Forthcoming Current Anthropology Wenner-Gren Symposium Curren t Supplementary Issues (in order of appearance)

VOLUME 54 SUPPLEMENT 8 DECEMBER 2013

Crisis, Value, and Hope: Rethinking the Economy. Susana Narotzky and Anthropolog y Current Niko Besnier, eds. e Anthropology of Christianity: Unity, Diversity, New Directions. Joel Robbins and Naomi Haynes, eds. Anthropology Politics of the Urban Poor. Veena Das and Shalini Randeria, eds.

Previously Published Supplementary Issues THE WENNER-GREN SYMPOSIUM SERIES December 2013 Working Memory: Beyond Language and Symbolism. omas Wynn and ALTERNATIVE PATHWAYS TO COMPLEXITY: Frederick L. Coolidge, eds. EVOLUTIONARY TRAJECTORIES IN THE MIDDLE Engaged Anthropology: Diversity and Dilemmas. Setha M. Low and Sally PALEOLITHIC AND MIDDLE STONE AGE Engle Merry, eds. GUEST EDITORS: STEVEN L. KUHN AND ERELLA HOVERS Corporate Lives: New Perspectives on the Social Life of the Corporate Form. Damani Partridge, Marina Welker, and Rebecca Hardin, eds. Alternative Pathways to Complexity V e Origins of Agriculture: New Data, New Ideas. T. Douglas Price and 54 olum e Mediterranean and Red Sea Paleoclimate Ofer Bar-Yosef, eds. Neanderthal Demographic Estimates Agreements and Misunderstandings among Three Scientific Fields e Biological Anthropology of Living Human Populations: World Histories, National Styles, and International Networks. Susan Lindee Hominin Evolution in the Middle-Late Pleistocene and Ricardo Ventura Santos, eds. Variability in the Middle Stone Age of Eastern Africa

Supplement Roots of the Middle Paleolithic in Eurasia Human Biology and the Origins of Homo. Susan Antón and Leslie C. Aiello, eds. Middle Stone Age Hunting Strategies and Diet Breadth Trends versus Conservatism in the Predatory Niche Potentiality and Humanness: Revisiting the Anthropological Object in Technological Trends in the Middle Stone Age of South Africa Contemporary Biomedicine. Klaus Hoeyer and Karen-Sue Taussig, eds. Change and Stasis in the Iberian Middle Paleolithic 8 Lessons from the Levantine Middle Paleolithic Record Paleolithic Cultures in China Current Anthropology is sponsored by e Mechanisms behind MP and MSA Cultural Trajectories Wenner-Gren Foundation for Anthropological Population Size and the Paleolithic Archaeological Record

Research, a foundation endowed for scientific, Page s educational, and charitable purposes. e Measuring the Complexity of Lithic Technology Foundation, however, is not to be understood as

endorsing, by virtue of its financial support, any of S173–S406 the statements made, or views expressed, herein.

Sponso r e d b y the W enner-Gren Founda tion f o r Anth r opologic a l Rese a r c h

THE UNIVERSIT Y O F CHICAGO PRESS Wenner-Gren Symposium Series Editor: Leslie Aiello Wenner-Gren Symposium Series Managing Editors: Laurie Obbink and Daniel Salas Current Anthropology Editor: Mark Aldenderfer Current Anthropology Managing Editor: Lisa McKamy Book Reviews Editor: Holley Moyes Corresponding Editors: Claudia Briones (IIDyPCa-Universidad Nacional de Rı´o Negro, Argentina; [email protected]), Michalis Kontopodis (Humboldt Universita¨t zu Berlin, Germany; [email protected]), Jose´Luis Lanata (Universidad Nacional de Rı´o Negro San Carlos de Bariloche, Argentina; [email protected]), David Palmer (Hong Kong University, China; [email protected]), Anne de Sales (Centre National de la Recherche Scientifique, France; [email protected]), Zhang Yinong (Shanghai University, China; [email protected])

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Alternative Pathways to Complexity: Evolutionary Trajectories in the Middle Paleolithic and Middle Stone Age

Leslie C. Aiello Alternative Pathways to Complexity: Evolutionary Trajectories in the Middle Paleolithic and Middle Stone Age: Wenner-Gren Symposium Supplement 8 S173 Steven L. Kuhn and Erella Hovers Alternative Pathways to Complexity: Evolutionary Trajectories in the Middle Paleolithic and Middle Stone Age: An Introduction to Supplement 8 S176 Eelco J. Rohling, Katharine M. Grant, Andrew P. Roberts, and Juan-Cruz Larrasoan˜a Paleoclimate Variability in the Mediterranean and Red Sea Regions during the Last 500,000 Years: Implications for Hominin Migrations S183 Jean-Pierre Bocquet-Appel and Anna Degioanni Neanderthal Demographic Estimates S202 Carles Lalueza-Fox Agreements and Misunderstandings among Three Scientific Fields: Paleogenomics, Archaeology, and Human Paleontology S214 Osbjorn M. Pearson Hominin Evolution in the Middle-Late Pleistocene: Fossils, Adaptive Scenarios, and Alternatives S221 Christian A. Tryon and J. Tyler Faith Variability in the Middle Stone Age of Eastern Africa S234 Steven L. Kuhn Roots of the Middle Paleolithic in Eurasia S255 Jamie L. Clark and Andrew W. Kandel The Evolutionary Implications of Variation in Human Hunting Strategies and Diet Breadth during the Middle Stone Age of Southern Africa S269

http://www.journals.uchicago.edu/CA MaryC.Stiner An Unshakable Middle Paleolithic? Trends versus Conservatism in the Predatory Niche and Their Social Ramifications S288 Sarah Wurz Technological Trends in the Middle Stone Age of South Africa between MIS 7 and MIS 3 S305 Ignacio de la Torre, Jorge Martı´nez-Moreno, and Rafael Mora Change and Stasis in the Iberian Middle Paleolithic: Considerations on the Significance of Mousterian Technological Variability S320 Erella Hovers and Anna Belfer-Cohen On Variability and Complexity: Lessons from the Levantine Middle Paleolithic Record S337 Xing Gao Paleolithic Cultures in China: Uniqueness and Divergence S358 Francesco d’Errico and William E. Banks Identifying Mechanisms behind Middle Paleolithic and Middle Stone Age Cultural Trajectories S371 Mark Collard, Briggs Buchanan, and Michael J. O’Brien Population Size as an Explanation for Patterns in the Paleolithic Archaeological Record: More Caution Is Needed S388 Charles Perreault, P. Jeffrey Brantingham, Steven L. Kuhn, Sarah Wurz, and Xing Gao Measuring the Complexity of Lithic Technology S397 Current Anthropology Volume 54, Supplement 8, December 2013 S173

Alternative Pathways to Complexity: Evolutionary Trajectories in the Middle Paleolithic and Middle Stone Age Wenner-Gren Symposium Supplement 8

by Leslie C. Aiello

Figure 1. Seated: Leslie Aiello, Xing Gao, Francesco d’Errico, Christian Tryon, Erella Hovers. Standing: Ignacio de la Torre, Jamie Clark, Sarah Wurz, Anna Degioanni, Eelco Rohling, Mary Stiner, Osbjorn Pearson, Mark Collard, Charles Perreault, Steve Kuhn, Ariel Malinsky-Buller. A color version of this photo appears in the online edition of Current Anthropology.

The Wenner-Gren Foundation for Anthropological Research supplement of Current Anthropology (see http://www has a long tradition of organizing symposia that deal with the .wennergren.org/history/conferences-seminars-symposia “big” questions in contemporary anthropology. Alternative /wenner-gren-symposia for a complete list of symposia and Pathways to Complexity: Evolutionary Trajectories in the Middle the history of the symposium program). The Alternative Path- Paleolithic and Middle Stone Age is the 145th in the sympo- ways symposium was organized by Steven L. Kuhn (University sium series and the eighth to be published as an open-access of Arizona, U.S.A.) and Erella Hovers (Hebrew University, Israel) and was held at Ha¨ringe Slott, Stockholm, Sweden, Leslie C. Aiello is President of the Wenner-Gren Foundation for June 1–8, 2012 (fig. 1). Anthropological Research (470 Park Avenue South, 8th Floor North, Ha¨ringe Slott was the country home of Axel Wenner-Gren New York, New York 10016, U.S.A.). from the late 1930s until his death in 1961, and it was par-

᭧ 2013 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved. 0011-3204/2013/54S8-0001$10.00. DOI: 10.1086/673284 S174 Current Anthropology Volume 54, Supplement 8, December 2013

Figure 2. Ha¨ringe Slott, the summer home of Axel Wenner-Gren, located in Va¨sterhaninge, Sweden, approximately 25 km south of Stockholm. A color version of this photo appears in the online edition of Current Anthropology. ticularly enjoyable to spend an academically stimulating week are summarized in the introduction to Human Biology and in a beautiful environment relevant to the history of the Foun- the Origins of Homo (Aiello 2012), a recent Current Anthro- dation (fig. 2). It is also noteworthy that this symposium pology supplementary issue on human origins (Anto´n and followed on from a workshop cosponsored by the Wenner- Aiello 2012). The majority of these deal with the earlier phases Gren Foundation for Anthropological Research and the Swed- of the human evolutionary record, reflecting the focus on ish Wenner-Gren Foundations (the Wenner-Gren Center australopith research in eastern and southern Africa in the Foundation for Scientific Research, the Axel Wenner-Gren 1960s and 1970s. A notable exception is After the Australo- Foundation for International Exchange of Scientists, and the pithecines: Stratigraphy, Ecology and Culture Change in the Foundation Wenner-Grenska Samfundet). The meeting was Middle Pleistocene (Butzer and Isaac 1975). In more recent titled Reality and Myth: A Symposium on Axel Wenner-Gren years the Foundation has supported meetings on broader hu- and was held at the Wenner-Gren Center in Stockholm, Swe- man origins topics, for example, Ancestors: The Hard Evidence den (May 30–31, 2012). It provided the first opportunity in (Delson 1985), and on broader topics, Tools, Language and recent years for the independent U.S. and Swedish founda- Cognition in Human Evolution (Gibson and Ingold 1993), tions to meet and explore the fascinating history of their Roots of Human Sociality: Culture, Cognition and Interaction mutual founder. Information on this workshop and the life (Enfield and Levinson 2006), and Working Memory: Beyond of Axel Wenner-Gren can be found at http://wennergren.org Language and Symbolism (Wynn and Coolidge 2010). /history/-story-and-people-wenner-gren/people-wenner Alternative Pathways combines past interests in data-rich -gren/axel-wenner-gren. discussions of human origins with more theoretical discus- Alternative Pathways is the most recent symposium in the sions of the significance of paleoanthropology for understand- Foundation’s long history of interest in and support for re- ing the course of human evolution. As discussed in the in- search on human origins. One of the first large meetings troduction to this issue (Kuhn and Hovers 2013), Middle organized by the Foundation (in 1956 in collaboration with Paleolithic Neanderthals and Middle Stone Age humans pro- the city of Dusseldorf, Germany, and H. R. von Koenigswald, vide a unique opportunity “to study the evolution of different Wilhelm Gieseler, and Horst Sieloff), marked the centenary kinds of intelligence and different modes of behavioral com- of the discovery of Neanderthal Man (Neanderthal 1; fig. 3; plexity in sister taxa, using multiple lines of archeological and and see http://www.wennergren.org/history/conferences biological evidence.” One purpose of the meeting was to in- -seminars-symposia/wenner-gren-symposia/wenner-gren form our understanding of why Neanderthals went to ex- -symposia1950-1970). tinction and modern humans were able to spread throughout Other Wenner-Gren Symposia dealing with human origins the world. However, going beyond this, the intention was to Aiello Alternative Pathways to Complexity: Evolutionary Trajectories in the Middle Paleolithic and Middle Stone Age S175

emerged from the symposium, and these will take the research forward. Additionally, the papers provide an assessment of state-of-the art knowledge in the Middle Paleolithic of Eurasia as well as the Middle Stone Age in Africa, along with relevant paleoclimatological, genetic, demographic, and biological per- spectives. This collection is destined to be an important re- source for the future. The Wenner-Gren Foundation is always looking for big questions and innovative new ideas in all areas of anthro- pology for future Foundation-sponsored and Foundation- organized symposia and eventual CA publication. We en- courage anthropologists to contact us with their proposals for future meetings. Information about the Wenner-Gren Foun- dation, the symposium program, application procedures and deadlines, and what constitutes a good symposium topic can be found on the Foundation’s website (http://wennergren.org /programs/international-symposia).

References Cited

Aiello, Leslie C. 2012. Human biology and the origins of Homo: Wenner-Gren symposium supplement 6. Current Anthropology 53(suppl. 6):S267–S268. Anto´n, Susan C., and Leslie C. Aiello, eds. 2012. Human biology and the origins of “Homo.” Current Anthropology, vol. 53, suppl. 6. Butzer, Karl W., and Glynn Isaac, eds. 1975. After the Australopithecines: stra- tigraphy, ecology and culture change in the Middle Pleistocene. World An- thropology Series. The Hague: Mouton. Figure 3. From the left: Gunter Behm-Blanke, G. H. R. von Delson, Eric, ed. 1985. Ancestors: the hard evidence. New York: Liss. Koenigswald, and Abbe H. Breuil examine the Ehringsdorf fossil Enfield, Nicholas J., and Stephen C. Levinson, eds. 2006. Roots of human remains. sociality: culture, cognition and interaction. Oxford: Berg. Gibson, Kathleen, and Tim Ingold, eds. 1993. Tools, language and cognition in human evolution. Cambridge: Cambridge University Press. explore the alternative elaborate and flexible cultural adaptive Kuhn, Steven L., and Erella Hovers. 2013. Alternative pathways to complexity: systems associated with two forms of cognitively sophisticated evolutionary trajectories in the Middle Paleolithic and Middle Stone Age: an introduction to supplement 8. Current Anthropology 54(suppl. 8):S176– hominins. S182. As with most successful meetings, more questions were Wynn, Thomas, and Frederick L. Coolidge, eds. 2010. Working memory: beyond raised than answers provided. However, collaborative projects language and symbolism. Current Anthropology, vol. 51, suppl. 1. S176 Current Anthropology Volume 54, Supplement 8, December 2013

Alternative Pathways to Complexity: Evolutionary Trajectories in the Middle Paleolithic and Middle Stone Age An Introduction to Supplement 8

by Steven L. Kuhn and Erella Hovers

The 145th symposium of the Wenner-Gren Foundation took place June 1–8, 2012, in Ha¨ringe Slott near Stockholm, Sweden. The primary goal of the symposium was to reframe discussions of behavioral evolution among Neanderthals and early modern humans. We hoped to replace conventions of a single scale of evolutionary progress (in which the primary benchmark is “modern human behavior”) with a more Darwinian framework that could allow for independent evolutionary trajectories in different areas. The 15 participants included archaeologists researching material culture and subsistence in Eurasia, Africa, and China; physical anthropologists; a demographer; a geneticist; modelers of cultural evolution; and a climatologist. Participants were asked to draw on evidence in their areas of expertise, focusing on evolutionary trends in both modal tendencies and levels of variation/diversity within various regions during the interval in which the Middle Stone Age and Middle Paleolithic developed, spread, and eventually disappeared. It was agreed that there is compelling evidence for very different trajectories of cultural evolution in different parts of the world but that we are not yet in a position to fully evaluate and understand the outcomes of the parallel cultural evolutionary pathways among modern Homo sapiens in Africa and Neanderthals in Europe. Answering questions this large in scope requires synthesis on a large geographic scale comparable to studies by climate scientists and biogeographers. Conventional approaches to collecting, reporting, and analyzing archaeological and skeletal data do not lend themselves to rigorous tests of alternative evolutionary models. At the same time, the intellectual tools needed to research these questions are well developed, and answers are within reach.

In his 1989 book Wonderful Life (Gould 1989), the biologist of these was the subject of the workshop from which the S. J. Gould proposed a singular thought experiment, the idea papers of this volume derive. of “replaying the tape of life.” Gould’s aim was to raise ques- Genetic evidence (Endicott, Ho, and Stringer 2010; Lalueza- tions about the role of contingency in the evolution of life Fox 2013) show that Neanderthals and anatomically modern on earth, to consider whether the same initial conditions Homo sapiens (AMHs) last shared a common ancestor roughly could have led to radically different forms of plants and an- 400–500 kya. Although they may have remained genetically imals because of slight differences in external conditions. compatible (Sankararaman et al. 2012), by the time of the most Scholars disagree about the likely outcomes of the exercise. recent of the human dispersals out of Africa and into western Gould (1989:51) believed that repeatedly playing the tape of Eurasia, the two metapopulations had been partially if not life forward would produce very different outcomes, whereas wholly isolated from one another for at least 300,000 yr. Be- Dennett (1995) argues the opposite. Regardless of the specific tween the time the two lineages separated and the time they views, such a line of inquiry perforce remains a thought ex- came into contact again in marine isotope stage (MIS) 4, a periment. And yet for more limited arenas there are natural certain amount of encephalization and a great deal of cultural situations that partially replicate Gould’s mental exercise. One evolution had occurred in both of them. The long period of geographic vicariance rendered Neanderthals and African H. Steven L. Kuhn is Professor at the School of Anthropology of the sapiens both behaviorally and anatomically differently derived University of Arizona (Building 30, Tucson, Arizona 85721-0030, descendants of their last common ancestor. By the time they U.S.A. [[email protected]]). Erella Hovers is Professor at the Institute of Archaeology of the Hebrew University of Jerusalem met again in Eurasia, both Neanderthals and African H. sap- (Mount Scopus, Jerusalem 91905, Israel). This paper was submitted iens had developed culturally aided adaptations that were 3 VII 13, accepted 14 VIII 13, and electronically published 20 XII more complex than those of any other organism in the history 13. of the planet, in essence creating new regimes of evolutionary

