plants in the diet of individual animals.

Chimpanzees from Ugalla, a savanna/woodland south of Uvinza in western Tanzania, had average δ 13 C values most similar to the Madagascar lepilemurs from the drought-afflicted forest. Chimpanzees from the Ishasha River (savanna/woodland of eastern Zaire) approximate the New World howlers and muriquis, and the two African galago species from dry, deciduous forests. In sharp contrast, chimpanzees from the Lake Telle region swamp forest of northern Congo approximate New World capuchins and spider monkeys from closed-canopy evergreen forests.

The patterns in the data support the use of similar analyses for reconstructing past ecological variations.

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The Ecological Impact of Projectile Weaponry in Late Pleistocene Human Evolution

John Shea

Department of Anthropology, Stony Brook University, NY, USA

Lethal projectile weaponry is uniquely human and a cultural universal. All historical and ethnographic human cultures use projectile weapons and tactics that allow them to kill or mortally wound a target from a safe distance. The antiquity of projectile weaponry remains unclear. Hypothetical wooden javelins from Schöningen, Germany, are significantly larger than ethnographic projectile weapons. Geochronological and contextual problems complicate the interpretation of hypothetical bone spear points from African Middle Stone Age contexts. Pointed stone artifacts whose ballistically-significant dimensions overlap with those of ethnographic projectile points occur relatively late in the Pleistocene, after 40-50 Ka, in Africa, Europe, and Southwest Asia, although the African evidence is complicated by uncertainties about the functions of various "backed pieces". Although it makes sense to seek the origins of projectile weaponry in the activities in which they are used ethnographically, mainly big-game hunting and warfare, the period 40-50 Ka witnesses ambiguous evidence for increases in either of these activities. Most scholarship on this issue envisions projectile technology emerging first as an aid to subsistence that was later recruited into warfare, but the hypothesis that warfare was a prime mover in the origins and development of projectile technology cannot be rejected.

Brain Size Change as Related to Diet in the Earliest Hominids

J. Josh Snodgrass 1, William R. Leonard 2, and Marcia L. Robertson 2

1 Department of Anthropology, University of Oregon, USA

2 Department of Anthropology, Northwestern University, USA

Bioenergetics, the study of the use and transfer of energy, can provide important insights into the ecology and evolution of early hominids. Despite a relatively large brain with high metabolic demands, contemporary humans and other primates have resting metabolic rates (RMRs) that are similar to those of other mammals. As a result, a comparatively large proportion of their resting energy budget is spent on brain metabolism among humans (~20-25%) and other primates (~8-10%) compared to other mammals (~3-5%). To understand this shift in energy budget, Aiello and Wheeler’s Expensive Tissue Hypothesis (ETH) posits a metabolic trade-off—a reduction in gut size with brain size increase—to explain this phenomenon. Here, we explore the interrelationships between brain size, body size, diet, and body composition to better understand brain evolution in early hominids using comparative data for humans, non-human primates, and other mammals. Although contemporary humans display relatively small guts, primates as a group have gut sizes that are similar to non-primate mammals. In contrast, humans and other primates have significantly less skeletal muscle for their size compared to other mammals. Among living primates, the relative proportion of energy allocated to brain metabolism is positively correlated with dietary quality. Contemporary humans fall at the positive end of this relationship, having both a high quality diet and a large brain size. Thus, high costs associated with the large human brain are supported, in part, by energy-rich diets. These comparative analyses suggest that alterations in body composition and a sufficiently high quality diet were necessary conditions for supporting substantial increases in brain size during hominid evolution. The fossil evidence indicates that the rapid brain expansion with the emergence of Homo erectus at about 1.8 million years ago was likely associated with important changes in both of these parameters (diet and body composition).

Biogeochemical Evidence for the Ecology of Australopiths and Early Homo

Matt Sponheimer

Department of Anthropology, University of Colorado at Boulder, USA

In the past decade researchers have begun to look increasingly to biogeochemical evidence to test hypotheses about early hominin ecology. Here, we present biogeochemical evidence of the diets, habitats, and mobility of early African hominins. Stable carbon isotope analysis has revealed that australopiths and early Homo, both in South and East Africa, consumed considerable quantities of C4 foods. This likely indicates that these hominins were fully integrated into emerging savanna and woodland communities, quite unlike modern chimpanzees, which can be found in such communities today but which consume no measurable abundance of C4 foods. There has also been renewed interest in using elemental ratio data to investigate the trophic position of early hominins, and we show that Sr/Ca and Ba/Ca data, contrary to previous studies, offer no prima facie support for the consumption of animal foods. We caution, however, that researchers have simply not done the necessary background work to meaningfully interpret such elemental ratio data. We also present new carbon, oxygen and strontium isotope data which bear directly on the question of early hominin habitats and mobility.