᭧ 2013 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved. 0011-3204/2013/54S8-0002$10.00. DOI: 10.1086/673501 Kuhn and Hovers Alternative Pathways to Complexity S177 change. Thus, the two taxa can be seen as the outcomes havioral records of the two taxa represent alternative elaborate of parallel evolutionary “experiments” resulting in large- and flexible cultural adaptive systems associated with two brained, behaviorally complex, yet quite distinct hominins. forms of cognitively sophisticated hominins. The parallel biological and cultural evolution in Europe The notion of biological and cultural evolution in Africa and Africa certainly differs from Gould’s original example and Eurasia as parallel experiments has significance beyond of the Burgess Shale fauna in that it involved evolutionary explaining the record of hominin evolution in both areas. processes unfolding in different habitats and leading to Comparison of the evolutionary trajectories taken by Nean- diversification, whereas Gould was referring to loss of di- derthals and AMHs affords a rare prospect in the history of versity. Still, the underlying principle that one can get very life on earth to study the evolution of different kinds of in- differently evolved outcomes from the same starting point telligence and different modes of behavioral complexity in remains the same. sister taxa, using multiple lines of archaeological and biolog- For most of the history of archaeology and paleoan- ical evidence. There is growing interest across the biological thropology, studies of Neanderthals and early AMHs have and social sciences in topics such as the evolution of culture typically adopted a very different perspective on the re- and niche construction. Humans are typically held up as niche lationship between the two taxa than the one implied by constructors, and modern humans are regarded as the ulti- Gould’s metaphor. Comparisons of material culture evi- mate expression of a culturally adapted organism with no dence from Eurasia and Africa have focused mainly on the equal in the animal kingdom. Because they are so different evolutionary status of the Middle Paleolithic (MP) and from other living organisms, they are regarded as an extreme Middle Stone Age (MSA) in relation to the presumed com- case. Treating the records of Neanderthals and AMHs as two plexity of later time periods. The behavioral evolution of independent pathways of cognitive and behavioral evolution both taxa has been evaluated using a single scale, defined opens the prospect of a much richer and more nuanced un- in terms of a shifting set of traits referred to as “modern derstanding of the potential for different kinds of culturally human behavior.” Studies tend to focus on the manifes- aided niche construction in cognitively sophisticated organ- tations of particular behavioral traits (ornaments, bone isms. It could shift the discussion from the human cultural tools, small-game hunting) among Neanderthals or AMHs niche to diverse sorts of cultural niches created and occupied as testament to what members of the two taxa could or by contemporaneous hominins. could not do. This evidence is not treated symmetrically. In organizing the Wenner-Gren symposium, we were in- For the most part, similarities are equated with equality terested in creating a forum for empirically based, theoretically between the two populations, but differences tend to be informed discussions of different evolutionary trajectories of inferred as deficiencies on the part of Neanderthals. Be- Neanderthals, African H. sapiens, and other hominins inde- cause the field remains focused on the question of what pendent of the eventual fate of the population. The goal was happened to the Neanderthals and why H. sapiens came not to formulate a consensus about which population first to be the only hominin on the planet, differences or sim- reached some threshold of “behavioral modernity” (a concept ilarities in turn are used to justify or undermine hypotheses that is epistemologically and practically problematic; see, e.g., about the behavioral, cultural, or cognitive superiority of Belfer-Cohen and Hovers 2010; Henshilwood and Marean AMHs. 2003; Hovers 2009a, 2009b; Hovers and Belfer-Cohen 2006; The differing behavioral tendencies of MP and MSA hom- Kuhn and Hovers 2006; Shea 2011). From the perspective inins may or may not help us explain the historical circum- that we adopt here, that particular question is anecdotal. In- stances of the Neanderthal’s disappearance and the rapid dis- stead, the specific aims of the symposium were to discuss and persal and persistence of modern humans across the globe. evaluate the evidence for independent histories of behavioral However, we believe that attempts to establish the relative evolution in western Eurasia and southern Africa. We were positions of the MP and MSA in a single progressive, devel- not so ambitious or so naive as to expect a final, compre- opmental scheme culminating in “behavioral modernity” are hensive account of the outcome and implications of the Eur- misguided (Hovers 2009a, 2009b; Hovers and Belfer-Cohen asian and African evolutionary “experiments.” Our aim was 2006; Kuhn and Hovers 2006). They represent a holdover considerably more modest. A previous symposium and edited from the anagenetic mind-sets of early twentieth-century cul- volume (Hovers and Kuhn 2006) explored the evidence for tural evolutionists, for whom all of human cultural variation behavioral change within the MP and MSA. The more recent could be fit into a single set of developmental stages that workshop turned to the question of outcomes—what hap- marked the gradual elevation of the human race from savagery pened in the two macroregions—and how variables such as to civilization or from acephalous bands to complexly or- environmental change and demographic dynamics shaped ganized states. Crucially, attempts to organize the MP and (pre)historical trajectories. Equally, we hoped to reframe the MSA into a linear framework of change toward “modernity” discussion and to reach a clearer understanding of the kinds distract us from a unique and more valuable opportunity of information still needed to answer the big questions we created by the long separation of two human lineages involved were posing. in authoring these records. The differences between the be- With these questions in mind, the 145th symposium of S178 Current Anthropology Volume 54, Supplement 8, December 2013 the Wenner-Gren Foundation was convened June 1–8, have developed everywhere out of local variants of the Acheu- 2012, in Ha¨ringe Slott near Stockholm, Sweden (befittingly, lean. a former residence of the foundation’s financial benefactor Stronger contrasts emerge when comparing the archae- and namesake, Axel Wenner-Gren). The location and tim- ological records of the later MSA and MP. Sites from across ing enabled participants to enjoy (or suffer) 20 hours of southern Africa show similar patterns of cultural succession sunlight per day (on the days when the sun made an ap- during MIS 4, with MSA 1 and 2 followed by distinctive pearance, that is). Invitees were asked to draw on evidence but short-lived Still Bay, Howiesons Poort, and “post–How- in their areas of expertise, focusing on evolutionary trends iesons Poort” technocomplexes (but see Brown et al. 2012, in both modal tendencies and levels of variation/diversity who suggest that the punctuated nature of technological within various regions. As dictated by the nature of the practices associated with these cultural phenomena is an questions asked, the 15 participants brought to the dis- artifact of sampling). The Still Bay and Howiesons Poort cussion a diversity of training and expertise. The group have received a great deal of attention in recent years because included paleoanthropologists who study material culture of evidence for “complex” behaviors such as bone-tool mak- and subsistence in Eurasia, Africa, and China; physical an- ing, personal ornaments, heat treatment of raw materials, thropologists; a demographer; a geneticist; modelers of cul- and abstract symbolic renderings (Wurz 2013). The orderly tural evolution; and a climatologist. The papers and dis- sequencing of assemblage types across southern Africa also cussions focused on the time period between 400 and 40 fits many researchers’ expectations for cumulative, progres- ka (MIS 9–3), the interval in which the MSA and MP sive cultural evolution or “cultural ratcheting” (Tennie, Call, developed, spread, and eventually disappeared.1 That many and Tomasello 2009; Tomasello 1999) similar to what is of the papers presented in the formal session were co- observed in the Eurasian Upper Paleolithic (UP). authored by two or more researchers is perhaps a reflection Well-studied areas of southwest Europe show rather di- of the scope and scale of the questions addressed. vergent evolutionary patterns during the late MP. As de la The first part of the workshop was devoted to establishing Torre and colleagues (2013) demonstrate, there are no strong the empirical evidence for evolutionary trajectories in Eurasia directional trends within the northern Spanish MP record and southern Africa. Though differing in the details, the his- through the Upper Pleistocene, whereas the southwestern tories of the African MSA and European MP seem to be French Mousterian shows a distinct sequence of industries similar, with a gradual coalescence of behavioral character- over time (Delagnes, Jaubert, and Meignen 2007; Delagnes istics over the second half of the Middle Pleistocene (Kuhn and Meignen 2006). The presence of very divergent regional 2013; Tryon and Faith 2013). Common features such as Le- trajectories in neighboring regions such as northern Iberia vallois technology are likely to represent convergent evolution and southern France exemplifies the geographically parti- rather than diffusion: in fact, there may be multiple Levallois tioned nature of the European MP record. These observations origins within both Africa and Eurasia (Villa 2001; White and also highlight one apparent difference in the Eurasian and Ashton 2003:605). Contrary to some characterizations, both African records, namely the apparently more progressive pat- the MSA and the MP show a notable level of technological tern of culture change in the MSA of southern Africa. This variability, particularly in methods for producing flake blanks. raises the question of whether the Still Bay and Howiesons Interestingly, the sorts of regional variability long noted in Poort represent the first evidence of cumulative, directional MP sequences from Eurasia are not as strongly expressed in human cultural evolution, as some have argued, or whether the East African record studied by Tryon and Faith (2013). they simply reflect more rapid diffusion of novel ideas across Although substantial differences in sample size potentially southern African landscapes. Conversely, the more localized temper any conclusions drawn from these observations, such and divergent patterns in southern Europe could be evidence geographic variability as exists among East African MSA and for different cultural capacities of the hominins for acquisition European assemblages may correspond with broadly defined and diffusion of cultural information, but they could also habitat types (e.g., d’Errico and Banks 2013). The grain of simply be a consequence of more fragmented habitats and variation in behavioral evidence in both Africa and Eurasia less thoroughly interconnected populations. may reflect the structure of populations on the landscape, The contribution by Gao (2013) brings into the discussion which in turn is strongly influenced by the scale of environ- a very different record. The workshop focused on the MSA mental variation and presence of ecological barriers (Hovers and western Eurasian MP simply because the preponderance and Belfer-Cohen 2013; Tryon and Faith 2013). Cultural di- of the evidence comes from those macroregions. A substantial versity in western Eurasia may also have deeper chronological body of archaeological and fossil evidence is now available roots, as there are strong regional differences in technology from China, even if it is not well known to Western scholars. as early as 400 kya. In contrast, the African MSA seems to In contrast to Africa and western Eurasia, where lithic tech- nology provides robust evidence of culture change and di- 1. The student moderator for the meeting, Mr. Ariel Malinsky-Buller, versification during the Paleolithic, the Chinese lithic record is currently writing his PhD thesis on the transition from the late Lower is remarkably stable from the Early Pleistocene through the to the early Middle Paleolithic in Eurasia. late Pleistocene or even into the early Holocene. It makes little Kuhn and Hovers Alternative Pathways to Complexity S179 sense to talk about a distinct “MP” phenomenon, and over Evidence of subsistence intensification in the form either of most of that vast country even early UP stone artifacts differ expanding diet breadth or intensified resource processing has remarkably little from much older Lower Paleolithic ones (see been widely associated with changes in population-resource Bar-Yosef and Wang 2012). At the same time, elements of balances. When the effects of changing climates are filtered material culture such as ornaments and bone and antler ar- out, the MP archaeofaunal record of Eurasia (summarized by tifacts made their way into the behavioral repertoire of Chi- Stiner 2013) provides very little evidence for sustained sub- nese hominins during the Upper Pleistocene. The Middle and sistence intensification. The hominins producing MP indus- Upper Pleistocene archaeology of China strongly suggests that tries seem to have focused their meat-oriented foraging efforts this region may represent yet another independent trajectory on high-ranked, large-body-sized ungulate prey. There is reg- in the evolution of behavioral complexity during the Upper ular use of some specific kinds of small game and sporadic Pleistocene. Recent analyses of ancient and modern DNA evidence for expansion of the meat diet to include more costly indicate that northeast Asia was also home to a distinct pop- and lower-ranked items such as birds and small mammals ulation of hominins, further demonstrating its independence even in the early MP. However, these cases seem isolated and from “the West.” We suspect that there are other parallel so far do not resolve into evidence for sustained intensification situations in macroregions such as south Asia and possibly of subsistence. The more rarefied archaeofaunal record from North Africa not covered in this workshop. southern Africa (synthesized by Clark and Kandel 2013) It was clear from presentations and discussions during the shows a clear response to climate, especially evident in faunas symposium that demography and its effects on processes of dating to MIS 4. Still, while shellfish exploitation appears quite cultural transmission have become important to many ac- early in southern Africa (as it does in Iberia and other parts counts of MP/MSA technological variation and foraging be- of the Mediterranean), there are no long-term trends indic- haviors. The majority of participants subscribed to the view ative of continuous pressure on high-ranked resources unless that Neanderthals were as a rule spread very thin on the the gradual increase in suid exploitation is considered as such. landscape. Demography can influence the flow of cultural Thus, the South African faunal evidence at least provides little information just as it does the flow of genetic information, support for the notion that demographic conditions were very and many interesting hypotheses about variation and inno- different there than in Eurasia. vation in cultural practices as well as trajectories of biological Vegetable foods may have played an important, even dom- change can be derived from this single premise. Interestingly, inant role in the diets of hominins in many times and places, the physical anthropologist who participated in the discussion but macroscopic plant remains seldom preserve over Paleo- (Pearson 2013) also concludes that demographic effects such lithic timescales. While occasional discoveries of preserved as multiple bottlenecks may explain many of the anatomical seeds from archaeological sites as well as recent findings of characteristic of native Eurasian hominins (i.e., Neanderthals) phytoliths and starch grains adhering to teeth or evidence for better than models of adaptation. plant residues on artifacts (e.g., Barton et al. 1999; Hardy and As compelling as demographic explanations may be, they Moncel 2011; Henry, Brooks, and Piperno 2010; Lev, Kislev, are still in their infancy. To date, models of demographic and Bar-Yosef 2005) open a new window on the diets of effects on Paleolithic cultural dynamics have been verified Neanderthals and other hominins, the unfortunate fact re- (checked for internal consistency) but have not been validated mains that various sources of data are incommensurate. An- (i.e., checked against real-world evidence; Collard, Buchanan, imal bones are common and quantifiable but reveal only part and O’Brien 2013). The papers by Boucquet-Appel and De- of the diet. Stable isotope data provide a better proportional gioanni (2013) and by Lalueza-Fox (2013) point out some of estimate of the importance of plant and animal foods, but the difficulties in testing these models. Estimates of demo- only as regards their contributions to protein intake. Mean- graphic potentials based on life-history characteristics are no- while, emerging but still sparse data on plant foods in the toriously sensitive to component variables and often produce Paleolithic tell us about diversity but not quantity; that is, we conflicting predictions, whereas estimates of effective popu- know what people might have eaten but not how much. In- lation size from genetic data are not easily translated into tegrating these disparate sources into a more coherent picture census population size. Attempts to reconstruct census pop- of dietary variation is a challenge yet to be addressed by the ulation levels based on archaeological evidence (number of field. The work of nutritionists and ecologists, who have iden- dated sites, site area, etc.; see, e.g., Mellars and French 2011) tified a number of patterns of the use of fish, meat, and plant are uncertain (Dogandzˇic´ and McPherron 2013). Thus, while resources among recent foraging populations, can be helpful there is both empirical and theoretical reason to believe that in contextualizing some of the archaeological data into a more Neanderthals lived at comparatively low population densities, nuanced view of dietary variations in various regions (e.g., we have a very coarse understanding of actual MP population Cordain et al. 2000; Hockett 2012; Hockett and Haws 2005; sizes or of the relative sizes of contemporaneous or later H. Kelly 1995; see review in Speth 2010). On the other hand, sapiens populations. these observations are themselves contingent on the particular Other sources of information may provide proxy evidence historical, climatic, and demographic circumstances of the last for changes in demography if not actual population sizes. few hundred years. Another line of evidence for dietary in- S180 Current Anthropology Volume 54, Supplement 8, December 2013 tensification—and indirectly, population-resource imbal- retical models that had been discussed to formulate testable ances—could come from technology. Maximally efficient use hypotheses about rates of innovation, change, and cu- of plant foods such as seeds and nuts often requires intensive mulative cultural evolution. One working group developed processing to achieve maximum nutrient yields. Moreover, a framework for evaluating complexity of technical pro- the technological accoutrements of this sort of food process- cesses as a means of assessing cumulative cultural evolu- ing—grinding or pounding stones—are durable and large, tion. The paper by Perrault et al. (2013) contains an initial unlike the botanical remains themselves. Reports of grind- application of this framework to Paleolithic lithic tech- stones from MSA sites in southern Africa may hint at an nological evidence, a “proof of concept” for this analytical unexplored dimension of subsistence intensification in the and methodological approach. A second group explored remote past, but until these artifacts are systematically ana- strategies for comparing assemblages of material culture lyzed in Africa as well as in Eurasia, they remain little more across regions with very different histories as a means of than tantalizing anecdotes. evaluating both cultural phylogenies and adaptive varia- Overall, the similarities in subsistence regimes between late tion. Pleistocene H. sapiens in southern Africa and late Pleistocene In many respects, the rise of population-based arguments Neanderthals in western Eurasia far outweigh the differences. in paleoanthropology today is similar to what occurred in This observation in turn highlights another interesting dif- population genetics in the 1960s or in ecology during the late ference in the outcomes of independent trajectories of cultural 1990s, when neutral theory was first introduced into those evolution in the two regions. Recent research has produced disciplines by Kimura (1968, 1983) and Hubbell (2001), re- indisputable evidence that both MSA and MP hominins reg- spectively. Recognition of the importance of replicator dy- ularly produced composite, multipart tools, some of which namics led to an increasing emphasis on stochastic, neutral were weapons (Rots 2013; Tryon and Faith 2013; Villa and processes such as drift or dispersal at the expense of selection Lenoir 2008; Villa et al. 2009; Wilkins et al. 2012; Wurz 2013). or adaptation. In fact, both stochastic and directional/selective However, the frequency, variety, and elaboration of projectile processes influence most genetic and cultural systems. The elements seems to be higher in the later MSA than the late big challenge for archaeology and human paleontology alike MP even though the Pleistocene environments of temperate is to sort out what is neutral from what is selectively or Eurasia should have favored greater dependence on large game adaptively salient in a particular case. Papers by Collard, than the subtropical and tropical habitats in which MSA pop- Buchanan, and O’Brien (2013) and Pearson (2013) show how ulations operated. Among recent foragers, greater dependence this could be approached for both anatomy and material cul- on large game has been shown to correlate with more in- ture. vestment in hunting technology (see Collard, Buchanan, and Obviously, climate is a key element in any evolutionary O’Brien 2013). The divergence was not caused by the absence scenario. A third theme that figured prominently both in this of key innovations or some cognitive difference, as the same workshop and in the current literature concerns the effects basic procedures and techniques had long existed in both of climatic events on the evolutionary histories of different areas. One potential explanation is that MP diets in fact in- hominin species and different regions. Attempts to match volved less use of large game than is normally thought. An trends in anatomical and archaeological data to global climate alternate explanation is that MP hominins used means such curves are basically exercises in pattern recognition, underlain as group-wide social cooperation to improve hunting effec- by the explicit assumption that association equals causation, tiveness. Finally, unstable demographic conditions in Eurasia rather than concerted attempts at explanation. As Rohling et could have prevented MP hominins from reliably passing on al. (2013) emphasize, matching scales of observation and anal- complex technological procedures. These kinds of contrasts ysis is a crucial prerequisite to integrating data from diverse point to possible fundamental divergences in the cultural fields. When paleoanthropology turns to climate sciences to niches of African MSA and Eurasian MP hominins. At the define the environmental envelope of MSA/MP behaviors in same time they reveal anomalies that must be resolved for various regions, spatial and temporal scales are often incom- the field to move forward. patible. Climate variability in the Upper Pleistocene is often In the end, understanding how the sizes and structures documented on a millennial, centennial, or (rarely) decadal of Neanderthal and AMH populations may have influenced scale of resolution, whereas dating of archaeological sites does cultural evolutionary processes will require more than just not normally achieve such accuracy. At the same time, sites more precise estimates of population sizes. The resulting are by nature of restricted spatial scales whereas major climatic models must also be made empirically testable using ar- shifts occur globally yet manifest themselves differently at chaeological data. Accomplishing this will require more smaller geographic scales. Hominins in Eurasia and southern precise definitions and replicable measures of key param- Africa were exposed to the same global climatic events, but eters such as complexity and diversity of cultural behaviors they experienced them in very different ways. Features of the analogous but not identical to some measures in genetics landscape (as well as technological repertoires and anatomy) and ecology. During the last days of the meeting, partic- play a major role in determining how these global events are ipants worked toward using the empirical data and theo- manifest at the scale of sites or research areas. An important Kuhn and Hovers Alternative Pathways to Complexity S181 focus of discussion during the latter part of the workshop using material culture data and attempts to identify phylo- was developing strategies for resolving such geographic and genetic relationships between MP and MSA cultural entities. temporal incongruities, ranging from development of local This fulfills one final aim of the workshop, which was to open records to focusing on periods of high and low variability up new avenues for collaboration among participants and rather than specific sets of conditions (Rohling et al. 2013). their colleagues in a range of disciplines. The notion of a “Cohesive Adaptive System,” or CAS, as outlined by d’Errico and Banks here and in other papers (Banks et al. 2006), is one approach to analyzing culture and environmental variation on similar spatial and temporal References Cited scales. Banks, William E., Francesco d’Errico, Harold L. Dibble, Leonard Krishtalka, Differences between southern Africa and western Eurasia Dixie West, Deborah I. Olszewski, A. Townsend Peterson, et al. 2006. Eco- cultural niche modeling: a new tool for reconstructing the geography and with respect to the magnitude of climate change over time ecology of past human populations. Paleoanthropology 2006:68–83. and spatial grain of environmental variation will make it Barton, R. N. E., Andrew Currant, Yolanda Fernandez-Jalvo, Clive Finlayson, difficult though not impossible to fully exploit the natural Paul Goldberg, Richard Macphail, Paul Pettit, and Chris Stringer. 1999. Gibraltar Neanderthals and results of recent excavations in Gorham’s, Van- experiments that inspired this workshop. However, other guard, and Ibex caves. Antiquity 73:13–24. areas may provide unique controls over these variables. Bar-Yosef, Ofer, and Youping Wang. 2012. Paleolithic archaeology in China. The Levant offers a particularly interesting and useful case. Annual Review of Anthropology 41:319–335. Belfer-Cohen, Anna, and Erella Hovers. 2010. Modernity, enhanced working Because of the strong habitat zonation and limited mag- memory, and the Middle to Upper Paleolithic record in the Levant. Current nitude of temperature and rainfall variation when com- Anthropology 51(suppl. 1):S167–S175. pared with more extreme latitudes, the region provided a Bocquet-Appel, Jean-Pierre, and Anna Degioanni. 2013. Neanderthal demo- graphic estimates. Current Anthropology 54(suppl. 8):S202–S213. relatively stable environmental setting over the course of Brown, Kyle S., Curtis W. Marean, Zenobia Jacobs, Benjamin J. Schoville, the Pleistocene compared with Europe. It also has a Simen Oestmo, Erich C. Fisher, Jocelyn Bernatchez, Panageorgis Karkanas, “checkered” history in terms of hominin dispersals, with and Thalassa Matthews. 2012. An early and enduring advanced technology originating 71,000 years ago in South Africa. Nature 491:590–593. multiple taxa, including both early AMHs and Neander- Clark, Jamie L., and Andrew W. Kandel. 2013. The evolutionary implications thals, coming and going. Thanks to many highly detailed of variation in human hunting strategies and diet breadth during the Middle studies of lithic material culture and of nonlithic related Stone Age of southern Africa. Current Anthropology 54(suppl. 8):S269–S287. Collard, Mark, Briggs Buchanan, and Michael J. O’Brien. 2013. Population behaviors combined with high resolution terrestrial cli- size as an explanation for patterns in the Paleolithic archaeological record: matic records, research on the Levantine MP may serve as more caution is needed. Current Anthropology 54(suppl. 8):S388–S396. a first approximation for constructing scale-appropriate Cordain, Lauren, Janette B. Miller, S. Boyd Eaton, Neil Mann, Susanne H. Holt, and John D. Speth. 2000. Plant-animal subsistence ratios and mac- research programs that are suitable to address the specific ronutrient energy estimations in worldwide hunter-gatherer diets. American questions raised in the meeting and a basis on which to Journal of Clinical Nutrition 71:682–692. incorporate new methods of analysis. The paper by Hovers Delagnes, Anne, Jacques Jaubert, and Liliane Meignen. 2007. Les technocom- plexes du Pale´olithique moyen en Europe occidentale dans leur cadre dia- and Belfer-Cohen (2013) explores some of the apparent chronique et ge´ographique. In Les ne´andertaliens: biologie et cultures.Ber- consequences of this unique environment for the behavior nard Vandermeersh and Bruno Maureille, eds. Pp. 213–229. Documents of successive hominin populations. South Africa and Pre´historiques 23. Paris: E´ ditions du Comite´ des Travaux Historiques et Scientifiques. southwest Europe (northern Iberia, southern France) were Delagnes, Anne, and Liliane Meignen. 2006. Diversity of lithic production also nominated as areas of roughly comparable size and systems during the Middle Palaeolithic in France: are there any chrono- data density, although they have very different environ- logical trends? In Transitions before the transition: evolution and stability in the Middle Paleolithic and Middle Stone Age. Erella Hovers and Steven L. mental histories. Kuhn, eds. Pp. 85–107. New York: Springer. One conclusion from the meeting was that we are not yet de la Torre, Ignacio, Jorge Martı´nez-Moreno, and Rafael Mora. 2013. Change in a position to fully evaluate and understand the outcomes and stasis in the Iberian Middle Paleolithic: considerations on the signifi- cance of Mousterian technological variability. Current Anthropology of the parallel evolutionary trajectories represented by AMHs 54(suppl. 8):S320–S336. and the MSA in Africa and Neanderthals and the MP in Dennett, Daniel C. 1995. Darwin’s dangerous idea: evolution and the meanings Europe. The meeting has underlined the fact that whereas the of life. New York: Simon & Schuster. d’Errico, Francesco, and William E. Banks. 2013. Identifying mechanisms basic notion is sound and some of the available evidence behind Middle Paleolithic and Middle Stone Age cultural trajectories. Cur- compelling, conventional approaches to collecting, reporting, rent Anthropology 54(suppl. 8):S371–S387. and analyzing data do not lend themselves to rigorous tests Dogandzˇic´, T., and S. P. McPherron 2013. Demography and the demise of Neandertals: a comment on “Tenfold population increase in Western Europe of alternative evolutionary models. Answering questions this at the Neandertal-to-modern human transition.” Journal of Human Evo- large in scope requires synthesis on a vast geographic scale lution 64:311–313. similar to efforts undertaken by climate scientists and bio- Endicott, Phillip, Simon Ho, and Chris Stringer. 2010. Using genetic evidence to evaluate four palaeoanthropological hypotheses for the timing of Ne- geographers. At the same time, the participants agreed that anderthal and modern human origins. Journal of Human Evolution 59:87– the intellectual tools needed to research these questions are 95. well within reach. Various members of the group have begun Gao, Xing. 2013. Paleolithic cultures in China: uniqueness and divergence. Current Anthropology 54(suppl. 8):S358–S370. collaborative efforts in this direction, including the design Gould, Stephen J. 1989. Wonderful life: the Burgess Shale and the nature of and implementation of pilot studies of cumulative evolution history. New York: Norton. S182 Current Anthropology Volume 54, Supplement 8, December 2013

Hardy, Bruce L., and Marie-Helene Moncel. 2011. Neanderthal use of fish, Western Europe at the Neandertal-modern human transition. Science 333: mammals, birds, starchy plants and wood 125–250,000 years ago. PLoS 623–627. ONE 6:e23768. Pearson, Osbjorn M. 2013. Hominin evolution in the Middle-Late Pleistocene: Henry, Amanda G., Alison S. Brooks, and Dolores R. Piperno. 2010. Micro- fossils, adaptive scenarios, and alternatives. Current Anthropology 54(suppl. fossils in calculus demonstrate consumption of plants and cooked foods in 8):S221–S233. Neanderthal diets (Shanidar III, Iraq; Spy I and II, Belgium). Proceedings Perreault, Charles, P. Jeffrey Brantingham, Steven L. Kuhn, Sarah Wurz, and of the National Academy of Sciences of the USA 108:486–491. Xing Gao. 2013. Measuring the complexity of lithic technology. Current Henshilwood, Christopher S., and Curtis W. Marean. 2003. The origin of Anthropology 54(suppl. 8):S397–S406. modern human behavior: critique of the models and their test implications. Rohling, Eelco J., Katharine M. Grant, Andrew P. Roberts, and Juan-Cruz Current Anthropology 44:627–651. Larrasoan˜a. 2013. Paleoclimate variability in the Mediterranean and Red Hockett, Bryan. 2012. The consequences of Middle Paleolithic diets on preg- Sea regions during the last 500,000 years: implications for hominin migra- nant Neanderthal women. Quaternary International 264:78–82. tions. Current Anthropology 54(suppl. 8):S183–S201. Hockett, Bryan, and Jonathan A. Haws. 2005. Nutritional ecology and the Rots, Veerle. 2013. Insights into early Middle Palaeolithic tool use and hafting human demography of Neandertal extinction. Quaternary International 137: in Western Europe: the functional analysis of level IIa of the early Middle 21–34. Palaeolithic site of Biache-Saint-Vaast (France). Journal of Archaeological Hovers, Erella. 2009a. The lithic assemblages of Qafzeh Cave. New York: Oxford Science 40:497–506. University Press. Sankararaman, Sriram, Nick Patterson, Heng Li, Svante Pa¨a¨bo, and David ———. 2009b. The Middle-to-Upper Paleolithic transition: what news? In Reich. 2012. The date of interbreeding between Neandertals and modern Sourcebook of Paleolithic transitions: methods, theories, and interpretations. humans. PLoS Genetics 8(10):e1002947, doi:10.1371/journal.pgen.1002947. Marta Camps and Parth Chauhan, eds. Pp. 455–462. New York: Springer. Shea, John J. 2011. Homo sapiens is as Homo sapiens was: behavioral variability Hovers, Erella, and Anna Belfer-Cohen. 2006. “Now you see it, now you versus “behavioral modernity” in Paleolithic archaeology. Current Anthro- don’t”: modern human behavior in the Middle Paleolithic. In Transitions pology 52:1–35. before the transition: evolution and stability in the Middle Paleolithic and Speth, John D. 2010. The paleoanthropology and archaeology of big-game hunt- Middle Stone Age. Erella Hovers and Steven L. Kuhn, eds. Pp. 295–304. ing: protein, fat, or politics? New York: Springer. Stiner, Mary C. 2013. An unshakable Middle Paleolithic? trends versus con- New York: Springer. servatism in the predatory niche and their social ramifications. Current ———. 2013. On variability and complexity: lessons from the Levantine Anthropology 54(suppl. 8):S288–S304. Middle Paleolithic record. Current Anthropology 54(suppl. 8):S337–S357. Tennie, Claudio, Josep Call, and Michael Tomasello. 2009. Ratcheting up the Hovers, Erella, and Steven L. Kuhn, eds. 2006. Transitions before the transition: ratchet: on the evolution of cumulative culture. Philosophical Transactions evolution and stability in the Middle Paleolithic and Middle Stone Age. New of the Royal Society B 364(1528):2405–2415. York: Springer. Tomasello, Michael. 1999. The cultural origins of human cognition. Cambridge, Hubbell, Stephen P. 2001. The unified neutral theory of biodiversity and bio- MA: Harvard University Press. geography. Princeton, NJ: Princeton University Press. Tryon, Christian A., and J. Tyler Faith. 2013. Variability in the Middle Stone Kelly, Robert L. 1995. The foraging spectrum: diversity in hunter-gatherer life- Age of eastern Africa. Current Anthropology 54(suppl. 8):S234–S254. ways. Washington, DC: Smithsonian Institution. Villa, Paola. 2001. Early Italy and the colonization of Western Europe. Qua- Kimura, Motoo. 1968. Evolutionary rate at the molecular level. Nature ternary International 75:113–130. 217(5129):624–626. Villa, Paola, Paolo Boscato, Filomena Ranaldo, and Annamaria Ronchitelli. ———. 1983. The neutral theory of molecular evolution. Cambridge: Cam- 2009. Stone tools for the hunt: points with impact scars from a Middle bridge University Press. Paleolithic site in southern Italy. Journal of Archaeological Science 36(3):850– Kuhn, Steven L. 2013. Roots of the Middle Paleolithic in Eurasia. Current 859. Anthropology 54(suppl. 8):S255–S268. Villa, Paola, and Michel Lenoir. 2008. Hunting and hunting weapons of the Kuhn, Steven L., and Erella Hovers. 2006. General introduction. In Transitions Lower and Middle Paleolithic of Europe. In The evolution of hominin diets. before the transition: evolution and stability in the Middle Paleolithic and Jean-Jacques Hublin and Michael Richards, eds. Pp. 59–87. Dordrecht: Middle Stone Age. Erella Hovers and Steven L. Kuhn, eds. Pp. 1–11. New Springer. York: Springer. White, Mark, and Nick Ashton. 2003. Lower Paleolithic core technology and Lalueza-Fox, Carles. 2013. Agreements and misunderstandings among three the origins of the Levallois method in north-western Europe. Current An- scientific fields: paleogenomics, archaeology, and human paleontology. Cur- thropology 44:598–608. rent Anthropology 54(suppl. 8):S214–S220. Wilkins, Jayne, Benjamin J. Schoville, Kyle S. Brown, and Michael Chazan. Lev, Ephraim, Mordechai E. Kislev, and Ofer Bar-Yosef. 2005. Mousterian 2012. Evidence for early hafted hunting technology. Science 338:942–946. vegetal food in Kebara Cave, Mt. Carmel. Journal of Archaeological Science Wurz, Sarah. 2013. Technological trends in the Middle Stone Age of South 32:475–484. Africa between MIS 7 and MIS 3. Current Anthropology 54(suppl. 8):S305– Mellars, Paul, and Jennifer C. French. 2011. Tenfold population increase in S319. Current Anthropology Volume 54, Supplement 8, December 2013 S183

Paleoclimate Variability in the Mediterranean and Red Sea Regions during the Last 500,000 Years Implications for Hominin Migrations

by Eelco J. Rohling, Katharine M. Grant, Andrew P. Roberts, and Juan-Cruz Larrasoan˜a

The Mediterranean–Red Sea region has been critical to dispersal of hominids and other species between Africa and the rest of the world, and climate and sea level are thought to be key controls on migration pathways. Assessing climate variations, we highlight increased millennial-scale variability at 480–460, 440–400, 380–360, 340–320, 260– 220, 200–160, 140–120, and 80–40 thousand years ago (ka), which likely caused intermittent habitat fragmentation. We also find that passageways across the Sahara Desert and the northern out-of-Africa route (from Egypt into the Levant) were intermittently open during pluvials associated with orbital insolation maxima. No such relationship is apparent for the southern out-of-Africa route (across the Red Sea). Instead, we present a novel interpretation of combined sea-level and regional climate control on potential migrations via the southern route, with “windows of opportunity” at 458–448, 345–340, 272–265, 145–140, and 70–65 ka. The 145–140 ka window seems relevant for ka, and the 70–65 ka window agrees with estimates of 65 ϩ5/Ϫ8 ka for 16 ע early colonization of Arabia at 127 the final out-of-Africa migration by the anatomically modern human founder group of all non-Africans. Once they reached Eurasian Mediterranean margins, populations benefited from a rich diversity of terrain and microclimates, with persistent favorable conditions in lowlands and potential to occupy higher elevations during milder periods.

Introduction contractions and expansions of the Sahara desert; and (4) the Indian Ocean monsoon, which causes seasonal wind reversals Њ Њ The Mediterranean–Red Sea region occupies a zone that is over the southern Red Sea region up to latitudes of 20 –25 N. influenced by four major climate systems. From northwest to No significant rainfall is associated with the Indian Ocean southeast, these are (1) the temperate westerlies that affect monsoon over the Red Sea and Arabian Peninsula today, but Europe, the western Mediterranean, and the northern sector the monsoon’s summer rainfall domain may have shifted onto of the eastern Mediterranean; (2) the dry subtropical con- the southeast margin of the Arabian Peninsula (i.e., Yemen ditions that dominate the southern and eastern sectors of the and Oman) during insolation-driven monsoon maxima such Mediterranean basin as well as the entire Red Sea region; (3) as those of the Early-Middle Holocene (Conroy and Overpeck the African monsoon, which affects Mediterranean conditions 2011). through inflow of major rivers (Nile, and in the past other Climatic gradients over the Mediterranean and Red Sea North African drainage systems) and that has caused past region are well illustrated by the updated Ko¨ppen-Geiger cli- mate classification (Kottek et al. 2006; fig. 1). Important mea- sures of climate variability through time are obtained from a Eelco J. Rohling is Professor, Katharine M. Grant is Postdoctoral variety of methods, such as pollen data, lake levels, stable Researcher, and Andrew P. Roberts is Professor at the Research isotopes from a variety of sedimentary archives, faunal School of Earth Sciences of the Australian National University changes, and others. These data especially supply information (Canberra, Australian Capital Territory 0200, Australia about changes in regional temperature and precipitation re- [[email protected]; [email protected]; andrew gimes. In addition, the Red Sea has in the past decade become [email protected]]). Juan-Cruz Larrasoan˜a is Staff Scientist at the Instituto Geolo´gico y Minero de Espan˜a of Unidad de Zaragoza a key region for reconstruction of continuous records of sea- (Zaragoza 50006, Spain [[email protected]]). This paper was level fluctuations (Grant et al. 2012; Rohling et al. 1998a, submitted 3 VII 13, accepted 4 IX 13, and electronically published 2008a, 2008b, 2009b, 2010; Siddall et al. 2003, 2004). Finally, 18 XII 13. the presence of vast deserts to the south of the Mediterranean

᭧ 2013 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved. 0011-3204/2013/54S8-0003$10.00. DOI: 10.1086/673882 S184 Current Anthropology Volume 54, Supplement 8, December 2013

Figure 1. Summary map of the main climate zones in the study region using the Ko¨ppen-Geiger climate classification. A p equatorial; B p arid; C p warm temperate; D p snow; s p summer dry; f p fully humid; S p steppe; W p desert; w p winter dry; m p monsoonal; a p hot summer; b p warm summer; k p cold arid; h p hot arid. Modified after Kottek et al. (2006). A color version of this figure is available in the online edition of Current Anthropology. and around the Red Sea gives rise to large windblown dust evaporation in the entire Mediterranean increases toward the fluxes into these basins, and reconstructions of those fluxes east, with an average of 1.45 m yϪ1 (Malanotte-Rizzoli and through time—notably using marine sediment cores—also Bergamasco 1991) to 1.57 m yϪ1 (Be´thoux and Gentili 1994). reveal important changes in regional climate conditions. The classical Mediterranean climate is a result of the re- In this paper, we present an overview of the modern climate gion’s location on the transition between temperate westerlies of the region and of the main changes that have been recon- that dominate over central and northern Europe and the sub- structed from paleoclimate proxy records for the last 500,000 tropical high-pressure belt over North Africa (fig. 2; Boucher yr. This overview provides a context for consideration of the 1975; Lolis, Bartzokas, and Katsoulis 2002). In summer, sub- role of climate variability in anthropological and archaeolog- tropical high-pressure conditions (and drought) extend from ical developments at the interface of Africa and Europe. the southeast in a northwestward direction over most of the Mediterranean. Polar-front depressions may still reach the Modern Climatic Setting western Mediterranean, but they only exceptionally penetrate the eastern Mediterranean (Rohling and Hilgen 1991). During Mediterranean winter, the subtropical conditions shift southward, and the We summarize modern Mediterranean climate conditions fol- northern sector of the Mediterranean becomes influenced by lowing the recent review of Rohling et al. (2009a). The clas- the temperate westerlies with associated Atlantic depressions sical Mediterranean climate is characterized by warm and dry that track eastward over Europe. These depression influences summers and mild and wet winters. Mean annual precipi- extend from the Mediterranean southeastward across the Le- tation along the Mediterranean ranges from less than 0.12 m vant and into the northernmost sector of the Red Sea (e.g., in North Africa, to over 2.00 m in portions of southwest Arz et al. 2003a; Bar-Matthews et al. 2003; Goodfriend 1991; Turkey and in the eastern Adriatic Sea along the slopes of the Matthews, Ayalon, and Bar-Matthews 2000; McGarry et al. Dinaric Alps (Naval Oceanography Command 1987). Total 2004; Trommer et al. 2010). Rohling et al. Mediterranean and Red Sea Paleoclimate S185

Figure 2. Atmospheric circulation pattern during Northern Hemisphere summer. The main winds are indicated by arrows. ITCZ p intertropical convergence zone; H p areas of high sea-level pressure; L p areas of low sea-level pressure. After an adaptation in Rohling et al. (2009), which compiled information from Rossignol-Strick (1985) and Reichart (1997).

Polar and continental air masses over Europe are channeled faces in the northern sectors of the basin. Thus, winter cy- into the Mediterranean basin through gaps in the mountain- clones are linked to North Atlantic systems, given that they ous topography of the northern Mediterranean margin. Dur- represent either (occasional) direct entries of Atlantic synoptic ing winter and spring, intense cold and dry air flows through systems into the Mediterranean basin or secondary lows the lower Rhone Valley to reach the Gulf of Lions (the “mis- formed when Atlantic systems interact with the Alps and lead tral”), and similar flows extend over the Adriatic and Aegean to cyclogenesis within the basin (Trigo, Davies, and Bigg Seas (the “Bora” and “Vardar”), where they cause strong evap- 2000). oration and sea surface cooling (e.g., Casford et al. 2003; Over the Mediterranean Sea, cyclogenesis is most frequent Leaman and Schott 1991; Maheras et al. 1999; Poulos, Drak- over the Gulf of Genoa and the Ligurian Sea, but the Aegean opoulos, and Collins 1997; Saaroni et al. 1996; and references Sea is also a major center for winter cyclogenesis (Boucher therein). The northerly air flows into the western and eastern 1975; Cantu 1977; Rumney 1968; Trewartha 1966; Trigo, Da- Mediterranean are determined by interaction between an in- vies, and Bigg 1999). Most Genoan depressions track over tense low over the central or eastern Mediterranean and Italy, thereby affecting the Adriatic region, and thence in a northeastward extension of the Azores High (over Iberia, generally eastward direction toward the Aegean Sea and/or France, and southern Britain) or westward ridging of the northern Levantine seas (Lolis, Bartzokas, and Katsoulis 2002; Siberian High toward northwestern Europe and southern Rumney 1968; Trewartha 1966; Trigo, Davies, and Bigg 1999). Scandinavia (Lolis, Bartzokas, and Katsoulis 2002; Maheras These depressions, along with those that develop over other et al. 1999). Persistent winter low-pressure conditions over centers of cyclogenesis, cause the winter precipitation that is the region result from high Mediterranean sea surface tem- characteristic of modern Mediterranean climate. Geological peratures (Lolis, Bartzokas, and Katsoulis 2002). The most pronounced basin-wide cold winter events com- archives indicate that Mediterranean depressions have con- plement cold conditions over Europe and develop in asso- trolled Mediterranean climate in the Levant as an enduring ciation with positive sea-level pressure anomalies to the west feature over glacial-interglacial timescales (Bar-Matthews or northwest of the British Isles and particularly low pressure et al. 2003; Goodfriend 1991; Matthews, Ayalon, and Bar- over the Mediterranean. An important aspect of winter var- Matthews 2000; McGarry et al. 2004; Rohling 2013; and ref- iability concerns cyclogenesis (formation of new depressions), erences therein). which governs precipitation in the northeastern and south- Summer rainfall is low around the Mediterranean region, central sectors of the Mediterranean. Some Atlantic depres- especially in eastern and southeastern sectors. Some cyclo- sions may enter the (western) basin, but most cyclones ob- genesis occurs around Cyprus and the Middle East in summer, served in the Mediterranean form over the basin itself but adiabatic descent in the upper troposphere—related to (Rumney 1968; Trigo, Davies, and Bigg 1999), when cold and the Asian summer monsoon—precludes deep convection over relatively dry northerly air flows extend over warm sea sur- the region and so causes the prevalence of dry summer con- S186 Current Anthropology Volume 54, Supplement 8, December 2013 ditions (Rodwell and Hoskins 1996; Trigo, Davies, and Bigg of only 2 mm yϪ1 when distributed over the entire Red Sea 1999). surface area. Thus, river inflow and precipitation are negligible The African monsoon does not reach directly into the Med- throughout the Red Sea region, especially when contrasted iterranean basin, and there is no evidence that it ever did with the high rates of evaporation, which seasonally reach 2 during the Quaternary. It does, however, have (or more ap- myϪ1 or more (Maillard and Soliman 1986; Morcos 1970; propriately used to have) a “remote” influence on the basin Pedgley 1974; Privett 1959; see also Fenton et al. 2000; Siddall through Nile River discharge. Before the anthropogenic con- et al. 2004). Evaporation over the Red Sea is seasonally af- trol of the Nile, its average discharge was 8.4 # 1010 m3 yrϪ1 fected by atmospheric circulation changes that are related to (4.5 # 1010 m3 yrϪ1 in low-flood years to 15.0 # 1010 m3 the Indian Ocean monsoon. The most notable feature is a yrϪ1 in high-flood years), which from the mid-1960s has dwin- seasonal wind reversal over the southern Red Sea, the general dled to nearly nothing (Be´thoux 1984; Nof 1979; Rohling and nature of which is discussed below. For more details and Bryden 1992; Said 1981; Wahby and Bishara 1981). During modeling, see Jiang et al. (2009). the predamming instrumental era, a strong (threefold) inter- During winter (October–May), reduction of sensible heat annual variability has been noted between high and low dis- radiation occurs over the cold landmass of central Asia that charge years, mainly due to variability in the monsoon-fed is enhanced by snow-induced high albedo (reflection of in- contribution of the Blue Nile and Atbara rivers (see data solation) over the Tibetan Plateau and Himalayas. This results summary in Rohling et al. 2009a). in a quasi-stable high-pressure system that extends from Mon- The Nile River comprises two different systems: the White golia to central Europe, Turkey, and Arabia. The cold, de- Nile, which drains the equatorial uplands of Uganda in a scending air leads to a radial outflow of cold dry air toward regular, permanent manner; and the Blue Nile and Atbara, low-pressure areas over the relatively warm Indian Ocean. which drain highly seasonal (summer) African monsoon pre- The intertropical convergence zone (ITCZ) is displaced south- cipitation from the Ethiopian highlands. Summarizing the ward in winter, to 20ЊS over East Africa, and the winter north- predamming Nile hydrology after Adamson et al. (1980) and east monsoon blows across the Arabian Sea and the Gulf of Williams et al. (2000), it appears that up to 30% of the annual Aden toward central Africa (Morcos 1970). A general south- discharge of the Nile originated from the White Nile and a easterly wind circulation results between a continental de- minimum of 70% of the annual discharge from the Blue Nile/ pression over central Africa and a continental anticyclone that Atbara. The winter flow was dominated (83%) by the steady extends from Asia to Arabia. Channeling by the rift geometry White Nile contribution, and the Blue Nile/Atbara compo- of the Red Sea coast then causes strong (6.7–9.3 m sϪ1) winds nent provided 90% of the summer flow (with a peak over to blow from the south or south-southeast over the southern August–October). The White Nile discharge has a much half of the Red Sea up to about 20ЊN (e.g., Morcos 1970; smaller ratio of change between its annual peak and lowest Patzert 1974). Throughout October–December, there is a con- monthly value, with a maximum between late September and vergence between these south-southeasterlies and the north- January. northwesterly winds that prevail year-round over the northern It should be noted that the Nile has not always been the Red Sea. This convergence occupies a north-south zone across only route for drainage of African monsoon precipitation into the Red Sea at around 20ЊN (Jiang et al. 2009; Pedgley 1974), the Mediterranean. During past monsoon maxima (related to which varies in size and is characterized by low-pressure calms orbitally induced insolation maxima), the extent of the Sahara (Morcos 1970). Jiang et al. (2009) demonstrated that the po- desert was much reduced, and there was northward routing sition of the convergence zone may be determined by gaps of drainage from the central Saharan watershed into the Med- in the topography, notably the Tokar Gap some 50 km inland iterranean along the wider North African margin (see below from the Tokar Delta on the Sudan coast, which acts as an for details). outlet of surface wind from the Red Sea basin in winter (and as an inlet in summer; see below). During summer (June–September), the monsoon system Red Sea is reversed. The spring melt reduces albedo over central Asia, The Red Sea basin is entirely situated in an arid zone with and insolation increases, thereby causing the landmass to very low humidity. Coastal stations record annual rainfall warm. This warms the air above, causing it to rise, creating figures of less than 20 mm in the north and 50–100 mm in a quasi-stable low-pressure system over northern India (!995 the south (Pedgley 1974). Riverine flow into the basin is neg- mb; Morcos 1970), which extends over Pakistan to the Persian ligible because of the basin’s small watershed (Maillard and Gulf. The updraft causes humid, relatively cool, maritime air Soliman 1986; Morcos 1970; Siddall et al. 2004; fig. 3). One to flow in from the area of higher pressure over the now of the larger systems that drains into the Red Sea is the Baraka relatively cool Indian Ocean. Latent heat release from con- (Tokar) wadi in Sudan, which today is active 40–70 d yrϪ1 densation/precipitation against the flanks of the Himalayas (mainly during autumn). Wadi Baraka discharges 200– fuels further deepening of the continental low-pressure cell 970#106 m3 water at 18.5ЊN into the Red Sea (Trommer et and consequently more inflow of maritime air, which rein- al. 2011; Whiteman 1971), which is equivalent to a maximum forces the system. The ITCZ reaches its most northerly po- Rohling et al. Mediterranean and Red Sea Paleoclimate S187

Figure 3. Topography of the Red Sea basin. The dashed line delineates the Red Sea watershed. Numbers refer to key sediment core locations (not used here). After Siddall et al. (2004). A color version of this figure is available in the online edition of Current Anthropology. sition (20ЊN) in July and becomes identified with the front Red Sea longitudinal axis. . . . This . . . can also drive dust of the southwest monsoon. This passes north of Aden, and storms.” the southwest monsoon flows in a clockwise direction over In the northernmost Red Sea, records of past climate East Africa, the Gulf of Aden, and the Arabian Sea toward change have detected an influence of southeast Mediterranean the main monsoon low of northern India (Morcos 1970). At climate influences that extend across the Middle East (Arz et this time, a general northwesterly circulation is set up on the al. 2003a; Legge, Mutterlose, and Arz 2006; Trommer et al. western side of the summer Asiatic low-pressure cell. This 2010). Contemporary climatology suggests that this link func- Ϫ causes relatively weak (2.4–4.4 m s 1) north-northwesterly or tions through winter frontal rainfall associated with Cyprus northwesterly winds to dominate over the entire length of the lows (El-Fandy 1946; Morcos 1970). Red Sea (e.g., Morcos 1970; Patzert 1974). Jiang et al. (2009) analyzed regional patterns superimposed on the general along-axis wind systems, which are more zonal Past Climate Variability (west to east or east to west) in nature and that are related Glacial-Interglacial Changes to gaps in the topography along the basin, such as the Tokar Gap. There are eastward-blowing wind jets in summer Throughout the last three million years, climate variability (mainly through the Tokar Gap) and westward-blowing wind over the entire study region has been dominated by the effects jets in winter from the Saudi Arabian margin (mainly over of global ice-age cycles, which were particularly prominent the northern Red Sea). Wind speeds in these surface jets can during the last 500,000 yr (e.g., Lisiecki and Raymo 2005), reach 10–15 m sϪ1, and Jiang et al. (2009) provide clear evi- with variability that is forced by cyclic changes in the earth- dence of atmospheric dust entrainment over the Red Sea. sun orbital configuration. The astronomical forcing of climate Jiang et al. (2009:5) further note that “other strong zonal takes place because of three main processes, namely: changes winds . . . blow from the Egyptian coast eastward across the in the eccentricity of the earth’s orbit around the sun, with S188 Current Anthropology Volume 54, Supplement 8, December 2013 climate effects in approximately 100- and 400-kyr cycles; In the Red Sea, glacial-interglacial differences in sea surface changes in the tilt of the earth’s rotational axis, with effects temperature were of the order of 3Њ–4ЊC (Arz et al. 2003b; on climate in 41-kyr cycles; and precession of the equinoxes, Trommer et al. 2011), with a bias toward summer. These with climate effects in cycles of 19- and 23-kyr durations (e.g., estimates for the Red Sea are in agreement with estimates for Berger 1977; Hays, Imbrie, and Shackleton 1976; Imbrie and the western Arabian Sea (see compilation of Rohling et al. Imbrie 1980, 1986; Milankovitch 1941). 2012). Glaciations were strongly focused on the North American Terrestrial temperature contrasts between glacials and in- and northern Eurasian locations of continental ice sheets, but terglacials are typically around 1.3 or 1.5 times larger than the effects on climate were global. This had global implications sea surface temperature contrasts (Braconnot et al. 2007; for sea level, with fluctuations between interglacial highstands Laıˆne´ et al. 2009). This agrees with estimates by Kuhlemann up to ϩ10 m (Kopp et al. 2009; Muhs et al. 2011; Rohling et al. (2008) of terrestrial temperature reductions of 12ЊCin et al. 2008b, 2009b) and glacial lowstands at Ϫ120 m or lower the northwest Mediterranean region, 7.5ЊC around Sicily, and relative to present sea level (Rohling et al. 1998a; Waelbroeck 6ЊC or less to the south and east of Crete. Thus, mean glacial- et al. 2002). Methods for constructing continuous records of interglacial temperature contrasts are expected on land sea-level variability now exist for the last half-million years around the Red Sea. The inferred summer bias in Arz et al. (Rohling et al. 2009b, 2010; Siddall et al. 2003; Waelbroeck (2003b) and Trommer et al. (2011) suggests that the 4Њ–6ЊC et al. 2002). Model-based deconvolutions of deep-sea stable contrast estimated here may also have some bias toward sum- oxygen isotope records have also been used to extend the mer (i.e., winter contrasts may have been smaller). record over many millions of years (Bintanja, van de Wal, A notable southward displacement of climate zones also and Oerlemans 2005; De Boer et al. 2010, 2011), but these occurred in glacials on both orbital and suborbital timescales still require independent validation (fig. 4). (e.g., Wang et al. 2004, 2006), as well as global lowering of Along with enlarged land-ice volumes, another defin- the snow line and associated vertical compression of vege- ing characteristic of glacials was widespread cooling (e.g., tation zones (e.g., Barmawidjaja et al. 1993; Broecker and CLIMAP Project Members 1976; MARGO Project Members Denton 1989; Klein, Isacks, and Bloom 1995; Kuhlemann et 2009; Rohling et al. 2012; Schneider von Deimling et al. 2006). al. 2008). Such snow line and vegetation displacements were In the study region, cooling produced a strong gradient from particularly strongly expressed in the mountain ranges along west to east through the Mediterranean and then to the south- the northern margin of the Mediterranean (Kuhlemann et al. east along the axis of the Red Sea. Relative to the present, 2008). glacial sea surface temperatures in summer were reduced by In a general sense, summer circulation/monsoons were ,2ЊC weakened in the Northern Hemisphere during glacial periodsע2ЊC in the westernmost Mediterranean, 5Њע8Њ ,.2ЊC in the easternmost Mediterra- and winter circulation/monsoons were strengthened (e.gעaround Sicily, and 3Њ Rohling, Mayewski, and Challenor 2003; Rohling et al. 2009c עnean; winter sea surface temperatures were reduced by 5Њ .(1ЊC, and 0Њ–2.5ЊC, respectively (Hayes et al. 2005). and references thereinע2ЊC, 3Њ

Figure 4. Comparison between three approaches for reconstructing continuous records of sea-level variations. Black is Waelbroeck et al. (2002), based on coral-calibrated deep-sea stable oxygen isotope data, with uncertainties in gray. The dashed line is the reconstruction of De Boer et al. (2010, 2011) from a model-based deconvolution of deep-sea stable oxygen isotope records into a temperature and a sea-level component following the method of Bintanja, van de Wal, and Oerlemans (2005). Dots with (2σ) uncertainty bars represent the Red Sea reconstruction on a U-Th-adjusted chronology (Rohling et al. 2009b, 2010). Modified after Rohling et al. (2012). A color version of this figure is available in the online edition of Current Anthropology. Rohling et al. Mediterranean and Red Sea Paleoclimate S189

Overall, atmospheric dust transport was strongly intensified smaller amplitude than that observed in Greenland (Blunier during glacials (e.g., Lambert et al. 2008; Larrasoan˜a et al. et al. 1998; EPICA Community Members 2006). Continuous 2003; Mayewski et al. 1997; Roberts et al. 2011; Rohling, records of sea-level variability, developed from Red Sea oxygen Mayewski, and Challenor 2003; Ruth et al. 2007; Trauth, Lar- isotope records, indicate that within the last glacial cycle, rasoan˜a, and Mudelsee 2009; Winckler et al. 2008), which global ice volume fluctuated on millennial timescales with a attests to increased aridity, stronger winds, and reduced veg- rhythm close to that of variability observed in Antarctic cli- etation cover (reduced soil cohesion). However, even within mate records (Grant et al. 2012; Rohling et al. 2004a, 2009b; the Mediterranean and Red Sea region, there can be consid- Siddall et al. 2003, 2008). erable spatial differences in dust flux histories because of spa- Greenland ice-core records and North Atlantic marine sed- tially different conditions in the various dust source areas. iment records provide evidence of particularly strong climate The western Mediterranean receives dust from northwest Af- fluctuations that have become known as Dansgaard-Oeschger rica/western Sahara, while the eastern Mediterranean receives cycles, which include the particularly cold Heinrich events dust from the eastern Sahara (Libya, Egypt). The Red Sea (e.g., Broecker 2000; Dansgaard et al. 1993; Grootes et al. receives influxes of windblown dust from the easternmost 1993; Hemming 2004). In many records, these millennial- Sahara, Sudan, and Saudi Arabia (Hickey and Goudie 2007; scale fluctuations appear as an alternation between more in- Jiang et al. 2009; Middleton and Goudie 2001; fig. 5). On tense and less intense glacial conditions, and the Heinrich millennial timescales, dust variability from various source ar- events are often particularly cold/intense. Western Mediter- eas varies considerably within and around the Mediterranean ranean sea surface temperature strongly fluctuated in close and Red Seas; these variations do not seem to be systematic agreement with the Dansgaard-Oeschger cycles and Heinrich between the two basins (Roberts et al. 2011; fig. 6). This events (Cacho et al. 1999, 2000, 2001; Frigola et al. 2007; suggests considerable regional differences in the temporal var- Martrat et al. 2004; Rohling et al. 1998b). iability of vegetation cover, soil cohesion, and wind patterns/ Dansgaard-Oeschger oscillations in the Northern Hemi- intensities. sphere were related to Antarctic (Southern Hemisphere) tem- Ice-core records from Greenland and Antarctica reveal that perature cycles through a systematic out-of-phase relationship glacial periods were characterized by strong temperature fluc- (e.g., Blunier and Brook 2001; Blunier et al. 1998; EPICA tuations on millennial timescales (fig. 7). Antarctic ice-core Community Members 2006; Stocker and Johnsen 2003). In records reveal climate variability that was less abrupt and of this relationship, which has become known as the “bipolar