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Late Pleistocene Subsistence Strategies and Resource Intensification in Africa

Teresa E. Steele

Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Leipzig Germany

Studies in southern Africa, western Europe, and the Mediterranean basin have documented changes in subsistence strategies and technologies during the Late Pleistocene, often relating the changes to differences between the Middle and Upper Paleolithic of Europe and the Middle and Later Stone Age of Africa. However, few explicit comparisons have been made between these geographic regions to create a global view of the evolution of human diet. In this paper, I begin by documenting the state of our knowledge in southern Africa. The lines of evidence that I consider here follow those identified by previous researchers: 1) large game exploitation as indicated by prey mortality profiles and processing intensity, 2) changes in the relative abundance and increasing diversity of species included in the diet, especially the addition of small, fast game such as hares, birds, fish, and small, slow game such as shellfish and tortoises, and 4) changes in shellfish size reflecting intensity of collection. I then expand the discussion to include all of Africa, highlighting similarities and differences in the record and the areas that require further research. Two of the major limitations to this research are the lack of archaeological sites that span the transition from Middle to Later Stone Age (60 to 20 kyr ago) and the lack of data from most regions of Africa. Finally, I consider how the documented subsistence changes relate to demographic changes, the modern human expansion out of Africa, and models of modern human origins.

Trends in the Economic and Demographic Resilience of Paleolithic Forager Societies in the Mediterranean Basin

Mary C. Stiner

Department of Anthropology, Building 30, University of Arizona, Tucson, AZ 85721-0030, USA

Human predator-prey relationships changed dramatically in the Mediterranean Basin from 250,000 to 9,000 years ago. Hoofed animals were the principal source of meat for virtually all foragers in Eurasia prior to the end of the Paleolithic. Large game resources were supplemented with a variety of small animals, the diversity of which underwent the most radical changes with time, leading to considerable expansion in dietary breadth. Small game species are particularly diagnostic of increases in human pressure on the supply of resources and are a major source of evidence for local increases in human population densities after 40-45,000 years ago.

An interesting corollary to the trend in small game use was the increasing use of more productive species in both the large game and small game categories. Rank-order variation in prey biomass indicates a comparatively early date for the onset of this phenomenon in the eastern Mediterranean Basin, a trend that ultimately may have contributed to the collapse of large game hunting adaptations by roughly 10,000 years ago. Productive prey species are those that mature more rapidly and whose populations normally experience high rates of turnover. Although high turnover species generally were more costly to procure, they are better at withstanding heavy hunting by predators and thus may have represented more reliable resources in the short term, when high rank resource supplies were in decline. The Mediterranean Paleolithic cases involve a consistently negative relation between resource rank and prey population resilience for the important prey species.

The trends in human predator-prey dynamics indicate significant restructuring of animal communities of the region. The trends appear to be the product of cyclical feedback mechanisms, provoked in part by a demographic advantage that some human groups experienced inadvertently from dietary diversification. The data suggest step-wise, irreversible shifts in human predatory niche in the Mediterranean Basin, beginning in the east and spreading westward. Evidence of demographic pressure and greater use of resiliant prey populations is followed by technological innovations to exploit these animals more efficiently.

Middle and Lower Paleolithic human reproductive units probably were not robust at the micropopulation scale, due to the rather narrow set of behavioral responses that characterized social groups at the time; localized extinctions at the micropopulation level were likely to have been common as a result. Upper Paleolithic groups were the quintessential colonizers and, in addition, uniquely good at holding on to habitat gained. Upper Paleolithic have shorter histories of existence than those of earlier periods, but they were even more widespread geographically. The demographic robustness of the Upper Paleolithic systems may stem from wholesale strategies for evening-out or sharing risk and volatility in technology. Micropopulations were larger and often denser on landscapes, more connected via cooperative ties, and thus more robust.

Lower and Middle Paleolithic Faunal Exploitation in Europe

Paola Villa

University of Colorado Museum, UCB 265, Boulder, Colorado, USA

This paper is a review of the state of our knowledge and ignorance on Early, Middle and early Upper Pleistocene subsistence behavior in Europe. There are undoubtedly differences in subsistence behavior between early hominids and Upper Paleolithic humans in Europe. Yet recent research has shown that some of the most extreme statements (i.e. about scavenging from previously ravaged carcasses practiced on a regular basis at some Middle and Upper Pleistocene sites) are not supported by the evidence and must be rejected. The following issues will be discussed in this paper:

(1) evidence of subsistence behavior at some of the earliest sites, that is sites dated to the end of the Lower Pleistocene or just after the Matuyama-Bruhnes boundary; (2) the nature of the European data base and problems associated with the scarcity of taphonomic analyses; (3) evidence concerning the exploitation of very large mammals (elephants, rhinoceroses) and very small vertebrates and invertebrates (leporids, birds, fish and shellfish) in the Lower and Middle Paleolithic; (4) hunting weapons of the Middle Paleolithic.

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