Figure 5. Major global dust sources and locations of dust records. Dust flux contours (mg mϪ2 yrϪ1) are shown in oceans surrounding dust sources (after Duce et al. 1991). Locations of dust records discussed are indicated (ocean drilling program sites 659: Tiedemann, Sarntheim, and Shackleton 1994; 721: deMenocal, Bloemendal, and King 1991; 967: Larrasoan˜a et al. 2003; and the Zhaojiachuan and Lingtai loess sections: Sun et al. 2006). Site KL09 is the Red Sea record presented in Rohling et al. (2008a, 2009b) and Roberts et al. (2011). Modified after Roberts et al. (2011). A color version of this figure is available in the online edition of Current Anthropology. Figure 6. Comparison of sea-level and dust records for the Red Sea, circum-Sahara region, and Chinese loess plateau. a, Central Red Sea sea-level reconstruction (Rohling et al. 2009b, 2010). b, Environmental magnetic IRM900 mT@AF120 mT proxy for windblown hematite (Hem.) in Red Sea core KL09 (gray) compared with the stacked Chinese loess grain size record from Zhaojiachuan Rohling et al. Mediterranean and Red Sea Paleoclimate S191 temperature seesaw,” the magnitude of warming in the South- eccentricity (e.g., COHMAP Members 1988; Kutzbach and ern Hemisphere is proportional to the duration of cold ep- Gallimore 1988; Kutzbach and Guetter 1986; Kutzbach and isodes in the Northern Hemisphere (e.g., EPICA Community Street-Perrott 1985; Rossignol-Strick 1985). African monsoon Members 2006; Stocker and Johnsen 2003). Siddall et al. maxima are well known to have been associated with inso- (2010) identified during which intervals of the last 500,000 lation maxima. Freshwater flooding from the African margin yr such millennial-scale climate variability has been particu- into the Mediterranean during these times caused collapse of larly pronounced and found that this was the case at 480– deepwater formation, eventually resulting in deepwater an- 460, 440–400, 380–360, 340–320, 260–220, 200–160, 140–120, oxia. These events are easily recognized in eastern Mediter- and 80–40 ka (fig. 7). Because of the global nature of this ranean sedimentary sequences because organic carbon pres- variability, it may be expected that these intervals of generally ervation led to the formation of characteristic black organic enhanced climate variability will be noticeable in records from rich layers, called sapropels (e.g., Rohling 1994 and references Africa and Eurasia even if the exact nature of the variability therein). may differ between regions. The Mediterranean sapropel record reflects the more in- Northern high-latitude cooling events particularly affected tense African monsoon maxima, and individual sapropels the northern sectors of the Mediterranean region because of have been dated (Emeis et al. 2000; Hilgen 1991; Hilgen et cold air outbreaks that were channeled toward the basin al. 1993; Kroon et al. 1998; Lourens, Wehausen, and Brumsack through gaps in the mountain ranges along its northern limits, 2001; Lourens et al. 1996) based on the ages of precession- which also triggered atmospheric instability over the Medi- driven insolation maxima that are known from astronomical terranean and had implications for regional precipitation re- solutions (Berger 1977; Laskar et al. 2004; Milankovitch 1941). gimes (e.g., Casford et al. 2003; Frigola et al. 2007; Kuhlemann A particularly straightforward summary table of ages is given et al. 2008; Rohling et al. 1998b, 2002b). Thus, vegetation for the main sapropels by Kroon et al. (1998), after Lourens (pollen) records from the western Mediterranean and north- et al. (1996). These datings may be important for archaeo- ern sector of the eastern Mediterranean also reflect the strong logical, anthropological, and biogeographical studies because effects of northerly (Greenland-style) climate influences (e.g., they provide a chronological framework that may help us to Allen et al. 1999; Kotthoff et al. 2008; Moreno et al. 2002; date and understand, for example, past migration pulses Mu¨ller and Pross 2007; Sa´nchez-Gon˜i et al. 2002; Tzedakis through the Sahara desert region (e.g., Drake et al. 2011; 1999, 2009; Tzedakis et al. 2004). Osborne et al. 2008) or past humid phases associated with From the Red Sea area, relatively little is known about fossil hominid finds (e.g., Brown, McDougall, and Fleagle climate variability. It was always dry, with high windblown 2012; McDougall, Brown, and Fleagle 2005, 2008). The sap- dust input, but this input nevertheless reveals intensity var- ropel record indicates that there have been more than a hun- iations that are closely similar to those observed on the Chi- dred of such strongly developed African monsoon maxima nese loess plateau (Roberts et al. 2011). This suggests that the over the past couple of million years (Larrasoan˜a, Roberts, main climate influences over the central Red Sea (with respect and Rohling 2013). to windblown dust input) were dominated by atmospheric During times with intensified African monsoon circulation, circulation/wind changes that reflect the larger westerlies- the spatial extent of the Sahara Desert was much reduced, dominated Northern Hemisphere climate variability and which has become known as “greening of the Sahara,” when Indian-Asian monsoon variability (e.g., Porter and An 1995; the African monsoon penetrated farther northward than to- Rohling, Mayewski, and Challenor 2003). day (Larrasoan˜a, Roberts, and Rohling 2013). This penetra- tion was partly due to orbital forcing and partly resulted from African Monsoon Changes northward expansion of vegetation into the previously more Superimposed on the glacial cycles, the entire Mediterranean reflective desert, which in turn triggered further northward region is strongly affected by monsoon intensity variations, penetration of the monsoon front in a “vegetation-albedo which are dominated by Northern Hemisphere insolation feedback process” (e.g., Brovkin et al. 1998; Claussen et al. changes that mainly reflect the influences of precession and 1998; Foley et al. 2003). During intense monsoon maxima,

and Lingtai, Chinese loess plateau (from Sun et al. 2006, with minor age adjustments in Roberts et al. 2011). c, d, e, and f, respectively, compare the Red Sea dust record (gray) with dust records from EPICA Dome C, Antarctica (Lambert et al. 2008); ODP Site 659, off northwest Africa (Tiedemann, Sarntheim, and Shackleton 1994); ODP Site 967, eastern Mediterranean Sea (Larrasoan˜a et al. 2003); and ODP Site 721, Arabian Sea (deMenocal, Bloemendal, and King 1991). Vertical dashed lines coincide with Red Sea dust peaks at the glacial terminations and are shown to assist comparisons between panels. The Red Sea records are shown for consistency on the same chronology (Rohling et al. 2009b, 2010) as used in Roberts et al. (2011). This chronology is (subtly) updated within the last 150,000 yr in figures 7 and 8 based on the latest age controls developed by Grant et al. (2012). A color version of this figure is available in the online edition of Current Anthropology. S192 Current Anthropology Volume 54, Supplement 8, December 2013

Figure 7. Comparison of different records of climate variability including polar ice-core records, sea level, and Mediterranean temperature variability. a, EPICA Dome C ice-core stable hydrogen isotope proxy for temperature, Antarctica (Jouzel et al. 2007). b, The Red Sea sea-level record in the pre-150 ka interval on the chronology of Rohling et al. (2009b, 2010) and in the post-150 ka interval on the latest chronology using the age constraints developed by Grant et al. (2012). c, Sea surface temperature in the Alboran Sea, westernmost Mediterranean (Martrat et al. 2004). d, Stacked North-GRIP ice-core oxygen isotope proxy for temperature, Greenland (gray; Wolff et al. 2010), and—for clarity—a 21-point moving average smoothing (black). Bars at the bottom indicate intervals of particularly pronounced millennial-scale climate variability after the analysis of Siddall et al. (2010). A color version of this figure is available in the online edition of Current Anthropology. the monsoon front appears to have penetrated northward past decreases/collapses in monsoon intensity occurred during sev- the central Saharan watershed (at about 21ЊN), and seasonal eral monsoon maxima and that these roughly coincided with runoff occurred from the central Saharan mountains into the intrusions of northerly cooling events into the Mediterranean eastern Mediterranean along the wider North African margin basin (e.g., Casford et al. 2003; Gasse 2000; Mercone et al. (Drake et al. 2011; Osborne et al. 2008; Paillou et al. 2009; 2001; Osborne et al. 2008; Rohling et al. 2002a, 2002b, 2004b; Rohling et al. 2002a, 2004a, 2004b). Such drainages will have Scrivner, Vance, and Rohling 2004). presented green corridors along which humans and animals It is important to note that there is no evidence that the may have migrated across the otherwise arid region (Drake African monsoon penetrated at any time directly into the et al. 2011; Osborne et al. 2008). These ideas integrate a wide Mediterranean basin. Rainfall gradient and isotope recon- variety of evidence for past increases in precipitation and structions in the Levant indicate that rain in that region was water availability throughout the currently hyperarid Sahara, always sourced from the north and west, from the Mediter- including the distributions of animal and human fossils, sed- ranean, even during monsoon maxima (Bar-Matthews et al. imentological data, lake-level reconstructions, etc. (e.g., Ar- 2003; Goodfriend 1991; Matthews, Ayalon, and Bar-Matthews mitage et al. 2007; Cremaschi 2002; Drake et al. 2011; Gasse 2000; McGarry et al. 2004; Vaks et al. 2007). Moreover, Tze- 2000; Gaven et al. 1981; Kuper and Kro¨pelin 2006; Larrasoan˜a, dakis (2009) presented a compelling case that times of Roberts, and Rohling 2013; Mandell and Simmons 2001; Pa- insolation-driven African monsoon maxima were not char- chur 2001; Pachur and Altmann 2006; Pachur and Braun acterized by enhanced summer precipitation around the 1980; Szabo, Haynes, and Maxwell 1995). Records of African northern and eastern Mediterranean, as had been often sug- monsoon variability reveal that centennial- to millennial-scale gested before (e.g., Rohling and Hilgen 1991; Tzedakis 2009; Rohling et al. Mediterranean and Red Sea Paleoclimate S193 and references therein). Pollen data indicate that summer for all three major monsoon systems (African, Indian, and aridity was enhanced at these times and that any enhanced Southeast Asian). rainfall likely took place in winter (with some possible regional Red Sea records contain no evidence for any major pre- exceptions); that is, typical Mediterranean climate conditions cipitation/vegetation changes associated with monsoon max- were intensified (Tzedakis 2009). ima; the area appears to have remained (hyper)arid. Monsoon During opposite phases of the precession cycle, the mon- variability, however, affected Red Sea oceanography through soon was weak, and the (hyper)arid Sahara desert was spatially changes in the wind field over the basin (Biton et al. 2010; extended, similar to the present, without watercourses Trommer et al. 2011), and strong windblown dust variations crossing it to the wider North African margin. Orbital cyclicity over time support the notion of important fluctuations in in the African monsoon is obvious in records of windblown wind forcing over the basin (Roberts et al. 2011; Rohling et dust fluxes from the Sahara (e.g., Larrasoan˜a et al. 2003; Lour- al. 2008a). ens, Wehausen, and Brumsack 2001; Trauth, Larrasoan˜a, Millennial-scale variability in windblown dust records of Mudelsee 2009; Wehausen and Brumsack 2000), which re- the central Red Sea is coherent with millennial-scale changes flects insolation forcing of the African monsoon. in Arabian Sea productivity (Rohling et al. 2008a; Schultz, von Rad, Erlenkeuser 1998) as well as with Chinese loess records (Porter and An 1995; Roberts et al. 2011; Sun et al. Indian Monsoon Changes 2006). This links Arabia with winter-dominated climate var- Variations in the Indian Ocean monsoon circulation, which iability as recorded in Greenland ice-core records (Rohling, affect the wind field over the southern sector of the Red Sea Mayewski, and Challenor 2003). In short, there is good evi- region, have been documented especially by marine sedi- dence that colder conditions in Greenland coincided with mentary records from the Arabian Sea (e.g., Almogi-Labin et intensified winter-type atmospheric circulation over Asia, al. 2000; Clemens and Prell 1990, 2003; Clemens et al. 1991; which also affected Arabia, possibly through the winter Ivanochko et al. 2005; Prell and Kutzbach 1987; Reichart, (northeast) monsoon. Lourens, Zachariasse 1998; Rostek et al. 1997; Schmiedl and Leuschner 2005; Schultz, von Rad, and Erlenkeuser 1998; Sir- ocko et al. 1993) and by speleothem records from Oman and Implications and Conclusions Socotra (Burns et al. 2003, 2004; Fleitmann et al. 2003a, Conditions Relevant to Proposed Migration Routes 2003b, 2004, 2007). Long Arabian Sea records provide con- across the Southern Red Sea vincing evidence for a predominant control of orbital pre- cession and thus insolation on the Indian monsoon intensity, Sea level. The intense ice-age cycles of the last 500,000 yr similar to the African monsoon. Speleothem data from Oman have been associated with important variability in global sea (Fleitmann et al. 2003a, 2003b, 2004, 2007) indicate that the level over a range of 120 m or more below the present to precipitation regime of the Indian Ocean monsoon expanded perhaps 10 m above the present (fig. 4). The Strait of Bab- to affect the southeastern margin of the Arabian Peninsula el-Mandab in the southern Red Sea, which connects the basin during the Early to Middle Holocene summer southwest with the open ocean, is highly sensitive to sea-level change Monsoon maximum, whereas that region currently (during because it is (today) only 137 m deep. This is of the same the opposite precession phase) is unaffected (Conroy and order as past sea-level drops during glacial maxima, and there Overpeck 2011). have consequently been many proposals of a potential mi- Trommer et al. (2011) described the timing of a somewhat gration route across the southern Red Sea, assuming that a more humid interval in the central Red Sea Bakala (Tokar) passage between Africa and Arabia may have emerged during wadi catchment and found that this started immediately fol- times of maximum glacial sea-level lowstands. lowing the last interglacial sea-level highstand and lasted sev- Fernandes, Rohling, and Siddall (2006) evaluated the con- eral thousand years. This relative timing agrees with the period cept of emergence of a southern land bridge between Africa of deposition of the deep-sea anoxic event known as sapropel and Arabia and concluded that there is no evidence for it at S5 in the eastern Mediterranean (Grant et al. 2012), which is any time during the last 500,000 yr. Emergence of a land also recognized as a humid interval in the speleothem record bridge in the strait would rapidly lead to desiccation within of Soreq cave, Israel, which reflects the last interglacial African the highly evaporative Red Sea, with deposition of evaporites monsoon maximum (Bar-Matthews, Ayalon, and Kaufman within a matter of several centuries in the open basin and 1997, 2000; Bar-Matthews et al. 1999, 2003). U-series dating within a matter of years to decades in shallow coastal envi- of the Soreq cave record demonstrates that the last interglacial ronments. Moreover, there is evidence of substantial persistent African monsoon maximum dates to 128–120 ka. This agrees local water depths above the sill in the strait of at least 15 m with datings from Dongge cave, China, for the last interglacial (Fernandes, Rohling, and Siddall 2006) and up to 35 m (Biton, to 119.6 Gildor, and Peltier 2008); for the Last Glacial Maximum, the 0.9 ע maximum of the Asian monsoon, from 129.3 m (Lambeck 4 ע ka (Yuan et al. 2004). On millennial scales, therefore, passage depth has been estimated as up to 25 0.6 ע the timing of monsoon maxima seems to be roughly similar et al. 2011). For lower water depths, inflow from the open S194 Current Anthropology Volume 54, Supplement 8, December 2013 ocean would have become sufficiently restricted for basin- et al. 2003b; Vaks et al. 2007), a little later than in the eastern wide development of extreme salinities in excess of about 75 Sahara (from 140 ka; Osmond and Dabous 2004; Szabo, practical salinity units (psu). There is evidence that salinities Haynes, and Maxwell 1995; Vaks et al. 2007). at times rose above 49 psu, causing local extinction of plank- It would be an oversimplification to assume a simple suc- tonic foraminifera (Fenton et al. 2000 and references therein). cession from low sea level to favorable climate for migration However, salinities remained below 75 psu given that higher across the Arabian Peninsula during the transition from the values would have also caused the local extinction of all pter- Saalian glacial to the last interglacial. The data instead reveal opods and all benthic foraminifera, which did not happen considerable complexity across that transition, highlighting (Fenton et al. 2000; Rohling et al. 1998a). that the sea-level lowstand and the climatically more favorable Regardless of the above, the strait passage was much shal- conditions were separated by at least 5,000 yr of arid regional lower and narrower than today during glacial sea-level low- conditions and sharp sea-level rise (Roberts et al. 2011). This stands (Lambeck et al. 2011; Rohling et al. 1998a; Siddall et does not exclude the possibility that the southern route out al. 2004). This is especially clearly illustrated by a recent ad- of Africa was employed at this time. But if it was, then the vanced reconstruction of strait morphology during the Last animals/humans involved must have been resilient to signif- Glacial Maximum (Lambeck et al. 2011), which takes into icantly adverse environmental conditions. We note that a sim- account detailed hydrographic data for the present-day strait, ilar sequence of events is observed for almost all glacial ter- sea-level change during the Last Glacial Maximum, and a minations. In the next section, we elaborate a new view of model for isostatic change components. Lambeck et al. (2011) more promising migration intervals. also present arguments about older glacial lowstands and sup- port the notion that an open passage remained in existence. A new concept: “windows of opportunity” for southern mi- In summary, it is evident from the combined studies that any gration out of Africa. Our sea-level and windblown dust rec- migration across the southern Red Sea during glacial sea-level ords, sampled from the same central Red Sea sediment se- lowstands would have (a) benefited from the fact that the quence (Rohling et al. 2008a, 2009b, 2010; Roberts et al. 2011) marine passage was strongly reduced in width, and (b) def- are shown in figure 8. The last 150,000 yr are shown on a initely included some element of swimming, rafting, or nav- new chronology (Grant et al. 2012, which is tightly con- igation. strained relative to the U-Th dated Soreq Cave record from Israel; Bar-Matthews, Ayalon, and Kaufman 1997; Bar- Climate conditions. Another important control on potential Matthews et al. 1999, 2000, 2003). Before 150 ka, the records migrations across the southern Red Sea concerns regional are shown on a chronology developed (Rohling et al. 2009b) climatic conditions. For example, was enough water and food by correlation with the European Project for Ice Coring in available to sustain migrating animals/humans on either side Antarctica (EPICA) Dome C temperature proxy record on of the strait? This question has come to the fore because of the EDC3 chronology (Jouzel et al. 2007) with an adjustment a recent proposal (Armitage et al. 2011) that an early wave that accounts for radiometric datings of past sea-level high- of human migration out of Africa occurred across the south- stands (Rohling et al. 2010). ern Red Sea, with migration toward and across the strait The horizontal light gray bar in figure 8 highlights relatively region during the penultimate glacial maximum (the “Saa- low dust fluxes similar to those of the Holocene. During times lian”) and subsequent spreading across/along the Arabian with such relatively low dust fluxes, we suggest that the re- Peninsula during the monsoon maximum of the last inter- gional climate may have been most favorable for habitation/ glacial (about 128–120 ka). migration in contrast to the intervening intervals with high Red Sea sedimentary records provide important insights dust fluxes, which attest to more intense winds and lower soil into regional climatic conditions. Windblown dust concen- cohesion due to even lower soil moisture and even more trations have been measured in exactly the same sedimentary scarce vegetation cover than today. The vertical light gray bars sequences that were used to reconstruct the Red Sea sea-level identify periods when both (a) sea level stood 100 m or more record. This dust record clearly illustrates that the transition below the present, so that the marine connection to be crossed between the Saalian glacial and the last interglacial was char- in the Strait of Bab el Mandab would have been only some acterized by extreme fluxes of windblown dust, which reflect 6 km wide (cf. Last Glacial Maximum reconstruction of high winds and pronounced aridity in the region (Roberts et Lambeck et al. 2011); and (b) dust fluxes were relatively low. al. 2011). Organic geochemical data, also from the same sam- These highlighted intervals therefore represent periods with ple sequence, demonstrate that (lightly) enhanced humidity relatively favorable conditions (windows of opportunity) for in the Red Sea region—the most likely local expression of the potential migrations between northeast Africa and southwest last interglacial summer monsoon maximum—first developed Arabia via a southern route. There is increasing evidence that only after the sea-level highstand had peaked and sea level the southern route was used for the final migration out of had started to drop again (Trommer et al. 2011). Data from Africa by AMHs (e.g., Derricourt 2005; Fernandes et al. 2012). Oman suggest that relatively more humid conditions may The highlighted periods of significant sea-level lowstands have started to develop earlier, from about 135 ka (Fleitmann were also characterized by emerged continental shelves Rohling et al. Mediterranean and Red Sea Paleoclimate S195

Figure 8. Direct comparison between (a) the latest version of the Red Sea sea-level reconstruction, with the pre-150 ka interval on the chronology of Rohling et al. (2009b, 2010) and the post-150 ka interval on the new chronology of Grant et al. (2012), and (b) central Red Sea dust proxy data, including the Ti/Ca ratio from core-scanning XRF analysis (gray) and hematite concentration data from environmental magnetic analyses (smooth line; see also Roberts et al. 2011; Rohling et al. 2008a). Data originate from a single sampling of the same sedimentary sequence, which ensures unambiguous phase relationships between the various records. Light gray bars are explained in the text. Horizontal bars at bottom indicate intervals of particularly pronounced millennial-scale climate variability, after the analysis of Siddall et al. (2010; see also fig. 7). A color version of this figure is available in the online edition of Current Anthropology. around the Arabian Peninsula. These provided excellent hab- ϩ5/Ϫ8 ka is in remarkable agreement with our window of itation and migration potential for animals and humans, es- opportunity of 70–65 ka for a southern route migration out pecially if the enhanced hydraulic head due to sea-level low- of Africa. The chronological uncertainties for our records in ering led to enhanced freshwater seepage on the emerged that interval are less than 1,000 yr (Grant et al. 2012). Hap- shelves from groundwater reservoirs (e.g., Parker and Rose lotype dispersal maps suggest that the migration at around 2008). Such seepage exists even today on the shelves, in sub- 65 ka followed the southern route out of Africa and that marine form (Ghoneim 2008; Parker and Rose 2008). The Arabia was the first staging post in the spread of AMHs emerged shelves may therefore have presented an excellent around the world (Fernandes et al. 2012). habitat with gentle topography, freshwater availability, and an abundance of coastal/marine resources. If emerged shelves Conditions Relevant to Habitation of and Migration were key migration/habitation zones, then much of the an- through the Sahara thropological record may now be under water. Several windows of opportunity for migration along the Migration potential through the Sahara Desert was compre- southern route out of Africa within the last half-million years hensively assessed by Drake et al. (2011). Their abstract says are highlighted in figure 8. These windows date to about 458– it all: 448, 345–340, 272–265, 145–140, and 70–65 ka. The interval Both animals and humans populated it [the Sahara] during of 145–140 ka may be relevant with respect to a tentative early past humid phases....Moreanimals crossed via this route migration of AMHs that led to early inhabitation at the Hor- than used the Nile corridor . . . [and] many of these species muz region of the Arabian Gulf with an oldest dating of 127 are aquatic. This dispersal was possible because during the ka (Armitage et al. 2011). That population is thought Holocene humid period the region contained a series of 16 ע to have been “unsuccessful” in that it did not leave any de- linked lakes, rivers, and inland deltas comprising a large scendants (Fernandes et al. 2012). Instead, all non-African interlinked waterway, channeling water and animals into humans originate from a more recent migration out of Africa and across the Sahara. . . . This system was last active in dated with various molecular clock approaches at around 57– the early Holocene when many species appear to have oc- 65 ka, with an upper age bound of 70–65 ka based on East cupied the entire Sahara. Human dispersals were influenced African data (Fernandes et al. 2012). This age range of 65 by this distribution. . . . Lacustrine sediments show that the S196 Current Anthropology Volume 54, Supplement 8, December 2013

“green Sahara” also existed during the last interglacial (∼125 region was intermittently more humid (especially between 133 ka) and provided green corridors that could have formed and 122 ka) and thus more hospitable (Vaks et al. 2007). This dispersal routes at a likely time for the migration of modern closely matches data that indicate more humid conditions in humans out of Africa. the Egyptian Sahara (Osmond and Dabous 2004), which sug- gests a high probability that a pathway existed through the Both periods highlighted in that study are well-known Levant for migrations out of Africa between about 140 and insolation-driven African monsoon maxima. The Holocene 110 ka, with possible extension to 85 ka, after which hyperarid period is known to correspond to eastern Mediterranean sap- conditions were reestablished (Vaks et al. 2007). ropel S1 and the last interglacial period to sapropel S5 (e.g., Derricourt (2005) reviewed climatic and archaeological Larrasoan˜a, Roberts, and Rohling 2013; Osborne et al. 2008; studies to conclude that the Levantine route was the most Rohling and Hilgen 1991; Rohling et al. 2002a,2004b; Scriv- likely route of early (pre-85 ka) migrations out of Africa in ner, Vance, and Rohling 2004). contrast to the youngest (post-85 ka) migration, which he Archaeological observations around exclusively rain-fed de- links more to a southern route across the Bab-el-Mandab pressions on the Libyan Plateau suggest that monsoonal sum- Strait (see above). Early AMH finds from Qafzeh and Skhul ka 13 ע and 81 18 ע mer rains from central Africa periodically penetrated at least in the Near East date to between 119 Њ as far north as Kharga (roughly 25 N) during the Holocene (e.g., Armitage et al. 2011; Gru¨n et al. 2005; Petraglia 2011; monsoon maximum despite the fact that conditions during Shea 2008). Genetic data indicate that this migration did not that pluvial phase seem to have remained drier than during leave any descendants in the modern human population out- earlier Quaternary pluvial phases (Mandel and Simmons side Africa, which instead can be traced back to a migration 2001). This suggestion that the Holocene monsoon maximum out of Africa that took place around 65 ka, likely via a south- was of a relatively low intensity compared with previous Qua- ern route (Fernandes et al. 2012; see above). ternary monsoon maxima has been corroborated by quan- For Levantine migration routes, the key control may have titative reconstructions of effects of the Holocene and last been exerted by development of pluvial episodes as recorded interglacial monsoon maxima in the eastern Mediterranean and accurately dated in cave speleothem deposits (e.g., Der- (Rohling 1999; Rohling et al. 2004b). Hence, past monsoon ricourt 2005; Shea 2008; Vaks et al. 2007). These intermittently maxima—datings for which can be obtained from the astro- more humid conditions in the Levant are closely related to nomical ages of insolation-driven monsoon maxima (Emeis monsoon maxima associated with Northern Hemisphere in- et al. 2000; Hilgen 1991; Hilgen et al. 1993; Kroon et al. 1998; solation maxima. It should be emphasized that monsoon Lourens et al. 1996, 2001)—represent times of enhanced hu- maxima, with “greening of the Sahara,” were not limited to midity that may have been crucial for migrations through the only interglacial insolation maxima but occurred also during otherwise hyperarid Sahara “barrier” between sub-Saharan insolation maxima within glacial times (e.g., Larrasoan˜a, Rob- Africa and the Mediterranean/Levantine regions (Larrasoan˜a, erts, and Rohling 2013; Larrasoan˜a et al. 2003; Liu et al. 2012). Roberts, and Rohling 2013). The control by pluvials that applies to the Levantine route As mentioned above, several African monsoon maxima is considerably more straightforward than the windows of (e.g., the Holocene and last interglacial) have been interrupted opportunity control we have proposed in “A new concept: by centennial- to millennial-scale periods of reduced mon- ‘windows of opportunity’ for southern migration out of Af- soon intensity. Hence, potential routes for migration through rica” for the southern route, which requires a combination the Sahara region remained intermittent, subject to periodic of low sea level and concomitant interludes of relatively fa- returns of harsh conditions. vorable climate conditions. These windows of opportunity Finally, hypotheses that invoke an importance of periods are not simply aligned with monsoon maxima but instead of rapid, millennial-scale climate variability for developments reflect millennial-scale episodes of relatively favorable climate and/or migrations of hominins and their food sources—for within glacial maxima. example, through intermittent habitat fragmentation—may benefit from the objectively identified episodes of generally (globally) enhanced millennial-scale climate variability (Sid- Conditions along the European Margin of the Mediterranean dall et al. 2010). We have indicated these intervals (480–460, As recorded in Greenland ice cores, the European margin of 440–400, 380–360, 340–320, 260–220, 200–160, 140–120, and the Mediterranean has been strongly affected by intense cli- 80–40 ka) in figures 7 and 8. mate swings that originate in the North Atlantic region. Cold events were transmitted to the Mediterranean by northerly air outbreaks through gaps in the mountainous topography Conditions Relevant to Migrations through the Levant around the northern margin of the Mediterranean, especially Conditions for migrations through the Levant are normally over the northwest Mediterranean, the Adriatic Sea, and the poor because of hyperarid conditions in the Sinai-Negev re- Aegean (e.g., Casford et al. 2003; Frigola et al. 2007; Kuhle- gion. However, data from a north–south array of caves high- mann et al. 2008; Moreno et al. 2002; Rohling et al. 1998b, light a window of time between 140 and 110 ka when this 2002b). During the Holocene, a strong temporal coincidence Rohling et al. Mediterranean and Red Sea Paleoclimate S197 has been found between the northerly cooling events and Red Sea marine environment during termination I. Paleoceanography 18: 1053, doi:10.1029/2002PA000864. archaeological transitions in the northeastern sector of the Bar-Matthews, M., A. Ayalon, M. Gilmour, A. Matthews, and C. J. Hawkes- Mediterranean region (Clare et al. 2008). Because of resultant worth. 2003. Sea-land oxygen isotopic relationships from planktonic fo- atmospheric instabilities within the Mediterranean basin, raminifera and speleothems in the Eastern Mediterranean region and their implication for paleorainfall during interglacial intervals. Geochimica et Cos- northerly cooling events may have had an amplified effect on mochimica Acta 67:3181–3199. snow line lowerings and vegetation-zone migrations in the Bar-Matthews, M., A. Ayalon, and A. Kaufman. 1997. Late Quaternary pa- Mediterranean region (e.g., Kuhlemann et al. 2008 and ref- leoclimate in the eastern Mediterranean region from stable isotope analysis of speleothems at Soreq Cave, Israel. Quaternary Research 47:155–168. erences therein). ———. 2000. Timing and hydrological conditions of sapropel events in the Despite this glacial variability, the Mediterranean region Eastern Mediterranean, as evident from speleothems, Soreq Cave, Israel. during glacial times was a refugium with relatively mild con- Chemical Geology 169:145–156. Bar-Matthews, M., A. Ayalon, A. Kaufman, and G. J. Wasserburg. 1999. The ditions for plants and animals, avoiding the much more severe Eastern Mediterranean paleoclimate as a reflection of regional events: Soreq climatic stress over the main Eurasian landmass to the north Cave, Israel. Earth and Planetary Science Letters 166:85–95. of the Alps/Carpathians (e.g., Blondel 2009; Tzedakis 2009). Barmawidjaja, D. M., E. J. Rohling, W. A. Van der Kaars, C. Vergnaud-Graz- zini, and W. J. Zachariasse. 1993. Glacial conditions in the northern Molucca The region’s great diversity of terrain and microclimates, Sea region (Indonesia). Palaeogeography, Palaeoclimatology, Palaeoecology drainage patterns, and accentuated relief with many rock shel- 101:147–167. ters and caves would have been conducive to a relatively rich Berger, A. 1977. Support for the astronomical theory of climatic change. Nature 268:44–45. variety of exploitable plant and animal food sources and to Be´thoux, J. P. 1984. Paleo-hydrologie de la Mer Mediterranee au cours des camp locations. From a climatic point of view, therefore, it derniers 20 000 ans. Oceanologica Acta 7:43–48. is likely that habitation and migration were possible through- Be´thoux, J. P., and B. Gentili. 1994. The Mediterranean Sea: a test area for marine and climatic interactions. In Ocean processes in climate dynamics: out this region even during severe glacials, especially in low- global and Mediterranean examples. P. Malanotte-Rizzoli and A. R. Rob- lands and coastal plains, and that there was the potential to inson, eds. Pp. 239–254. Dordrecht: Kluwer. range into higher elevations during periods with milder con- Bintanja, R., R. S. W. van de Wal, and J. Oerlemans. 2005. Modelled atmo- spheric temperatures and global sea levels over the past million years. Nature ditions. 437:125–128. Biton, E., H. Gildor, and W. R. Peltier. 2008. Relative sea level reduction at the Red Sea during the Last Glacial Maximum. Paleoceanography 23:PA1214, doi:10.1029/2007PA001431. Acknowledgments Biton, E., H. Gildor, G. Trommer, M. Siccha, M. Kucera, M. T. J. van der Meer, and S. Schouten. 2010. Sensitivity of Red Sea circulation to monsoonal Thanks to the Wenner-Gren Foundation for organizing the variability during the Holocene: an integrated data and modeling study. conference and arranging for publication of the papers pre- Paleoceanography 25:PA4209, doi:10.1029/2009PA001876. sented there and to Erella Hovers and Steve Kuhn and two Blondel, J. 2009. The nature and origin of the vertebrate fauna. In The physical geography of the Mediterranean. J. C. Woodward, ed. Pp. 139–163. Oxford: anonymous reviewers for their constructive assessments of Oxford University Press. earlier versions of the manuscript. This paper contributes to Blunier, T., and E. Brook. 2001. Timing of millennial-scale climate change in the United Kingdom Natural Environment Research Council Antarctica and Greenland during the last glacial period. Science 291:109– 112. projects NE/E01531X/1 (RESET), NE/I009906/1 (iGlass), and Blunier, T., J. Chapellaz, J. Schwander, A. Da¨llenbach, B. Stauffer, T. F. Stocker, NE/H004424/1 as well as E. J. Rohling’s Royal Society– D. Raynaud, et al. 1998. Asynchrony of antarctic and Greenland climate Wolfson Research Merit Award and Australian Laureate Fel- change during the last glacial period. Nature 394:739–743. Boucher, K. 1975. Global climate. London: English University Press. lowship FL120100050. Braconnot, P., B. Otto-Bliesner, S. Harrison, S. Joussaume, J. Y. Peterchmitt, A. Abe-Ouchi, M. Crucifix, et al. 2007. Results of PMIP2 coupled simu- lations of the mid-Holocene and Last Glacial Maximum. 1. Experiments References Cited and large-scale features. Climate of the Past 3:261–277. Adamson, D. A., F. Gasse, F. A. Street, and M. A. J. Williams. 1980. Late Broecker, W. S. 2000. Abrupt climate change: causal constraints provided by Quaternary history of the Nile. Nature 288:50–55. the paleoclimate record. Earth-Science Reviews 51:137–154. Allen, J. R. M., U. Brandt, A. Brauer, H. W. Hubberten, B. Huntley, J. Keller, Broecker, W. S., and G. H. Denton. 1989. The role of ocean-atmosphere M. Kraml, et al. 1999. Rapid environmental changes in southern Europe reorganizations in glacial cycles. Geochimica et Cosmochimica Acta 53:2465– during the last glacial period. Nature 400:740–743. 2501. Almogi-Labin, A., G. Schmiedl, C. Hemleben, R. Siman-Tov, M. Segl, and D. Brovkin, V., M. Claussen, V. Petoukhov, and A. Ganopolski. 1998. On the Meischner. 2000. The influence of the NE winter monsoon on productivity stability of the atmosphere-vegetation system in the Sahara/Sahel region. changes in the Gulf of Aden, NW Arabian Sea, during the last 530 ka as Journal of Geophysical Research 103:31613–31624. recorded by benthic foraminifera. Marine Micropaleontology 40:295–319. Brown, F. H., I. McDougall, and J. G. Fleagle. 2012. Correlation of the KHS Armitage, S. J., N. A. Drake, S. Stokes, A. El-Hawat, M. Salem, K. White, P. Tuff of the Kibish Formation to volcanic ash layers at other sites, and the Turner, and S. J. McLaren. 2007. Multiple phases of North African humidity age of early Homo sapiens (Omo I and Omo II). Journal of Human Evolution recorded in lacustrine sediments from the Fazzan Basin, Libyan Sahara. 63:577–585. Quaternary Geochronology 2:181–186. Burns, S. J., D. Fleitmann, A. Matter, J. Kramers, and A. A. Al-Subbary. 2003. Armitage, S. J., S. A. Jasim, A. E. Marks, A. G. Parker, V. I. Usik, and H. P. Indian Ocean climate and an absolute chronology over Dansgaard/Oeschger Uerpmann. 2011. The southern route “Out of Africa”: evidence for an early events 9 to 13. Science 301:1365–1367. expansion of modern humans into Arabia. Science 331:453–456. ———. 2004. Indian Ocean climate and an absolute chronology over Dans- Arz, H. W., F. Lamy, J. Pa¨tzold, P. J. Mu¨ller, and M. Prins. 2003a. Mediter- gaard/Oeschger events 9 to 13: correction. Science 305:1567. ranean moisture source for an Early-Holocene humid period in the Cacho, I., J. O. Grimalt, M. Canals, L. Sbaffi, N. J. Shackleton, J. Scho¨nfeld, Northern Red Sea. Science 300:118–121. and R. Zahn. 2001. Variability of the western Mediterranean Sea surface Arz, H. W., J. Pa¨tzold, P. J. Mu¨ller, and M. O. Moammar. 2003b. Influence temperatures during the last 25,000 years and its connection with the of Northern Hemisphere climate and global sea level rise on the restricted Northern Hemisphere climatic changes. Paleoceanography 16:40–52. S198 Current Anthropology Volume 54, Supplement 8, December 2013

Cacho, I., J. O. Grimalt, C. Pelejero, M. Canals, F. J. Sierro, J. A. Flores, and Fenton, M., S. Geiselhart, E. J. Rohling, and C. Hemleben. 2000. Aplanktonic N. J. Shackleton. 1999. Dansgaard-Oeschger and Heinrich event imprints zones in the Red Sea. Marine Micropaleontology 40:277–294. in Alboran Sea paleotemperatures. Paleoceanography 14:698–705. Fernandes, C., E. J. Rohling, and M. Siddall. 2006. Absence of Quaternary Cacho, I., J. O. Grimalt, F. J. Sierro, N. J. Shackleton, and M. Canals. 2000. Red Sea land bridges: biogeographic implications. Journal of Biogeography Evidence of enhanced Mediterranean thermohaline circulation during rapid 33:961–966. climate coolings. Earth and Planetary Science Letters 183:417–429. Fernandes, V., F. Alshamali, M. Alves, M. D. Costa, J. B. Pereira, N. M. Silva, Cantu, V. 1977. The climate of Italy. In Climates of central and southern Europe. L. Cherni, et al. 2012. The Arabian cradle: mitochondrial relicts of the first C. C. Wallen, ed. Pp. 127–183. World Survey of Climatology, vol. 6. Am- steps along the southern route out of Africa. American Journal of Human sterdam: Elsevier. Genetics 90:1–9. Casford, J. S. L., E. J. Rohling, R. H. Abu-Zied, F. J. Jorissen, M. Leng, and Fleitmann, D., S. J. Burns, A. Mangini, M. Mudelsee, J. Kramers, I. Villa, U. J. Thomson. 2003. A dynamic concept for eastern Mediterranean circulation Neff, et al. 2007. Holocene ITCZ and Indian monsoon dynamics recorded and oxygenation during sapropel formation. Palaeogeography, Palaeocli- in stalagmites from Oman and Yemen (Socotra). Quaternary Science Reviews matology, Palaeoecology 190:103–119. 26:170–188. Clare, L., E. J. Rohling, B. Weninger, and J. Hilpert. 2008. Warfare in Late Fleitmann, D., S. J. Burns, M. Mudelsee, U. Neff, J. Kramers, A. Mangini, Neolithic/Early Chalcolithic Pisidia, southwestern Turkey: climate induced and A. Matter. 2003a. Holocene forcing of the Indian monsoon recorded social unrest in the late 7th millennium cal BC. Documenta Praehistorica in a stalagmite from Southern Oman. Science 300:1737–1739. 35:65–92. Fleitmann, D., S. J. Burns, U. Neff, A. Mangini, and A. Matter. 2003b. Chang- Claussen, M., V. Brovkin, A. Ganopolski, C. Kubatski, and V. Petoukhov. ing moisture sources over the last 330,000 years in Northern Oman from 1998. Modelling global terrestrial vegetation-climate interaction. Philosoph- fluid-inclusion evidence in speleothems. Quaternary Research 60:223–232. ical Transactions of the Royal Society B 353:53–63. Fleitmann, D., S. J. Burns, U. Neff, M. Mudelsee, A. Mangini, and A. Matter. Clemens, S. C., and W. L. Prell. 1990. Late Pleistocene variability of Arabian 2004. Palaeoclimatic interpretation of high-resolution oxygen isotope pro- Sea summer monsoon winds and continental aridity: eolian records from files derived from annually laminated speleothems from Southern Oman. the lithogenic component of deep-sea sediments. Paleoceanography 5:109– Quaternary Science Reviews 23:935–945. 145. Foley, J. A., M. T. Coe, M. Scheffer, and G. Wang. 2003. Regime shifts in the ———. 2003. A 350,000 year summer-monsoon multi-proxy stack from the Sahara and Sahel: interactions between ecological and climatic systems in Owen Ridge, northern Arabian Sea. Marine Geology 201:35–51. Northern Africa. Ecosystems 6:524–539. Clemens, S. C., W. L. Prell, D. Murray, G. Shimmield, and G. Weedon. 1991. Frigola, J., A. Moreno, I. Cacho, M. Canals, F. J. Sierro, J. A. Flores, J. O. Forcing mechanisms of the Indian Ocean monsoon. Nature 353:720–725. Grimalt, D. A. Hodell, and J. H. Curtis. 2007. Holocene climate variability CLIMAP Project Members. 1976. The surface of the ice-age earth. Science in the western Mediterranean region from a deepwater sediment record. 191:1131–1137. Paleoceanography 22:PA2209, doi:10.1029/2006PA001307. COHMAP Members. 1988. Climatic changes of the last 18,000 years: obser- Gasse, F. 2000. Hydrological changes in the African tropics since the Last vations and model simulations. Science 241:1043–1052. Glacial Maximum. Quaternary Science Reviews 19:189–211. Conroy, J. L., and J. T. Overpeck. 2011. Regionalization of present-day pre- Gaven, C., C. Hillaire-Marcel, and N. Petit-Maire. 1981. A Pleistocene lacus- cipitation in the greater monsoon region of Asia. Journal of Climate 24: trine episode in southeastern Libya. Nature 290:131–133. 4073–4095. Ghoneim, E. 2008. Optimum groundwater locations in the northern United Cremaschi, M. 2002. Late Pleistocene and Holocene climatic changes in the Arab Emirates. International Journal of Remote Sensing 29:5879–5906. central Sahara: the case study of the south-western Fezzan, Libya. In Goodfriend, G. A. 1991. Holocene trends in 18O in land snail shells from the Droughts, food and culture: ecological change and food security in Africa’s Negev Desert and their implications for changes in rainfall source areas. later prehistory. F. A. Hassan, ed. Pp. 65–81. New York: Kluwer/Plenum. Quaternary Research 35:417–426. Dansgaard, W., S. J. Johnsen, H. B. Clausen, D. Dahl-Jensen, N. S. Gundestrup, Grant, K. M., E. J. Rohling, M. Bar-Matthews, A. Ayalon, M. Medina-Elizalde, C. U. Hammer, C. S. Hvidberg, et al. 1993. Evidence for general instability C. Bronk Ramsey, C. Satow, and A. P. Roberts. 2012. Rapid coupling be- of past climate from a 250 kyr ice core. Nature 364:218–219. tween ice volume and polar temperature over the past 150 kyr. Nature 491: De Boer, B., R. S. W. van de Wal, R. Bintanja, L. J. Lourens, and E. Tuenter. 744–747. 2010. Cenozoic global ice-volume and temperature simulations with 1-D Grootes, P. M., M. Stuiver, J. W. C. White, S. Johnsen, and J. Jouzel. 1993. ice-sheet models forced by benthic δ18O records. Annals of Glaciology 51: Comparison of oxygen isotope records from the GISP2 and GRIP Greenland 23–33. ice cores. Nature 366:552–554. De Boer, B., R. S. W. van de Wal, L. J. Lourens, and R. Bintanja. 2011. Transient Gru¨n, R., C. Stringer, F. McDermott, R. Nathan, N. Porat, S. Robertson, L. nature of the earth’s climate and the implications for the interpretation of Taylor, G. Mortimer, S. Eggins, and M. McCulloch. 2005. U-series and ESR benthic records. Palaeogeography, Palaeoclimatology, Palaeoecology 335/336: analyses of bones and teeth relating to the human burials from Skhul. 4–11. Journal of Human Evolution 49:316–334. deMenocal, P., J. Bloemendal, and J. King. 1991. A rock-magnetic record of Hayes, A., M. Kucera, N. Kallel, L. Sbaffi, and E. J. Rohling. 2005. Glacial monsoonal dust deposition to the Arabian Sea: evidence for a shift in the Mediterranean sea surface temperatures reconstructed from planktonic fo- mode of deposition at 2.4 Ma. In Proceedings of the Ocean Drilling Program, raminiferal assemblages. Quaternary Science Reviews 24:999–1016. Scientific Results, vol. 117. Pp. 389–407. College Station, TX: Ocean Drilling Hays, J. D., J. Imbrie, and N. J. Shackleton. 1976. Variations in the earth’s Program. orbit: pacemaker of the ice ages. Science 194:1121–1132. Derricourt, R. 2005. Getting “out of Africa”: sea crossings, land crossings and Hemming, S. R. 2004. Heinrich events: massive Late Pleistocene detritus layers culture in the hominin migrations. Journal of World Prehistory 19:119–132. of the North Atlantic and their global imprint. Reviews of Geophysics 42:1– Drake, N. A., R. M. Blench, S. J. Armitage, C. S. Bristow, and K. H. White. 43. 2011. Ancient watercourses and biogeography of the Sahara explain the Hickey, B., and A. S. Goudie. 2007. The use of TOMS and MODIS to identify peopling of the desert. Proceedings of the National Academy of Sciences of dust storm source areas: the Tokar delta (Sudan) and the Seistan basin the USA 108:458–462. (south west Asia). In Geomorphological variations. A. S. Goudie, and J. Duce, R. A., P. S. Liss, J. T. Merrill, E. L. Atlas, P. Buat-Menard, B. B. Hicks, Kalvoda, J., eds. Pp. 37–57. Prague: P3K. J. M. Miller, et al. 1991. The atmospheric input of trace species to the world Hilgen, F. J. 1991. Astronomical calibration of Gauss to Matuyama sapropels ocean. Global Biogeochemical Cycles 5:193–259. in the Mediterranean and implication for the geomagnetic polarity time El-Fandy, M. G. 1946. Barometric lows of Cyprus. Quarterly Journal of the scale. Earth and Planetary Science Letters 104:226Ϫ244. Royal Meteorological Society 72:291–306. Hilgen, F. J., L. J. Lourens, A. Berger, and M. F. Loutre. 1993. Evaluation of Emeis, K. C., T. Sakamoto, R. Wehausen, and H. J. Brumsack. 2000. The the astronomically calibrated time-scale for the late Pliocene and earliest sapropel record of the eastern Mediterranean Sea: results of Ocean Drilling Pleistocene. Paleoceanography 8:549Ϫ565. Program Leg 160. Palaeogeography, Palaeoclimatology, Palaeoecology 158: Imbrie, J., and J. Z. Imbrie. 1980. Modeling the climatic response to orbital 371–395. variations. Science 207:943–953. EPICA (European Project for Ice Coring in Antarctica) Community Members. Imbrie, J., and K. P Imbrie. 1986. Ice ages: solving the mystery. Cambridge, 2006. One-to-one coupling of glacial climate variability in Greenland and MA: Harvard University Press. Antarctica. Nature 444:195–197. Ivanochko, T. S., R. S. Ganeshram, G. J. A. Brummer, G. Ganssen, S. J. A. Rohling et al. Mediterranean and Red Sea Paleoclimate S199

Jung, S. G. Moreton, and D. Kroon. 2005. Variations in tropical convection circum-Mediterranean region from magnetic and geochemical observations as an amplifier of global climate change at the millennial scale. Earth and of sapropels S1, S5 and S6. Palaeogeography, Palaeoclimatology, Palaeoecology Planetary Science Letters 235:302–314. 333/334:1–12. Jiang, H., J. T. Farrar, R. C. Beardsley, R. Chen, and C. Chen. 2009. Zonal Lolis, C. J., A. Bartzokas, and B. D. Katsoulis. 2002. Spatial and temporal 850 surface wind jets across the Red Sea due to mountain gap forcing along hPa air temperature and sea-surface temperature covariances in the Med- both sides of the Red Sea. Geophysical Research Letters 36:L19605, doi: iterranean region and their connection to atmospheric circulation. Inter- 10.1029/2009GL040008. national Journal of Climatology 22:663–676. Jouzel, J., V. Masson-Delmotte, O. Cattani, G. Dreyfus, S. Falourd, G. Hoff- Lourens, L. J., A. Antonarakou, F. J. Hilgen, A. A. M. van Hoof, C. Vergnaud- mann, B. Minster, et al. 2007. Orbital and millennial Antarctic climate Grazzini, and W. J. Zachariasse. 1996. Evaluation of the Plio-Pleistocene variability over the past 800,000 years. Science 317:793–797. astronomical timescale. Paleoceanography 11:391–413. Klein, A. G., B. Isacks, and A. L. Bloom. 1995. Modern and Last Glacial Lourens, L. J., R. Wehausen, and H. J. Brumsack. 2001. Geological constraints Maximum snowline in Peru and Bolivia: implications for regional climatic on tidal dissipation and dynamical ellipticity of the earth over the past three change. Bulletin de l’Institut Franc¸ais d’Etudes Andines 24:607–617. million years. Nature 409:1029–1033. Kopp, R. E., F. J. Simons, J. X. Mitrovica, A. C. Maloof, and M. Oppenheimer. Maheras, P., E. Xoplaki, T. Davies, J. Martin-Vide, M. Bariendos, and M. J. 2009. Probabilistic assessment of sea level during the last interglacial stage. Alcoforado. 1999. Warm and cold monthly anomalies across the Mediter- Nature 462:863–868. ranean basin and their relationship with circulation 1860–1990. Interna- Kottek, M., J. Grieser, C. Beck, B. Rudolf, and F. Rubel. 2006. World map of tional Journal of Climatology 19:1697–1715. the Ko¨ppen-Geiger climate classification updated. Meteorologische Zeitschrift Maillard, C., and G. Soliman. 1986. Hydrography of the Red Sea and exchanges 15:259–263. with the Indian Ocean in summer. Oceanologica Acta 9:249–269. Kotthoff, U., U. C. Muller, J. Pross, G. Schmiedl, I. T. Lawson, and H. Schulz. Malanotte-Rizzoli, P., and A. Bergamasco. 1991. The wind and thermally 2008. Late glacial and Holocene vegetation dynamics in the Aegean region: driven circulation of the eastern Mediterranean Sea. 2. The baroclinic case. an integrated view based on pollen data from marine and terrestrial archives. Dynamics of Atmospheres and Oceans 15:355–419. Holocene 18:1019–1032. Mandel, R. D., and A. H. Simmons. 2001. Prehistoric occupation of Late Kroon, D., I. Alexander, M. Little, L. J. Lourens, A. Matthewson, A. H. F. Quaternary landscapes near Kharga Oasis, western desert of Egypt. Geo- Robertson, and T. Sakamoto. 1998. Oxygen isotope and sapropel stratig- archaeology 16:95–117. raphy in the Eastern Mediterranean during the last 3.2 million years. In MARGO Project Members. 2009. Constraints on the magnitude and patterns Proceedings of the Ocean Drilling Program, Scientific Results, vol. 160. Pp. of ocean cooling at the Last Glacial Maximum. Nature Geoscience 2:127– 181–190. College Station, TX: Ocean Drilling Program. 132. Kuhlemann, J., E. J. Rohling, I. Kumrei, P. Kubik, S. Ivy-Ochs, and M. Kucera. Martrat, B., J. O. Grimalt, C. Lopez-Martinez, I. Cacho, F. J. Sierro, J. A. 2008. Regional synthesis of Mediterranean atmospheric circulation during Flores, R. Zahn, M. Canals, J. H. Curtis, and D. A. Hodell. 2004. Abrupt the Last Glacial Maximum. Science 321:1338–1340. temperature changes in the western Mediterranean over the past 250,000 Kuper, R., and S. Kro¨pelin. 2006. Climate-controlled Holocene occupation in years. Science 306:1762–1765. the Sahara: motor of Africa’s evolution. Science 313:803–807. Matthews, A., A. Ayalon, and M. Bar-Matthews. 2000. D/H ratios of fluid Kutzbach, J. E., and R. G. Gallimore. 1988. Sensitivity of a coupled atmo- inclusions of Soreq Cave (Israel) speleothems as a guide to the Eastern sphere/mixed layer ocean model to changes in orbital forcing at 9000 years. Mediterranean Meteoric Line relationships in the last 120 ky. Chemical Journal of Geophysical Research 93:803–821. Geology 166:183–191. Kutzbach, J. E., and P. J. Guetter. 1986. The influence of changing orbital Mayewski, P. A., L. D. Meeker, M. S. Twickler, S. Whitlow, Q. Yang, W. B. parameters and surface boundary conditions on climate simulations for the Lyons, and M. Prentice. 1997. Major features and forcing of high-latitude past 18,000 years. Journal of Atmospheric Science 43:1726–1759. Kutzbach, J. E., and F. A. Street-Perrott. 1985. Milankovitch forcing in the Northern Hemisphere atmospheric circulation using a 110,000-year-long level of tropical lakes from 18 to 0 kyr. Nature 317:130–134. glaciochemical series. Journal of Geophysical Research 102:26345–26366. Laıˆne´, A., M. Kageyama, P. Braconnot, and R. Alkama. 2009. Impact of green- McDougall, I., F. H. Brown, and J. G. Fleagle. 2005. Stratigraphic placement house gas concentration changes on surface energetics in IPSL-CM4: re- and age of modern humans from Kibish, Ethiopia. Nature 433:733–736. gional warming patterns, land-sea warming ratios, and glacial-interglacial ———. 2008. Sapropels and the age of hominins Omo I and II, Kibish, differences. Journal of Climate 22:4621–4635. Ethiopia. Journal of Human Evolution 55:409–420. Lambeck, K., A. Purcell, N. C. Fleming, C. Vita-Finzi, A. M. Alsharekh, and McGarry, S., M. Bar-Matthews, A. Matthews, A. Vaks, B. Schilman, and A. G. M. Bailey. 2011. Sea level and shoreline reconstructions for the Red Sea: Ayalon. 2004. Constraints on hydrological and paleotemperature variations isostatic and tectonic considerations and implications for hominin migra- in the eastern Mediterranean region in the last 140 ka given by the δD tion out of Africa. Quaternary Science Reviews 30:3542–3574. values of speleothem fluid inclusions. Quaternary Science Reviews 23:919– Lambert, F., B. Delmonte, J. R. Petit, M. Bigler, P. R. Kaufmann, M. A. Hutterli, 934. T. F. Stocker, U. Ruth, J. P. Steffensen, and V. Maggi. 2008. Dust-climate Mercone, D., J. Thomson, R. H. Abu-Zied, I. W. Croudace, and E. J. Rohling. couplings over the past 800,000 years from the EPICA Dome C ice core. 2001. High-resolution geochemical and micropalaeontological profiling of Nature 452:616–619. the most recent eastern Mediterranean sapropel. Marine Geology 177:25– Larrasoan˜a, J. C., A. P. Roberts, and E. J. Rohling. 2013. Dynamics of green 44. Sahara periods and their role in hominin evolution. PLoS One 8:e76514, Middleton, N. J., and A. S. Goudie. 2001. Saharan dust: sources and trajec- doi:10.1371/journal.pone.0076514. tories. Transactions of the Institute of British Geographers 26:165–181. Larrasoan˜a, J. C., A. P. Roberts, E. J. Rohling, M. Winklhofer, and R. Wehausen. Milankovitch, M. 1941. Kanon der Erdbestrahlung und seine Anwendung auf 2003. Three million years of monsoon variability over the northern Sahara. das Eiszeitenproblem. Royal Serbian Academy Special Publications, vol. 132; Climate Dynamics 21:689–698. Section Mathematics and Natural Sciences, vol. 33. Belgrade: Royal Serbian Laskar, J., M. Gastineau, F. Joutel, P. Robutel, B. Levrard, et al. 2004. A long- Academy. term numerical solution for the insolation quantities of the Earth. Astron- Morcos, S. A. 1970. Physical and chemical oceanography of the Red Sea. omy and Astrophysics 428:261–285. Oceanography and Marine Biology 8:73–202. Leaman, K. D., and F. A. Schott. 1991. Hydrographic structure of the con- Moreno, A., I. Cacho, M. Canals, M. A. Prins, M. F. Sa´nchez-Gon˜i,J.O. vection regime in the Gulf of Lions: winter 1987. Journal of Physical Ocean- Grimalt, and G. J. Weltje. 2002. Saharan dust transport and high-latitude ography 21:575–598. glacial climatic variability: the Alboran Sea record. Quaternary Research 58: Legge, H. L., J. Mutterlose, and H. W. Arz. 2006. Climatic changes in the 318–328. northern Red Sea during the last 22,000 years as recorded by calcareous Muhs, D., K. R. Simmons, R. R. Schumann, and R. B. Halley. 2011. Sea-level nannofossils. Paleoceanography 21:PA1003, doi:10.1029/2005PA001142. history of the past two interglacial periods: new evidence from U-series Lisiecki, L. E., and M. E. Raymo. 2005. A Plio-Pleistocene stack of 57 globally dating of reef corals from south Florida. Quaternary Science Reviews 30: distributed benthic δ18O records. Paleoceanography 20:PA1003, doi:10.1029 570–590. /2004PA001071. Mu¨ller, U., and J. Pross. 2007. Lesson from the past: present insolation min- Liu, Q., J. C. Larrasoan˜a, J. Torrent, A. P. Roberts, E. J. Rohling, Z. Liu, and imum holds potential for glacial inception. Quaternary Science Reviews 26: Z. Jiang. 2012. New constraints on climate forcing and variability in the 3025–3029. S200 Current Anthropology Volume 54, Supplement 8, December 2013

Naval Oceanography Command. 1987. U.S. Navy climatic study of the Med- and M. Kucera. 2009b. Antarctic temperature and global sea level closely iterranean Sea. Ashville, NC: Naval Oceanography Command Detachment. coupled over the past five glacial cycles. Nature Geoscience 2:500–504. Nof, D. 1979. On man-induced variations in the circulation of the Mediter- Rohling, E. J., K. Grant, C. Hemleben, M. Kucera, A. P. Roberts, I. Schmeltzer, ranean Sea. Tellus 31:558–564. H. Schulz, M. Siccha, M. Siddall, and G. Trommer. 2008a. New constraints Osborne, A. H., D. Vance, E. J. Rohling, N. Barton, M. Rogerson, and N. on the timing and amplitude of sea level fluctuations during Marine Isotope Fello. 2008. A humid corridor across the Sahara for the migration “out of Stage 3. Paleoceanography 23:PA3219, doi:10.1029/2008PA001617. Africa” of early modern humans 120,000 years ago. Proceedings of the Na- Rohling, E. J., K. Grant, C. Hemleben, M. Siddall, B. A. A. Hoogakker, M. tional Academy of Sciences of the USA 105:16444–16447. Bolshaw, and M. Kucera. 2008b. High rates of sea-level rise during the last Osmond, J. K., and A. A. Dabous. 2004. Timing and intensity of groundwater interglacial period. Nature Geoscience 1:38–42. movement during Egyptian Sahara pluvial periods by U-series analysis of Rohling, E. J., A. Hayes, D. Kroon, S. De Rijk, and W. J. Zachariasse. 1998b. secondary U in ores and carbonates. Quaternary Research 61:85–94. Abrupt cold spells in the NW Mediterranean. Paleoceanography 13:316– Pachur, H. J. 2001. Holoza¨ne Klimawechsel in den no¨rdlichen Subtropen. 322. Nova Acta Leopoldina NF88 331:109–131. Rohling, E. J., and F. J. Hilgen. 1991. The eastern Mediterranean climate at Pachur, H. J., and N. Altmann. 2006. Die Ostsahara im Spa¨tquarta¨r. Berlin: times of sapropel formation: a review. Geologie en Mijnbouw 70:253–264. Springer. Rohling, E. J., Q. Liu, A. P. Roberts, J. D. Stanford, S. O. Rasmussen, P. L. Pachur, H. J., and G. Braun. 1980. The paleoclimate of the central Sahara, Langen, and M. Siddall. 2009c. Controls on the East Asian monsoon during Libya and the Libyan Desert. Palaeoecology of Africa 12:351–363. the last glacial cycle, based on comparison between Hulu Cave and polar Paillou, P., M. Schuster, S. Tooth, T. Farr, A. Rosenqvist, S. Lopez, and J. M. ice-core records. Quaternary Science Reviews 28:3291–3302. Malezieux. 2009. Mapping of a major paleodrainage system in eastern Libya Rohling, E. J., R. Marsh, N. J. Wells, M. Siddall, and N. Edwards. 2004a. using orbital imaging radar: the Kufrah River. Earth and Planetary Science Similar melt-water contributions to glacial sea-level variability from ant- Letters 277:327–333. arctic and northern ice sheets. Nature 430:1016–1021. Parker, A. G., and J. I. Rose. 2008. Climate change and human origins in Rohling, E. J., P. A. Mayewski, and P. Challenor. 2003. On the timing and southeastern Arabia. Proceedings of the Seminar on Arabian Studies 38:25–42. mechanism of millennial-scale climate variability during the last glacial Patzert, W. C. 1974. Wind induced reversal in Red Sea circulation. Deep-Sea cycle. Climate Dynamics 20:257–267. Research 21:109–121. Rohling, E. J., P. A. Mayewski, A. Hayes, R. H. Abu-Zied, and J. S. L. Casford. Pedgley, D. E. 1974. An outline of the weather and climate of the Red Sea. 2002b. Holocene atmosphere-ocean interactions: records from Greenland Publications Centre National pour l’Exploitation des Oceans, Actes du Col- and the Aegean Sea. Climate Dynamics 18:587–593. loques 2:9–21. Rohling, E. J., M. Medina-Elizalde, J. G. Shepherd, M. Siddall, and J. D. Petraglia, M. D. 2011. Trailblazers across Arabia. Nature 470:50–51. Stanford. 2012. Sea surface and high-latitude temperature sensitivity to Porter, S. C., and Z. An. 1995. Correlation between climate events in the radiative forcing of climate over several glacial cycles. Journal of Climate North Atlantic and China during the last glaciation. Nature 375:305–308. 25:1635–1656. Poulos, S. E., P. G. Drakopoulos, and M. B. Collins. 1997. Seasonal variability Rohling, E. J., M. Sprovieri, T. R. Cane, J. S. L. Casford, S. Cooke, I. Bou- in sea surface oceanographic conditions in the Aegean Sea (eastern Med- loubassi, and K. C. Emeis. 2004b. Reconstructing past planktic foraminiferal iterranean): an overview. Journal of Marine Systems 13:225–244. habitats using stable isotope data: a case history for Mediterranean sapropel Prell, W. L., and J. E. Kutzbach. 1987. Monsoon variability over the past S5. Marine Micropaleontology 50:89–123. 150,000 years. Journal of Geophysical Research 92:8411–8425. Rossignol-Strick, M. 1985. Mediterranean Quaternary sapropels, an immediate Privett, D. W. 1959. Monthly charts of evaporation from the North Indian response of the African monsoon to variation of insolation. Palaeogeography, Ocean, including the Red Sea and Persian Gulf. Quarterly Journal of the Palaeoclimatology, Palaeoecology 49:237–263. Meteorological Society 85:424–428. Rostek, F., E. Bard, L. Beaufort, C. Sonzogni, and G. Ganssen. 1997. Sea Reichart, G. J. 1997. Late Quaternary variability of the Arabian Sea monsoon surface temperature and productivity records for the past 240 kyr in the and oxygen minimum zone. Geologica Ultraiectina, no. 154. PhD thesis, Arabian Sea. Deep Sea Research II 44:1461–1480. Universiteit Utrecht. Rumney, G. R. 1968. Climatology and the world’s climates. New York: Mac- Reichart, G. J., L. J. Lourens, and W. J. Zachariasse. 1998. Temporal variability millan. in the northern Arabian Sea oxygen minimum zone (OMZ) during the last Ruth, U., M. Bigler, R. Ro¨thlisberger, M. L. Siggaard-Andersen, J. Kipfstuhl, 225,000 years. Paleoceanography 13:607–621. K. Goto-Azuma, M. E. Hansson, S. J. Johnsen, H. Lu, and J. P. Steffensen. Roberts, A. P., E. J. Rohling, K. M. Grant, J. C. Larrasoan˜a, and Q. Liu. 2011. 2007. Ice core evidence for a very tight link between North Atlantic and Atmospheric dust variability from major global source regions over the last East Asian glacial climate. Geophysical Research Letters 34:L03706, doi: 500,000 years. Quaternary Science Reviews 30:3537–3541. 10.1029/2006GL027876. Rodwell, M. J., and B. J. Hoskins. 1996. Monsoons and the dynamics of deserts. Saaroni, H., A. Bitan, P. Alpert, and B. Ziv. 1996. Continental polar outbreaks Quarterly Journal of the Royal Meteorological Society 122:1385–1404. into the Levant and Eastern Mediterranean. International Journal of Cli- Rohling, E. J. 1994. Review and new aspects concerning the formation of matology 16:1175–1191. Mediterranean sapropels Marine Geology 122:1–28. Said, R. 1981. The geological evolution of the River Nile. New York: Springer. ———. 1999. Environmental controls on salinity and δ18O in the Mediter- Sa´nchez-Gon˜i, M. F., I. Cacho, J. L. Turon, J. Guiot, F. J. Sierro, J. P. Pey- ranean. Paleoceanography 14:706–715. pouquet, J. O. Grimalt, and N. J. Shackleton. 2002. Synchroneity between ———. 2013. Quantitative assessment of glacial fluctuations in the level of marine and terrestrial responses to millennial scale climatic variability dur- Lake Lisan, Dead Sea rift. Quaternary Science Reviews 70:63–72. ing the last glacial period in the Mediterranean region. Climate Dynamics Rohling, E. J., R. Abu-Zied, J. S. L. Casford, A. Hayes, and B. A. A. Hoogakker. 19:95–105. 2009a. The marine environment: present and past. In The physical geography Schmiedl, G., and D. C. Leuschner. 2005. Oxygenation changes in the deep of the Mediterranean. J. C. Woodward, ed. Pp. 33–67. Oxford: Oxford Uni- western Arabian Sea during the last 190,000 years: productivity versus deep- versity Press. water circulation. Paleoceanography 20:A2008, doi:10.1029/2004PA001044. Rohling, E. J., K. Braun, K. Grant, M. Kucera, A. P. Roberts, M. Siddall, and Schneider von Deimling, T., A. Ganopolski, H. Held and S. Rahmstorf. 2006. G. Trommer. 2010. Comparison between Holocene and marine isotope How cold was the Last Glacial Maximum? Geophysical Research Letters 33: stage-11 sea-level histories. Earth and Planetary Science Letters 291:97–105. L14709, doi:10.1029/2006GL026484. Rohling, E. J., and H. L. Bryden. 1992. Man-induced salinity and temperature Schulz, H., U. von Rad, and H. Erlenkeuser. 1998. Correlation between Ara- increases in Western Mediterranean Deep Water. Journal of Geophysical bian Sea and Greenland climate oscillations of the past 110,000 years. Nature Research 97:11191–11198. 393:54–57. Rohling, E. J., T. R. Cane, S. Cooke, M. Sprovieri, I. Bouloubassi, K. C. Emeis, Scrivner, A. E., D. Vance, and E. J. Rohling. 2004. New neodymium isotope R. Schiebel, et al. 2002a. African monsoon variability during the previous data quantify Nile involvement in Mediterranean anoxic episodes. Geology interglacial maximum. Earth and Planetary Science Letters 202:61–75. 32:565–568. Rohling, E. J., M. Fenton, F. J. Jorissen, P. Bertrand, G. Ganssen, and J. P. Shea, J. J. 2008. Transitions or turnovers? climatically-forced extinctions of Caulet. 1998a. Magnitudes of sea-level lowstands of the past 500,000 years. Homo sapiens and Neanderthals in the east Mediterranean Levant. Qua- Nature 394:162–165. ternary Science Reviews 27:2253–2270. Rohling, E. J., K. Grant, M. Bolshaw, A. P. Roberts, M. Siddall, C. Hemleben, Siddall, M., E. J. Rohling, A. Almogi-Labin, C. Hemleben, D. Meischner, I. Rohling et al. Mediterranean and Red Sea Paleoclimate S201

Schmeltzer, and D. A. Smeed. 2003. Sea-level fluctuations during the last Tzedakis, P. C. 1999. The last climatic cycle at Kopais, central Greece. Journal glacial cycle. Nature 423:853–858. of the Geological Society, London 156:425–434. Siddall, M., E. J. Rohling, T. Blunier, and R. Spahni. 2010. Patterns of millennial ———. 2009. Cenozoic climate and vegetation change. In The physical ge- variability over the last 500 ka. Climate of the Past 6:295–303. ography of the Mediterranean. J. C. Woodward, ed. Pp. 89–137. Oxford: Siddall, M., E. J. Rohling, W. G. Thompson, and C. Waelbroeck. 2008. Marine Oxford University Press. isotope stage 3 sea level fluctuations: data synthesis and new outlook. Re- Tzedakis, P. C., M. R. Frogley, I. T. Lawson, R. C. Preece, I. Cacho, and L. views of Geophysics 46:RG4003, doi:10.1029/2007RG000226. de Abreu. 2004. Ecological thresholds and patterns of millennial-scale cli- Siddall, M., D. A. Smeed, C. Hemleben, E. J. Rohling, I. Schmeltzer, and W. mate variability: the response of vegetation in Greece during the last glacial R. Peltier. 2004. Understanding the Red Sea response to sea level. Earth period. Geology 32:109–112. and Planetary Science Letters 225:421–434. Vaks, A., M. Bar-Matthews, A. Ayalon, A. Matthews, L. Halicz, and A. Frumkin. Sirocko, F., M. Sarnthein, H. Erlenkeuser, H. Lange, M. Arnold, and J. C. 2007. Desert speleothems reveal climatic window for African exodus of Duplessy. 1993. Century-scale events in monsoonal climate over the past early modern humans. Geology 35:831–834. 24,000 years. Nature 364:322–324. Waelbroeck, C., L. Labeyrie, E. Michel, J. C. Duplessy, J. F. McManus, K. Stocker, T. F., and S. J. Johnsen. 2003. A minimum thermodynamic Lambeck, E. Balbon, and M. Labracherie. 2002. Sea-level and deep water model for the bipolar seesaw. Paleoceanography 18:1087, doi:10.1029 temperature changes derived from benthic foraminifera isotopic records. /2003PA000920. Quaternary Science Reviews 21:295–305. Sun, Y., S. C. Clemens, Z. An, and Z. Yu. 2006. Astronomic timescale and Wahby, S. D., and N. F. Bishara. 1981. The effect of the river Nile on Med- palaeoclimatic implication of stacked 3.6-myr monsoon records from the iterranean water, before and after the construction of the High Dam at Chinese loess plateau. Quaternary Science Reviews 25:33–48. Aswan. In River inputs to ocean systems. J. M. Martin, J. D. Burton, and D. Eisma, eds. Pp. 311–318. Rome: UNEP-UNESCO. Szabo, B. J., C. V. Haynes Jr., and T. A. Maxwell. 1995. Ages of Quaternary Wang, X., A. S. Auler, R. L. Edwards, H. Cheng, E. Ito, and M. Solheid. 2006. pluvial episodes determined by uranium-series and radiocarbon dating of Interhemispheric anti-phasing of rainfall during the last glacial period. Qua- lacustrine deposits of eastern Sahara. Palaeogeography, Palaeoclimatology, ternary Science Reviews 25:3391–3403. Palaeoecology 113:227–242. Wang, X. F., A. S. Auler, R. L. Edwards, H. Cheng, P. S. Cristalli, P. L. Smart, Tiedemann, R., M. Sarnthein, and N. J. Shackleton. 1994. Astronomical time- D. A. Richards, and C. C. Shen. 2004. Wet periods in northeastern Brazil scale for the Pliocene Atlantic δ18O and dust flux records of Ocean Drilling over the past 210 ka linked to distant climate anomalies. Nature 432:740– Program site 659. Paleoceanography 9:619–638. 743. Trauth, M. H., J. C. Larrasoan˜a, and M. Mudelsee. 2009. Trends, rhythms and Wehausen, R., and H. J. Brumsack. 2000. Chemical cycles in Pliocene sapropel- events in Plio-Pleistocene African climate. Quaternary Science Reviews 28: bearing and sapropel-barren eastern Mediterranean sediments. Palaeogeog- 399–411. raphy, Palaeoclimatology, Palaeoecology 158:325–352. Trewartha, G. T. 1966. The earth’s problem climates. London: Methuen. Whiteman, A. 1971. The geology of the Sudan Republic. Oxford: Clarendon. Trigo, I. F., T. D. Davies, and G. R. Bigg. 1999. Objective climatology in the Williams, M. A. J., D. A. Adamson, B. Cock, and R. McEvedy. 2000. Late Mediterranean region. Journal of Climate 12:1685–1696. Quaternary environments in the White Nile region, Sudan. Global and ———. 2000. Decline in Mediterranean rainfall caused by weakening of Med- Planetary Change 26:305–316. iterranean cyclones. Geophysical Research Letters 27:2913–2916. Winckler, G., R. F. Anderson, M. Q. Fleisher, D. McGee, and N. Mahowald. Trommer, G., M. Siccha, E. J. Rohling, K. Grant, M. T. J. van der Meer, S. 2008. Covariant glacial-interglacial dust fluxes in the equatorial Pacific and Schouten, U. Baranowski, and M. Kucera. 2011. Sensitivity of Red Sea Antarctica. Science 320:93–96. circulation to sea level and insolation forcing during the last interglacial. Wolff, E. W., J. Chappellaz, T. Blunier, S. O. Rasmussen, and A. Svensson. Climate of the Past 7:941–955. 2010. Millennial-scale variability during the last glacial: the ice core record. Trommer, G., M. Siccha, E. J. Rohling, K. Grant, M. T. J. van der Meer, S. Quaternary Science Reviews 29:2828–2838. Schouten, C. Hemleben, and M. Kucera. 2010. Millennial scale variability Yuan, D., H. Cheng, R. L. Edwards, C. A. Dykoski, M. J. Kelly, M. Zhang, J. in Red Sea circulation in response to Holocene insolation forcing. Paleo- Qing, et al. 2004. Timing, duration, and transitions of the last interglacial ceanography 25:PA3203, doi:10.1029/2009PA001826. Asian monsoon. Science 304:575–578. S202 Current Anthropology Volume 54, Supplement 8, December 2013

Neanderthal Demographic Estimates

by Jean-Pierre Bocquet-Appel and Anna Degioanni

CAϩ Online-Only Material: Supplement A

This article offers a critical review of population estimates for the Neanderthal metapopulation based on (paleo-) biological, archaeological, climatic, and genetic data. What do these data tell us about putative Neanderthal de- mography? Biological data suggest a similar demographic frame (life-history traits, such as potential maximum longevity, age at menarche, and duration of gestation) between Neanderthals and modern humans. Archaeological data have revealed a contradiction between the mortality pattern corresponding to 45ϩ yr in Neanderthals and the longevity displayed by the manifest continuum of extant mammals, including primates. Paleoclimatic data suggest that the demography of Neanderthals, living as they did under highly fluctuating climatic conditions, was subject to frequent bottlenecks. This demographic instability combined with the fragmentation of geographical areas and variations in their distribution and extent could account for the fact that potential for technical creativity in the Neanderthal metapopulation would have been limited precisely because of its small numbers, leading it into what is known as a “Boserupian trap” in macrodemographic theory. Finally, genetic literature reports different—but

always very low—estimations of the effective size (Ne) of the Neanderthal metapopulation. It is not easy to relate

Ne to the census size of a population, but by combining different demographic values, this study produced nine different scenarios that were used to obtain an order of magnitude ranging from 5,000 to 70,000 individuals. The cause of the cultural limitation of the Neanderthal metapopulation, compared with that of modern humans, may well have resided in its small numbers alone.

In population biology, it is known that the number of indi- of every kind that one must try to interpret demographically viduals of a species (its metapopulation)—that is, its demo- and to the amount of this information, which decreases dras- graphic size in terms of census data—is a measure of its tically with temporal depth. In this paper, inferences about adaptive success. Thus, a population explosion is the result the Neanderthal metapopulation are critically reviewed on the of successful adaptation. We also know that this number of basis of (paleo-) biological, climatic, archaeological, and pa- individuals determines the rate of evolutionary change. The leogenetic data. What do these data tell us about the putative rate is fast when the population size is small, initiating a demography of Neanderthals? stochastic evolutionary direction; it is slow when a large pop- ulation pushes through the filter of purifying selection and against the resistance of niche construction (Kendal, Tehrani, Demographic Inferences from (Paleo-) and Odling-Smee 2011). This shows the value of linking de- Biological Data: The Frame of mography to paleoanthropology. But there are many diffi- Reference of Extant Mammals culties, mainly due to the nature of nondedicated information Estimation of Potential Maximum Longevity in Mammals

Jean-Pierre Bocquet-Appel is Professor at the E´ cole Pratique des For nearly 50 yr, thanks to the research initiated by Sacher Hautes E´ tudes (Paris) and Research Director at the Centre National (1959, 1975) and Sacher and Staffeldt (1974), a close rela- de la Recherche Scientifique (UPR2147 44, rue de l’Amiral Mouchez, tionship has been known in extant mammals, including pri- 75014 Paris, France [[email protected]]). mates, between maximum potential longevity (L) and the Anna Degioanni is Assistant Professor in the Department of biometric characters of brain weight (E) and body weight (P; Anthropology at Aix-Marseille Universite´, Centre National de la Cutler 1975; Hofman 1993; for a review, see Hawkes 2006). Recherche Scientifique (UMR 7269, Centre National de la Recherche This relationship is based on the biological quasi continuum Scientifique, Maison Me´diterrane´enne des Sciences de l’Homme, Laboratoire Me´diterrane´endePre´histoire [Europe-Afrique], BP 647, between related species, which are connected by their phy- 5 rue du Chaˆteau de l’Horloge, 13094 Aix-en-Provence cedex 2, logenies, and the many similarities they generate. Used as an France). This paper was submitted 3 VII 13, accepted 13 VIII 13, estimator of L for fossil hominins, this relationship has pro- and electronically published 19 XII 13. duced values of 52, 78, 93, and 94 yr for Homo habilis, Homo

᭧ 2013 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved. 0011-3204/2013/54S8-0004$10.00. DOI: 10.1086/673725 Bocquet-Appel and Degioanni Neanderthal Demographic Estimates S203 erectus, Neanderthals, and modern humans, respectively, as and Pfifferling 1976), preindustrial humans (Henry and Blayo well as age at sexual maturity taken as one-fifth of L, that is, 1975; Jannetta and Preston 1991), or ethnographic humans 12–13, 13–14, 18–19, and 18–19 yr, respectively (Cutler 1975; (Hill and Hurtado 1996; Howell 1979).3 Sacher 1975). These estimates were incorporated into paleo- Caspari and Lee (2004, 2005a, 2005b, 2006) have detected demography 35 yr ago as parameters influencing the shape “increased longevity, expressed as the number of individuals of the death distribution of fossil hominins (Bocquet and surviving to adulthood” (Caspari and Lee 2004:10895) from Masset 1977, 1982; Bocquet-Appel 1982). Updated estimates the ratios of old (30ϩ yr) to young (15–29 yr) adults (noted of L, from biometric (estimated) data in more recent literature OY, which is written in demographic notation as ωd30/15d15 and 1 using the Hofman regression (1993), have produced 111.7 read as ωd30, the number of deaths at age 30 plus a number of and 111.2 yr for Neanderthals and modern humans, respec- years coinciding with the end of life, noted ω, over 15d15,the tively, that is, identical figures between the two metapopu- number of deaths at age 15, plus 15 yr (i.e., between 15 and lations and figures similar to those of Sacher (1975) and Cutler 29.9 yr). With samples of australopithecines, early Homo,Ne- (1975). These estimates also suggest that other important de- anderthals, and Early Upper Paleolithic (modern human), Cas- terminants of life history, such as age at menarche or duration pari and Lee obtained OY values of 0.12, 0.25, 0.39, and 2.08, of gestation, were similar between Neanderthals and modern respectively, with an abrupt change between Neanderthals and humans, which allows them to be set within a common de- anatomically modern humans. If the information provided by mographic frame. the data is true, then these ratios, with an increasing number of older individuals relative to younger ones, express a trend Paleodemographic Estimates toward an increasing average life span in the metapopulation of extinct hominins (Caspari and Lee 2004, 2005a, 2005b, Two death distributions by age, pre-Neanderthal (Homo hei- 2006). The robustness of Caspari and Lee’s demonstration re- delbergensis; Bermu´dez de Castro and Nicola´s 1997; Bocquet- sides especially in the relatively large sizes of the paleontological p Appel and Arsuaga 1999; minimum number of individuals samples collected and in the use of the age estimation technique 32, now 29; Bermu´ dez de Castro et al. 2004) and Neanderthal of Miles, which is homogeneous between groups and allows p (Trinkaus 1995; N 206 registration numbers), were obtained, human groups to be set into a common comparative frame along with life span estimates of the australopithecines, un- without which this demographic signal, based on the OY in- til modern humans (Caspari and Lee 2004). The pre- dicator alone, would not have been detected. Neanderthal distribution was obtained by a technique of age Derived OY values express another interesting point of in- 2 estimation by self-reference based on dental attrition developed formation that combined with the estimates of potential lon- by Miles (1963, 2001; Bocquet-Appel and Arsuaga 1999). The gevity L given above (111.7 and 111.2) produces the outlines Neanderthal age distribution was obtained using the usual sta- of living population pyramids. Assuming that populations are tistical techniques exploiting morphological age indicators for stable on average, the pyramids for these populations are adults (Trinkaus 1995), which are known to be biased toward regular and can be represented here by a triangular polygon a younger age (Bocquet and Masset 1982). These two sampled where the height represents the ages and the width the age paleontological distributions produce very high proportions of distribution of the living population. The technical details for young and mature adults (in the Neanderthal sample from the construction of the population pyramid from Caspari and Trinkaus 1995, 80% of adults aged 20 yr and over die before Lee’s data and an observed statistical relationship between OY the age of 40, not counting the question of children under 5 values for dead and living in sample life tables of modern yr of age, who are still significantly underrepresented in paleo- human populations (table 1; fig. 1) are given in CAϩ online demographic data). These proportions have no equivalent in supplement A. What is obtained is a highly schematic rep- the many controlled distributions of attritional death in other resentation of a population pyramid at 15ϩ yr. The pyramids primates (Rawlins and Kessler 1987; Richard 1985), apes for the two groups are represented in figure 2. They show the (Courtenay and Santow 1989; Hill et al. 2001; Teleki, Hunt, flattening effect produced by the departure of the pyramid at the point of the statistical longevity ω rather than at the 1. Brain weight E p 1.036 # cranial capacity (Isler et al. (2008); average cranial capacity 1,519 cc, n p 9 (Trinkaus and Tompkins 1990); 3. It is this observation of the systematic structural deviation between kg; controlled demographic patterns in primates, including humans, and the 4.8 ע kg; 9 females, 66.4 4.5 ע body weight P, 17 males, 77.6 kg (Ruff, Trinkaus, and Holliday 1997, supplemental patterns obtained most frequently from paleodemographic data that 4.6 ע averaged, 72 data); Hofman regression no. 4, given with no standard deviation, R p brought the results of the latter into question 40 yr ago (Bocquet and 0.896 (Hofman 1993:214). Masset 1982, 1985; Bocquet-Appel 1986; Masset 1973). The alternative 2. This technique is described as “self-referencing” because the sample position was to accept the paleodemographic distributions as true, to analyzed is its own anthropological age/indicator reference sample, unlike consider them as ancestral patterns of the demographic distributions of in other estimation techniques that require reference samples that are extant primate populations, and to consider the conflicting references of external to the sample analyzed. Moreover, the self-referencing approach controlled distributions of extant primates as expressing the tyranny of is the technique of choice in paleoanthropology (Mann 1968), where it actualist demography. This was the main thrust of the arguments given is impossible to make up anthropological age/indicator reference samples by the American Journal of Physical Anthropology to one of us in 1977 of extinct hominins. in rejecting a manuscript that discussed past orthodoxy. S204 Current Anthropology Volume 54, Supplement 8, December 2013

Table 1. Paleodemographic statistical data

Death at 15 Ϫ 29 yr p Death at 15ϩ yr p Y/(Y ϩ O)deadp Estimated OY Estimated maximum p p a p Group Y 15d15 O ωd15 15d15/ωd15 living 15L30/15L15 longevity L Australopithecinesb 316 353 .895 .505 51.5c Early Homo 166 208 .798 .539 73.5d Neanderthal 96 133 .721 .569 111a,b Early Upper Paleolithic 24 74 .324 .805 111a,b R2 p 0.940 ,N p 44 , preindustrial life tables with low life expectancy,0.01904 ע a From OY living p 0.47274 Ϫ 0.29547 # log [Y/(Y ϩ O)] dead at birth, excluding Yanomamo, stationary population (see Bocquet and Masset 1977; Bocquet-Appel 2002),F p 658.1 , df p 1, 42 ,P ! .00001 . b Number of Australopithecines to Early Upper Paleolithic, aged by Miles’s technique (Caspari and Lee 2004). c Australopithecine and early Homo (Cutler 1975, table 2, excluding Homo habilis; see Caspari and Lee 2004). d Homo erectus, average. maximum longevity L, which gives the population pyramids cluding primates, and the paleodemographic signal detected a more acute angle. by Caspari and Lee that does not fit in the frame. We have The first pyramid for the australopithecines, compared with no solution to this contradiction, but it cannot be ignored the others, resembles those known for the great apes. Partic- (see also Trinkaus 2011). It raises two issues: one is technical ularly notable is the similarity of the Neanderthal and Early and concerns the recurrent problems of paleodemographic Upper Paleolithic pyramids, which both have a base width at age estimation, while the other concerns population biology 15 yr similar to that for early Homo, estimated from pale- and the assumption of faster maturation in Neanderthals than ontological data, and a height identical to the Early Upper modern humans, which we will now discuss. Paleolithic, estimated from zoological data. If L and ω are Miles’s technique is based on a self-reference in the ana- correctly estimated, then in the Neanderthal paleontological lyzed sample. It therefore seems to escape from the a priori data, the proportion of deaths corresponding to 45 Ϫ ω of probabilities that are encysted in the anthropological ages/ the population pyramid is either missing or not recognized indicators reference sample used in other techniques, which among the skeletal remains. The similarity in the represen- predetermine the age estimates and for which there is now a tation of population pyramids for Neanderthals and the Early range of solutions currently under evaluation (Bocquet-Appel Upper Paleolithic highlights the contradiction between the and Bacro 2008; Caussinus and Courgeau 2010; Hoppa and biodemographic pattern of longevity of extant mammals, in- Vaupel 2002; Lucy, Ackroyd, and Pollard 2002; Lucy et al. 1996). Nevertheless, it may seem surprising that Miles’s tech- nique, which has been used many times in paleoanthropology since its publication in 1963 (for an overview, see Miles 2001), has never, to the best of our knowledge, been tested with a sample of skeletons of known ages, with its share of immature individuals to calibrate the standards and its share of adults to apply them to. An old idea is that tooth wear is not linear with chronological age, as implied in the technique, but de- creases asymptotically in older individuals. As long as this validation test has not been done, in particular to estimate the confidence intervals (CIs), this technique should be con- sidered conservatively, as indicated by Miles himself, as “an art, not a precise science” (Miles 2001:980). The hypothesis of significantly faster biological maturation in Neanderthals than in modern humans has been put for- ward in recent years as estimated from the growth of tooth enamel (Guatelli-Steinberg et al. 2005; Ramirez-Rozzi and Bermudez de Castro 2004; Smith et al. 2007). As there is a correlation between tooth development and other life-history traits, then there are grounds for asking whether the life span of Neanderthals, despite their large brains, could also have been shorter, as revealed by the growth of tooth enamel. If this is indeed the case, it would suggest a death distribution Figure 1. Relationship between OY living (15L30/15L15, vertical axis) ϩ at a younger age, on average, than in anatomically modern and the variable Y/(Y O) dead (15d15/ωd15, horizontal axis) in a sample of 44 preindustrial life tables representing stationary humans. But there is no consensus over this hypothesis of populations (r p 0). faster maturation for reasons to do with the dental techniques Bocquet-Appel and Degioanni Neanderthal Demographic Estimates S205

Figure 2. Schematic representation of the population pyramids for ages 15ϩ yr of living populations of australopithecines, early

Homo, Neanderthals, and Early Upper Paleolithic (modern humans) estimated from the 15L30/15L15 ratios, derived values of OY (Caspari and Lee 2004), and estimated maximum potential longevity (regression: Hofman 1993). Regarding the australopithecines, because the statistical demographic longevity ω is lower than the upper limit of the area representing the living at 30–45 yr of age, of unit density, this limit of 15L30 was set at ω. used, the sizes of samples counted in units, and the failure logical sites in Europe (van Andel, Davies, and Weninger to consider interindividual variations in Neanderthals and 2003). This link is determined—via the amount of biome modern human in comparisons (for a summary, see Guatelli- distribution of primary biomass—by the biomass of ungulate Steinberg 2009). Moreover, the hypothesis of rapid matura- populations available for hunter populations. From these rel- tion does not imply, in paleodemographic distributions, an atively long-cycle climatic variations in the isotopic stages of absence of individuals with recognized aging markers, as celestial mechanics, it gradually becomes clear that medium- seems to be the case in the data. These faster-maturing in- cycle cold fluctuations (Heinrich events 10–8 kyr [H]) are dividuals would also have these markers of aging, but they superimposed over relatively short-cycle and short-duration would simply be chronologically younger than their modern temperate-cold fluctuations (Oeshger-Dansgaar events 1.5 kyr human counterparts. Finally, ethnographic assumptions [DO]). These DO changes, which occur rapidly over time, (Trinkaus 1995, 2011), or assumptions by analogy with eth- significantly amplify the predominant climate in the isotopic nohistorical environmental crises (Bocquet-Appel and Arsu- stages. From oxygen isotope stage (OIS) 5 of the Eemian aga 1999), have been put forward to explain the apparent onward, 25 DO events occurred in Europe (Sanchez-Gon˜i et deficit of adults over 40 yr old and even over 25 yr old in al. 2008). Others would be expected to appear in earlier pe- pre-Neanderthal and Neanderthal samples. riods when Neanderthals lived but are so far unknown for To summarize, the mortality pattern of Neanderthals, as lack of data. A correlation between the frequency of Upper revealed by paleontological data, does not correspond to the Paleolithic archaeological sites and the variation in DO events promise of longevity displayed by the manifest continuum of has been interpreted in terms of demographic and cultural extant mammals, including primates. Could one of the two changes in Europe, which roughly correspond via changes in be wrong? This question, which is beyond the scope of this the ecological zones of primary and secondary ungulate bio- paper, cannot be evaded. mass (d’Errico, Sanchez-Gon˜i, and Vanhaeren 2006). How did local populations react to these significant variations Demographic Inferences from Paleoclimatic across their expansion area? Rather than a model of the ebb and Archaeological Data and flow of small local populations on the geographical mar- We know there is a connection between climatic variations gins of the metapopulation to and from attested refuge zones in isotopic stages and the frequency of Mousterian archaeo- over evolutionary time—which were, in fact, already occupied S206 Current Anthropology Volume 54, Supplement 8, December 2013 by the cores of the metapopulation—a model of regional to the demographic sizes of both populations. This area is a extinction in situ is proposed (Hublin and Roebroeks 2009). refuge zone on the evolutionary timescale. It should be re- Other consequences of large-amplitude and long-duration membered that the size of the Aurignacian population in this climatic fluctuations—in their low points of glacial temper- same area has been estimated at 795–12,980 individuals with ature, moreover amplified by H and DO events—were likely a 95% CI (Bocquet-Appel et al. 2005:1665, fig. 5). By applying to create demographic bottlenecks for the Neanderthals, who the magnitude of the relative difference estimated by Mellars avoided permafrost zones (Aiello and Wheller 2003), that and French (2011) to this Aurignacian population size, we determine stochastic genetic drifts (Bruner and Manzi 2006). obtain a local Neanderthal population of 80–1,300 individuals But it is not certain, on the other hand, that the peak inter- in the Perigordian refuge area before the time of contact. glacial temperatures (isotopic stage 5e and 7e), which were similar to current temperatures and corresponded to a plant Hypothesis of a Boserupian Neanderthal recolonization on the latitudes, also corresponded to periods Population Trap of population growth. Recolonization can occur with a very low population density. This is because for the interglacial The homogeneity of the lithic cultural remains of Neander- periods, account must be taken of the result, in terms of thals during their last 150 kyr is striking, except, apparently, ungulate biomass, of the combined effects of (i) the expansion at the point of their extinction (Bocquet-Appel and Tuffreau of habitable areas due to the withdrawal of ice masses and 2009), suggesting very low technical elasticity despite the sig- despite the rise in sea level (3–7 m above the current level), nificant pressure of the environmental hazards summarized and (ii) the decrease in the area of the steppe tundra as it above, which should have favored innovations because of the shifted toward the high latitudes up to the pole. The con- overall change in the ungulate biomass. Questions about the ditions that brought this expansion of habitable areas were cognitive efficiency of Neanderthals are thereby raised (Neu- favorable to forests (van Andel and Tedzakis 1996; Kukla et bauer and Hublin 2012). But the hypothesis of technical lim- al. 2002), where ungulate biomass was very low. The north- itations in the production system of Neanderthal hunters due ward shift of steppe tundra toward the high latitudes resulted to a demographic trap (Bocquet-Appel and Tuffreau 2009) not only in a smaller area, as mentioned above, but also in must also be put forward again but more properly defined. lower productivity as available sunlight diminished. Until all In any population, the production of innovations depends net effects relative to the distribution of the major biomes not only on its cognitive biological capacities but also on its during the Eemian have been simulated, taking into account demographic size. With the same cognitive capacity, under the conflicting effects described above, it will not be possible the simple assumption that innovations are produced at a low to equate the interglacial with a demographic expansion of frequency in any population, then the most demographically the Neanderthal metapopulation (see also Frenzel 1985; Gam- numerous population, in absolute terms, will produce the ble 1987). greatest number of innovations (Kremer 1993; Kuznets 1973; Bottleneck situations and purifying selection are both chro- Simon 1977; see also Powell, Shennan, and Thomas 2009; nologically identifiable. These correspond to the low (Bruner Shennan 2001). If the size of the Neanderthal metapopulation and Manzi 2006) and likely midpoints of climate series. Nev- remained very low in terms of carrying capacity—that is to ertheless, and underlying the anatomical traits that appeared say, the maximum number of mouths that it was possible to in some glacial episodes because of drift (Hublin 1998), even- feed per square kilometer of ungulate biomass given its pro- tually producing the classic Neanderthal morphotype de- duction system (the technical and social relationships in an scribed as “hyperarctic” (Aiello and Wheeler 2003; Holliday environment)—then the metapopulation could have main- 1997), the metapopulation had to reach a significant size in tained itself in a state of demographic equilibrium at a “critical these cold and harsh environments for the filter of purifying level of density” (Boserup 1965:33), but its potential technical selection to act on the phenotypes carrying selected genotypes, creativity would have been strongly limited in what is known in particular via Bergman and Allen’s rule. These periods, as a “Boserupian trap” in macrodemography theory. when populations were relatively more numerous, may have The technical and social characteristics of the Neanderthal matched phases of OIS 8 and 6, in which long, cold, but not production system might, in central and northern Europe, extreme periods emerge (except H and DO events, which are have operated in an open environment: high residential mo- not yet listed). bility but not to great distances (Conarda, Bolusc, and Mu¨n- Finally, from the differential distribution of well-docu- zeld 2012) and targeted to large gregarious herbivores and mented archaeological remains in the Perigordian region— resources (horses, bison, reindeer, and ibex) with a smaller between the Chaˆtelperronian 45,000–40,000 yr ago and the amount of many other larger game species typically domi- Aurignacian 40,000–35,000 yr ago, attributed respectively to nating the assemblages (Conarda, Bolusc, and Mu¨nzeld 2012; Neanderthals and to anatomically modern humans—Mellars Gamble 1999; Patou-Mathis 2000); consumption of shellfish, and French (2011) have estimated vestige quantity as 10 times birds, and turtles in the peripheral southern latitudinal zones larger for the modern human population. The authors equate of the expansion area (Finlayson et al. 2001); direct and dan- this relative difference in the amount of remains in this area gerous contact with prey animals by killing with lances rather Bocquet-Appel and Degioanni Neanderthal Demographic Estimates S207 than killing at a distance using projectiles (spears; Gamble at distance). But 150 kyr of apparent Neanderthal techno- 1999) with the aid of beaters and with no division of labor logical stability do not make the case for an escape from the by gender (or by age? Kuhn and Stiner 2006) between hunting Boserupian trap during the climate windows of OIS 5 and 7. and gathering, as observed ethnographically, that is, with both males and females working as hunters and beaters. The car- Demographic Inference from Ancient rying capacity of this hunter-gatherer production system Neanderthal DNA along with high incidental mortality (Trinkaus 1995) should be lower than in other systems in which clearly more efficient While there has been general agreement since the 1970s over hunter-gatherer hunting techniques (spears, bows) were used. the effective population size (Ne) of 10,000 individuals for In addition, the energy balance of this putative production modern humans (Hammer and Zegura 1996; Harding et al. system, combining high energy expenditure due to mobility 1997; Nei and Graur 1984; Rogers and Jorde 1995; Takahata (following herds) and energy gain from a low calorie diet 1993; Takahata, Satta, and Klein 1995; Wilson et al. 1985),

(essentially game), determines long birth intervals in modern there is no such consensus over the Neanderthal Ne. There human females and, therefore, low fertility (see Bocquet- is now enough data from Neanderthal DNA analysis to pro- Appel 2008). By analogy, it may be thought that the energy vide an outline. Analysis of Neanderthal DNA sequences be- balance/fertility reaction norm was similar in Neanderthals gan in 1997 (Krings et al. 1997) from the original specimen and that their female fertility (total fertility rate) therefore discovered in 1856, but information has since become avail- tended, on average, toward low values (!Kung: 5 children and able on mitochondrial sequences from 14 additional Nean- less) rather than high values. The relatively low fertility and derthal specimens (see Degioanni, Fabre, and Condemi 2011 high mortality of the Neanderthal hunter-gatherer production for a review), including complete mitochondrial genome se- system would have been a contributing factor in locking them quences from nine Neanderthal individuals (Briggs et al. 2009; into the Boserupian trap. Green et al. 2008, 2010). Neanderthal genomic DNA has also Although ethnographic demographic control data are lack- been sequenced since 2006 (Green et al. 2006; Noonan et al. ing on this point, there are two conceivable circumstances in 2006: 65,000 and 1,000,000 base pairs, respectively). Finally, which a population of hunters may have escaped stagnation in 2010, the genes of three Neanderthal individuals to 1.3- in a Boserupian trap: (i) the outcome of the rate of innovation, fold genomic coverage were sequenced by the Neanderthal which would sporadically increased the carrying capacity, and Genome Project (Green et al. 2010). This project mainly aims (ii) favorable environmental change (Wood 1998:112). Unlike to determine the complete sequence of Neanderthal DNA, to the realization of the rate of innovation, which is a simple answer the question of recent interbreeding between Nean- statistical function of the rate and the metapopulation size derthals and modern humans, and to provide a catalog of and cannot be located precisely in time, favorable environ- differences between the human and Neanderthal genomes. mental changes were a certainty in western Eurasia, especially The Neanderthal Genome Project has also revealed the se- over the relatively long durations of OIS 5 and 7. These fa- quence of several Neanderthal protein-coding genes that have vorable environmental changes resulted in the extension of recently evolved adaptively in modern humans (Green et al. steppe savannah and grassland areas together with the cor- 2010). Several publications on Neanderthal DNA sequences responding ungulate biomass and its carnivorous predators, focus on the time of the most recent common ancestor of which included the increasing Neanderthal metapopulation. Neanderthals and modern humans, and several publications

Along with this demographic growth, the population expan- report estimations for Neanderthal Ne. sion area was gradually homogenized, particularly through a The following is a brief digest of the demographic findings defragmentation of the geographical structure inherited from of these studies. A first study comparing three short mtDNA the previous cold period, the increase in interpopulation mi- sequences (Krings et al. 2000) suggested that Neanderthals gration, and the redensification of local populations, all of had expanded from a small population to explain the lower which favored the spread of innovations. diversity of Neanderthal mtDNA than in the great apes and With no change in hunting technique (“in the herd”: Gam- the same order of magnitude in modern humans. This result ble 1999) but simply by virtue of the increase in the overall was confirmed by analyzing five Neanderthal mtDNA ge- biomass of ungulates during OIS 5 and 7, the logistic mobility nomes (Briggs et al. 2009): the Ne was small and probably p of the Neanderthal hunter system could have diminished. The included fewer than 3,500 females (mean Ne 1,476; 268 to effect of reduced logistic mobility (with the corresponding 3,510, 95% HPD), and the authors proposed that the low decrease in energy expenditure) is to improve the energy bal- mtDNA diversity might reflect a low Neanderthal Ne over a p ance, which results in a rise in female fertility through a large part of their history. Estimating the θ Neμ parameter reduction in the birth interval (Bocquet-Appel 2008). An in- (θ is the nucleotide diversity among sequences, and μ is the crease in the metapopulation or local populations could then substitution rate per generation), assuming constant popu- occur through an opportunistic adjustment of female fertility lation size and 20 yr per generation over a short sequence of to the new carrying capacity rather than a “technological” 9 HVI mtDNA, Lalueza and colleagues (2005) propose an Ne reduction in mortality due to hunting accidents (via killing ranging from 5,000 to 9,000 individuals and confirm that this S208 Current Anthropology Volume 54, Supplement 8, December 2013 population would have been constant over time. The com- N 4r(1 Ϫ r)T e p , plete Vindija 33.16 mitochondrial genome sequence (Green ϩ ϩ Ϫ ϩ ϩ Ϫ ϩ N [rA f(1 IAfm) (1 r)A m(1 IAb) (1 r)I m rIbf] et al. 2008) showed a significantly higher ratio of nonsynon- ymous to synonymous evolutionary (d /d ) rates in Nean- N S where N is the effective size, N the population size, r the sex derthals versus modern humans. This result could be ex- e ratio; A is the adult life span; T is the generation time; I is plained by a smaller Neanderthal N and contrasts with HVI A e the standardized variance (variance/mean2) in the adult life 1 sequence results from Teshik Tash and Okladnikov indi- 2 span; Ib is the standardized variance (variance/mean )inthe viduals (Krause et al. 2007)—based on mean pairwise differ- reproductive success of sex; m indicates male and f female. ences, which suggested that Neanderthals had an Ne similar N corresponds exactly to the size of the population (Nc)of to that of modern Europeans or Asians but lower than that animals with early sexual maturity and to the size of the adult of modern Africans—and also contrasts with the results of population only (Na) in late-maturing species (see Nunney Ovchinnikov and Kholina (2010), where the dN/dS ratio in- and Elan 1994). To obtain the N census value for Neander- dicates that the Ne for their common ancestor (the human- thals, being presumably also late maturing like modern hu- Neanderthal branch) tends to be larger than in either Ne- mans, the fraction of the juvenile population (0–19.9 yr old, anderthals or modern humans. or 20L0 in demographic notation) must therefore be added to On mtDNA data again, but using a modeling approach, Na. While these parameters are readily available for ecological Fabre, Condemi, and Degioanni (2009) found that the de- studies on living animals (e.g., livestock), estimating Na for mographic scenario that best explains the variability of Ne- past populations, for which none of the parameters of the anderthal is a population with an Ne ranging from 3,000 to formula are known, is much more complicated. It is never- 25,000 individuals that will grow in size up to 50,000 yr BP theless possible to put forward known values for living hunter- and then decline slowly until extinction. A larger distribution gatherer populations and to use these values in the knowledge value for Neanderthal Ne, whose median corresponds to that although they are not “real” values for Neanderthal pop- 32,263 individuals, was proposed instead by the best scenario ulations, it is probable that the real values would be in the analyzing a possible admixture process between Neanderthals suggested range. Our aim is to obtain an order of magnitude and early European modern humans (Ghirotto et al. 2011). rather than exact values. By cross-referencing the values, sev-

Ne values from nuclear DNA are very rare but confirm that eral scenarios are obtained. To reduce their number, and since Neanderthals derived from a very small ancestral population we have no evidence to the contrary at present, we consider with an N of about 3,000 ranging up to 12,000 (Green et al. that the population consisted of equal numbers of men and e p 2006). This estimated N turns out to be similar to that pro- women (r 0.5), that the men and women had the same e p p posed for a population of modern humans: this “small pop- adult life span (Af Am Ai) and the same reproductive p p ulation” character seems to be a feature of both Neanderthal success (Ibm Ibf Ibi). Regarding the adult life span, there and modern human evolution. are only two detailed demographic studies of hunter-gatherers As we have seen, the literature offers different N values, (Hill and Hurtado 1996; Howell 1979) giving this information e ϩ but all publications agree that the Neanderthal N would be (for individuals of 20 yr, both sexes). The adult life spans e for both sexes are virtually identical, at 36.5 for !Kungs and very low. But what is the N ? N is defined as the size of an e e 37 yr for Aches. These two poulations live in the tropics, ideal population, a Wright-Fisher population (Fisher 1930; unlike the Neanderthals, who lived in temperate and, most Wright 1931) that has the same rate of change of allele fre- frequently, subarctic regions. These studies include inconsis- quencies or heterozygosity as the population under study. It tencies, which are discussed by the authors (e.g., Hill and is not easy to relate the N to the census size of a population e Hurtado 1996:258; Howell 1979:116). However, in the ab- (N ). The N is almost always smaller than the actual size N . c e c sence of any other information, these demographic data on Some authors (Belle et al. 2006; Ray 2003; Wood 1987; for a living hunter-gatherers are used in this article to estimate a review, see Hawk 2008) propose that the N is approximately yr: 33–40 yr), which are extended 3ע) e range of adult life span one-half of the census size corresponding to the reproductive to lower values for the lower class boundary to take the effect individuals of the population. Moreover, many parameters of a cool or cold climate into account as well as the demo- can affect the Ne value; this raises the question of whether Ne graphic reality possibly reflected by the very small proportion is in fact a reliable predictor of Nc. of adult Neanderthal skeletons found. The result is a relatively In the field of animal species conservation, where it is es- broad range of 25–40 yr for the hypothetical life span of sential not only to maintain a sufficient population size for Neanderthal adults, with an estimated SD (deviation/mean2) the survival of the species but also some degree of genetic in the adult life span (both sexes) of 38.88888/(25 or 30 or variability, researchers were already focusing in the 1970s on 40)2. the relationship between Ne and Nc, for which Hill (1972) Combining these values produced nine different scenarios. proposed an initial formula. This formula, modified by Nun- For each scenario, we apply the formula to estimate the Na ney (1993) and widely used in animal studies, is written value of the Neanderthal population given the extreme values Bocquet-Appel and Degioanni Neanderthal Demographic Estimates S209

of Ne estimated from genomic DNA (Ne range from 3,000 to Table 2. Values of the parameters used to calculate the

12,000). Ne/Na ratio

We used Ne values estimated from the analysis of genomic DNA for two reasons. First, because the formulas available Parameter Value p to calculate Na are more suited to this type of data, and second, r sex ratio .5 because we believe that the information derived from the T p generation time 17.5, 20, 25 p p p a Ai Af Am adult life span 25, 30, 40 entire genome is less biased than the information contained p p p p IAi IAf IAm standardized variance 38.8888/Ai .06222, in mitochondrial DNA alone (females only, very wide chro- (variance/mean2) in the adult life .04320, .02430 nological dispersion of sequences). spanofsexi p p p 2 p We also tested three differentL 0 20 proportions (juvenile Ibi Ibf Ibm standardized variance 6/2 1.5 2 population): 40%, 50%, or 60% of the total population. The (variance/mean ) in the reproductive data used for the nine scenarios are shown in table 2, and success of sex i the results are shown in figure 3. The figure clearly shows a Based on Hill and Hurtado (1996:196) and Howell (1979:88). that the Ne/Na ratio reaches its maximum when the generation time (T) is the longest and life expectancy (Ai) the shortest: mating long-term and short-term Ne. This research field will in this case the N (Na and Nc) values are the lowest, even therefore provide answers to our question within a short time. with 60% of young individuals in the population, ranging from 5,000 to 50,000. On the other hand, when the ratio N / e Concluding Remarks Na is the lowest, that is to say, when T is short and Ai is long,

Na and Nc are the highest, with a maximum of 70,000 indi- Paleodemographic data are eclectic by nature, but the effort viduals. must be made to integrate their interpretation without hiding An estimated population size of 5,000 to 70,000 individuals the difficulties and to resolve contradictions, especially re- should not be considered as an exact value but rather as an garding paleontological metapopulations such as the Nean- order of magnitude. It should be remembered, furthermore, derthals, where the mists of time become increasingly im- that the formula proposed by Nunney applies to a population penetrable with chronological depth. In Neanderthal with no subdivisions and no generation overlap. If this is not paleodemographic death distributions by age, very few adults the case, then generation overlap can reduce Ne to 25%–75% are older than 40, while the promise of potential maximum of Na (Felsenstein 1971). Reproductive variance (variation in longevity implied by the quasi-biological continuum of mam- the contribution to the next generation) between males and mals points to much more. One could even venture to assume females (Ai) implies that the Ne of portions of the genome that the Neanderthal and modern human death distributions with different inheritance patterns can be different (the higher should be similar. the reproductive variance, the lower the Ne). This means that Increasingly detailed reconstitutions of climate, layering

Ne estimated for the same population but with different mark- multiple sequences of variations that range from very long to ers (Y chromosome, mtDNA, and autosomal markers) can short periods (Sanchez Gon˜i et al. 2008), and their connec- be very different and therefore difficult to compare. Nonran- tions to the Neanderthal population through geographically dom mating, in particular assortative mating (mate chosen distributed primary and secondary biomass raise new ques- on the basis of phenotypic similarities) decreases the Ne and tions. It is as if the last 10,000 yr of the Holocene, during increases the genetic drift. In a subdivided population, non- which the modern human metapopulation will reach 9 billion random mating can have a greater effect on members of the people, were a temperate niche of stability, as the latest similar same subpopulation: the Ne of a subdivided population can niches date back to OIS 5 (Eemian: 114–130 kyr) and OIS be different (lower) compared with a randomly mating pop- 11 (core: 400–420 kyr; see also Richerson, Boyd, and Bettinger ulation of the same size. 2009). Except in these three temperate niches outside the Finally, but crucially, it is important to keep in mind that Mediterranean zone, the vegetation was mainly cold steppe the average Ne over the long term is not the “classical” arith- tundra and was regularly devastated by what would now be metic mean but rather the harmonic mean over several gen- akin to catastrophic DO and H climate events. It can be erations (Crow and Kimura 1970; Wright 1938). This means hypothesized that the demography of the Neanderthal meta- that the Ne is strongly affected by the smaller Ne values and population, living under conditions where extreme environ- will be close to the smallest Ne over several generations; that mental instability with short periods was the norm, was pri- is, bottlenecks can mask previously high Ne values. marily stagnant, with frequent bottlenecks and episodes of The question we must ask is to what period the value of decline.

Ne calculated from Ne corresponds: beginning, average value, Finally, the Neanderthals may have suffered the additional or end? The current proliferation of studies geared to the handicap of a system of specialized hunters, described as top- conservation and management of endangered or exploited level carnivores (Richards and Trinkaus 2009), even though species (Gomez-Uchida et al. 2013; Serbezov et al. 2012; plants were also consumed (Henry, Brooks, and Piperno Whiteley et al. 2012) has produced new methods for esti- 2011). Relative to a modern human, the estimated individual p p p Figure 3. Ne/Na and Na and Nc variation depending on T, Ai value, and 20L0 proportions. a, T generation time 17.5. b, T 20. c, T 25. Bocquet-Appel and Degioanni Neanderthal Demographic Estimates S211 metabolic cost of an adult Neanderthal is very high (3,500– Tjeerd H. van Andel and William Davies, eds. Pp. 147–166. Cambridge: McDonald Institute. 5,000 kcal/d vs. 2,150–2,400 kcal/d for a male modern human; Belle, Elise M. S., Uma Ramakrishnan, Joanna L. Mountain, and Guido Bar- Churchill 2007; Sorensen and Leonard 2001; Steegman, Cerny, bujani. 2006. Serial coalescent simulations suggest a weak genealogical re- and Holliday 2002). Assuming a similar distribution of nu- lationship between Etruscans and modern Tuscans. Proceedings of the Na- tional Academy of Sciences of the USA 103:8012–8017. trients between Neanderthals and modern humans, food in- Bermu´dez de Castro, Jose´ Maria, Maria Martinon-Torres, Marina Lozano, take per capita for Neanderthals would have required almost Susana Sarmiento, and Ana Muela. 2004. Paleodemography of the Ata- twice the mass of ungulate meat as that consumed by modern puerca-Sima de los Huesos hominin sample: a revision and new approaches to the paleodemography of the European Middle Pleistocene population. humans. Mechanically, given similar natural hunting condi- Journal of Anthropological Research 60(1):5–26. tions, this level of ungulate consumption implies that the Bermu´dez de Castro, Jose´ Maria, and Marıa Elena Nicola´s. 1997. Palaeode- Neanderthal population density was half that of modern hu- mography of the Atapuerca-SH Middle Pleistocene hominid sample. Journal mans. The demographic instability of this metapopulation of of Human Evolution 33(2/3):333–355. Bocquet, Jean-Pierre, and Claude Masset. 1977. Estimateurs en pale´ode´mo- specialized hunters, which was small on average, and the var- graphie. L’Homme 17:65–90. iation of its geographical area of expansion and fragmentation, ———. 1982. Farewell to paleodemography. Journal of Human Evolution 11: should help to understand why it stagnated technologically 321–333. ———. 1985. Paleodemography: resurrection or ghost? matters of moment. (Bocquet-Appel and Tuffreau 2009) and probably also socially Journal of Human Evolution 14:107–111. (see the array of Hayden 2012), as it spent most of its evo- Bocquet-Appel, Jean-Pierre. 1982. Une esquisse de quelques me´canismes de´- lutionary time moldering in the depths of a Boserupian de- mographiques au pale´olithique. In 1er Congre`s international de pale´ontologie humaine. Jean Piveteau and Henry de Lumley, eds. Pp. 971–992. Paris: mographic trap. Institute de pale´ontologie humaine. Measuring the acquisition or lack of acquisition by Ne- ———. 1986. Small populations: demography and paleoanthropological in- anderthals of a modern form of human behavior—that is, ferences. Journal of Human Evolution 14:683–691. ———. 2002. Paleoanthropological traces of Neolithic demographic transi- the ability, usually shown in modern humans, to express a tion. Current Anthropology 43:638–650. very wide range of cognitive responses to contrasting so- ———. 2008. Explaining the Neolithic demographic transition. In The Ne- cionatural situations—by simply comparing lists of archae- olithic demographic transition and its consequences. Jean-Pierre Bocquet- Appel and Ofer Bar-Yosef, eds. Pp. 35–56. Dordrecht: Springer. ological cultural items between the two groups is not an Bocquet-Appel, Jean-Pierre, and Juan Luis Arsuaga. 1999. Age distributions appropriate approach (see Shea 2011). If one of the meta- of hominid samples at Atapuerca (SH) and Krapina indicate accumulation populations (the Neanderthals) remained largely within the by catastrophe. Journal of Archaeological Science 26(3):327–338. confines of the experience of the Boserupian traps men- Bocquet-Appel, Jean-Pierre, and Jean-Noe¨l Bacro. 2008. Estimation of an age distribution with its confidence intervals using an iterative Bayesian pro- tioned above while the other (modern human), numbering cedure and a bootstrap sampling approach. In Recent advances in paleo- several million, was able to experience the African and Eur- demography: data, techniques, patterns. Jean-Pierre Bocquet-Appel, ed. Pp. asian population expansion, then comparing their lists of 63–82. New York: Springer. Bocquet-Appel, Jean-Pierre, Pierre-Yves Demars, Lorette Noiret, and Dmitry cultural items at a point of fortuitous spatiotemporal contact Dobrowsky. 2005. Estimates of Upper Palaeolithic meta-population size in (say in the Pe´rigord at 40–35 kyr) does not inform us about Europe from archaeological data. Journal of Archaeological Science 32:1656– their different biological cognitive potential. The differences 1668. Bocquet-Appel, Jean-Pierre, and Alain Tuffreau. 2009. Technological responses between these lists will mirror those of the amplitudes of of Neanderthals to macroclimatic variations (240,000–40,000 BP). Human the socionatural experiences of these contemporaneous Biology 81:287–307. metapopulations whose demographic numbers varied by a Boserup, Ester. 1965. The conditions of agricultural growth. London: Allen & factor of 100 and perhaps even more. The cause of the Unwin. Briggs, Adrian W., Jeffrey M. Good, Richard E. Green, Johannes Krause, cultural limitation of the Neanderthal metapopulation com- Tomislav Maricic, Udo Stenzel, Carles Lalueza-Fox, et al. 2009. Targeted pared with that of modern humans may well have resided retrieval and analysis of five Neandertal mtDNA genomes. Science in its small numbers alone. 325(5938):318–321. Bruner, Emiliano, and Giorgio Manzi. 2006. Saccopastore 1: the earliest Ne- anderthal? a new look at an old cranium. In Neanderthals revisited: new approaches and perspectives. Harvati Katerina and Terry Harrison, eds. Pp. 23–36. Dordrecht: Springer. Acknowledgments Caspari, Rachel, and Sang-Hee Lee. 2004. Older age becomes common late in human evolution. Proceedings of the National Academy of Sciences of the We thank the organizers and the Wenner-Gren Foundation USA 101(30):10895–10900. for taking the opportunity to participate in this very stimu- ———. 2005a. Are OY ratios invariant? a reply to Hawkes and O’Connell. lating conference, Christine Verna for her comments, and two Journal of Human Evolution 49:654–659. ———. 2005b. Taxonomy and longevity: a reply to Minichillo. Journal of insightful anonymous reviewers. The figures were produced Human Evolution 49:646–649. by Danie`le Fouchier, Nathan Bocquet-Appel, and Ste´phane ———. 2006. Is human longevity a consequence of cultural change or modern Renault. biology? American Journal of Physical Anthropology 129(4):512–517. Caussinus, Henri, and Daniel Courgeau. 2010. Estimer l’aˆge sans le mesurer. Population 65(1):117–146. Churchill, Steven E. 2007. Bioenergetic perspectives on Neandertal thermo- References Cited regulatory and activity budgets. In Neanderthals revisited: new approaches Aiello, Leslie C., and Peter Wheeler. 2003. Neanderthal thermoregulation and and perspectives. Harvati Katerina and Terry Harrison, eds. Pp. 113–133. the glacial climate. In Neanderthals and modern humans in the European Dordrecht: Springer. landscape during the last glaciation: archaeological results of the Stage 3 project. Coale, Ansley J., Paul George Demeny, and Barbara Vaughan. 1983. Regional S212 Current Anthropology Volume 54, Supplement 8, December 2013

model life tables and stable populations. Princeton, NJ: Princeton University B. Clegg. 1997. Archaic African and Asian lineages in the genetic ancestry Press. of modern humans. American Journal of Human Genetics 60:772–789. Conarda, Nicholas J., Michael Bolusc, and Susanne C. Mu¨nzeld. 2012. Middle Hawk, John. 2008. From genes to numbers: effective population sizes in hu- Paleolithic land use, spatial organization and settlement intensity in the man evolution. In Recent advances in palaeodemography. Jean-Pierre Boc- Swabian Jura, southwestern Germany. Quaternary International 247:236– quet-Appel, ed. Pp. 9–30. Dordrecht: Springer. 245. Hawkes, Kristen. 2006. Life history theory and human evolution: a chronicle Courtenay, Jackie, and Gigi Santow. 1989. Mortality of wild and captive chim- of ideas and findings. In The evolution of human life history. Kristen Hawkes panzees. Folia Primatologia 52:67–177. and Richard R. Paine, eds. Pp. 45–93. Oxford: Currey. Crow, James F., and Motoo Kimura. 1970. An introduction to population ge- Hayden, Brian. 2012. Neandertal social structure? Oxford Journal of Archae- netics theory. New York: Harper & Row. ology 31(1):1–26. Cutler, Richard G. 1975. Evolution of human longevity and the genetic com- Henry, Amanda G., Alison S. Brooks, and Dolores R. Piperno. 2011. Micro- plexity governing aging rate. Proceedings of the National Academy of Sciences fossils in calculus demonstrate consumption of plant and cooked foods in of the USA 72(11):4664–4668. Neanderthal diets (Shanidar III, Iraq; Spy I and II, Belgium). Proceedings Degioanni, Anna, Virginie Fabre, and Silvana Condemi. 2011. Ge´ne´tique et of the National Academy of Sciences of the USA 108(2):486–491. pale´oanthropologie: deux approches pour un dialogue autour des Ne´an- Henry, Louis, and Yves Blayo. 1975. La population de la France de 1740 a` dertaliens. Bulletins et Me´moires de la Socie´te´ d’anthropologie de Paris 23(1): 1860. Population 30:71–122. 1–18. Hill, Kim, Christophe Boesch, Jane Goodall, Anne Pusey, Jennifer Williams, d’Errico, Francesco, Maria Fernanda Sanchez-Gon˜i, and Marian Vanhaeren and Richard Wrangham. 2001. Mortality rates among wild chimpanzees. 2006. L’impact de la variabilite´ climatique rapide des OIS 3–2 sur le peu- Journal of Human Evolution 40:437–450. plement de l’Europe. In L’homme face au climat. Paris: Odile Jacob. Hill, Kim Ronald, and A. Magdalena Hurtado. 1996. Ache life history: the Fabre, Virginie, Silvana Condemi, and Anna Degioanni. 2009. Genetic evi- ecology and demography of a foraging people. New York: Aldine. dence of geographical groups among Neanderthals. PLoS ONE 4(4):e5151. Hill, William G. 1972. Effective size of populations with overlapping gener- Felsenstein, Joseph. 1971. Inbreeding and variance effective numbers in pop- ations. Theoretical Population Biology 3:278–289. ulations with over-lapping generations. Genetics 168:581–597. Hofman, Michel A. 1993. Encephalization and the evolution of longevity in Finlayson, Clive, Darren A. Fa, Geraldine Finlayson, Francisco Giles Pacheco, mammals. Journal of Evolutionary Biology 6:209–227. Jose´Marı´a Gutierrez-Lopez, and Antonio Santiago Perez. 2001. Use of the Holliday, Trenton W. 1997. Body proportions in Late Pleistocene Europe and landscape by humans from the Middle Palaeolithic to the Neolithic: the modern human origins. Journal of Human Evolution 32:423–447. case of the northern shore of the Strait of Gibraltar. Almoraima 25:65–71. Hoppa, Robert D., and James W. Vaupel. 2002. Paleodemography: age distri- Fisher, Ronald Aylmer. 1930. The genetic theory of natural selection. Oxford: butions from skeletal samples. Cambridge: Cambridge University Press. Oxford University Press. Howell, Nancy. 1979. Demography of the Dobe !Kung. New York: Academic Frenzel, Burkhard. 1985. Die Umwelt des Menschen im Eiszeitalter. Quarta¨r Press. Hublin, Jean Jacques. 1998. Climatic changes, paleogeography, and the evo- 35/36:7–33. lution of the Neandertals. In Neandertals and modern humans in western Gamble, Clive. 1987. Man the shoveler: alternative models for Middle Pleis- Asia. Akazawa Takeru, Kenichi Aoki, and Ofer Bar-Yosef, eds. Pp. 295–310. tocene colonization in northern latitudes. In The Pleistocene old world: re- New York: Plenum. gional perspectives. Olga Soffer, ed. Pp. 81–98. New York: Plenum. Hublin, Jean Jacques, and Wil Roebroeks. 2009. Ebb and flow or regional ———. 1999. The Palaeolithic societies of Europe. Cambridge: Cambridge Uni- extinctions? on the character of Neandertal occupation of northern envi- versity Press. ronments. Comptes rendus Palevol 8(5):503–509. Ghirotto, Silvia, Francesca Tassi, Andrea Benazzo, and Guido Barbujani. 2011. Isler, Karin, Christopher E. Kirk, Joseph M. A. Miller, Gene A. Albrecht, Bruce No evidence of Neandertal admixture in the mitochondrial genomes of R. Gelvin, and Robert D. Martin. 2008. Endocranial volumes of primate early European modern humans and contemporary Europeans. American species: scaling analyses using a comprehensive and reliable data set. Journal Journal of Physical Anthropology 146(2):242–252. of Human Evolution 55:967–978. Gomez-Uchida, Daniel, Friso P. Palstra, Thomas W. Knight, and Daniel E. Jannetta, Ann Bowman, and Samuel H. Preston. 1991. Two centuries of mor- Ruzzante. 2013. Contemporary effective population and metapopulation tality change in central Japan: the evidence from a temple death register. size (Ne and meta-Ne): comparison among three salmonids inhabiting a Population Studies 45:417–436. fragmented system and differing in gene flow and its asymmetries. Ecology Kendal, Jeremy, Jamshid J. Tehrani, and John Odling-Smee. 2011. Human and Evolution 3(3):569–580. niche construction in interdisciplinary focus. Philosophical Transaction of Goux, Jean-Michel. 1982. L’estimation du rapport de masculinite´etdesa the Royal Society B 366:785–792. variance en fonction de l’effectif. In 1er Congre`s international de pale´ontologie Krause, Johannes, Ludovic Orlando, David Serre, Bence Viola, Kay Pru¨fer, humaine. Jean Piveteau and Henry de Lumley, eds. Pp. 993–996. Paris: Michael P. Richards, Jean-Jacques Hublin, Catherine Ha¨nni, Anatoly P. Institute de pale´ontologie humaine. Derevianko, and Svante Pa¨a¨bo. 2007. Neanderthals in central Asia and Green, Richard E., Johannes Krause, Adrian W. Briggs, Tolmislav Maricic, Siberia. Nature 449(7164):902–904. Udo Stenzel, Martin Kircher, Nick Patterson, et al. 2010. A draft sequence Kremer, Michael. 1993. Population growth and technological change: one of the Neandertal genome. Science 328:710–722. million BC to 1990. Quarterly Journal of Economics 108(3):681–716. Green, Richard E., Johannes Krause, Susan E. Ptak, Adrian W. Briggs, Michael Krings, Matthias, Cristian Capelli, Frank Tschentscher, Helga Geisert, Sonja T. Ronan, Jan F. Simons, Lei Du, et al. 2006. Analysis of one million base Meyer, Arndt von Haeseler, Karl Grossschmidt, Go¨ran Possnert, Maja Pau- pairs of Neanderthal DNA. Nature 444(7117):330–336. novic, and Svante Pa¨a¨bo. 2000. A view of Neandertal genetic diversity. Green, Richard E., Anna-Sapfo Malaspinas, Johannes Krause, Adrian W. Nature Genetics 26:144–146. Briggs, Philip L. F. Johnson, Caroline Uhler, Matthias Meyer, et al. 2008. Krings, Matthias, Anne Stone, Ralf W. Schmitz, Heike Krainitzki, Mark Stone- A complete Neandertal mitochondrial genome sequence determined by king, and Svante Pa¨a¨bo. 1997. Neandertal DNA sequences and the origin high-throughput sequencing. Cell 134:416–426. of modern humans. Cell 90:19–30. Guatelli-Steinberg, Debbie. 2009. Recent studies of dental development in Kuhn, Steven L., and Mary C. Stiner. 2006. What’s a mother to do? a hy- Neandertals: implications for Neandertal life histories. Evolutionary An- pothesis about the division of labor among Neandertals and modern hu- thropology 18:9–20. mans in Eurasia. Current Anthropology 47(6):953–980. Guatelli-Steinberg, Debbie, Donald J. Reid, Thomas A. Bishop, and Clark Kukla, George J., Michael L. Bender, Jacques-Louis de Beaulieu, Gerard Bond, Spencer Larsen. 2005. Anterior tooth growth periods in Neandertals were Wallace S. Broecker, Piet Cleveringa, Joyce E. Gavin, et al. 2002. Last in- comparable to those of modern humans. Proceedings of the National Acad- terglacial climates. Quaternary Research 58(1):2–13. emy of Sciences of the USA 102(40):14197–14202. Kuznets, Simon. 1973. Modern economic growth: findings and reflections. Hammer, Michael F., and Stephen L. Zegura. 1996. The role of the Y chro- American Economic Review 63(3):247–258. mosome in human evolutionary studies. Evolutionary Anthropology 5:116– Lalueza-Fox, Carles, Maria L. Sampietro, David Caramelli, Yvonne Puder, 134. Martina Lari, Francesc Calafell, Cayetana Martinez-Maza, et al. 2005. Nean- Harding, Rosalind M., Stephanie Malia Fullerton, Robert C. Griffiths, Jac- dertal evolutionary genetics: mitochondrial DNA data from the Iberian quelyn Bond, Martin J. Cox, Julie A. Schneider, Danielle S. Moulin, and J. peninsula. Molecular Biology and Evolution 22:1077–1081. Bocquet-Appel and Degioanni Neanderthal Demographic Estimates S213

Lucy, Dave, Robert. G. Ackroyd, and A. Mark. Pollard. 2002. Non-parametric transect modulated by orbital parameters. Quaternary Science Reviews calibration for age estimation. Applied Statistics 52(2):185–196. 27(11/12):1136–1151. Lucy, Dave, Robert G. Ackroyd, A. Mark. Pollard, and Tore Solheim. 1996. Serbezov, Dimitar, Per Erik Jorde, Louis Bernatchez, Esben Moland Olsen, A Bayesian approach to adult human age estimation from dental obser- and Leif Asbjørn Vøllestad. 2012. Life history and demographic determi- vations by Johanson’s age changes. Journal of Forensic Sciences 41:189–194. nants of effective/census size ratios as exemplified by brown trout (Salmo Mann, Alan. 1968. The paleodemography of Australopithecus. PhD disserta- trutta). Evolutionary Applications 5:607–618. tion, University of California, Berkeley. Shea, John J. 2011. Neanderthal news: extinct species exhibits variability. Evo- Masset, Claude. 1973. La de´mographie des populations inhume´es: essai de lutionary Anthropology 20(5):198–200. pale´ode´mographie. L’Homme 13:95–131. Shennan, Stephen. 2001. Demography and cultural innovation: a model and Mellars, Paul, and Jennifer C. French. 2011. Tenfold population increase in its implications for the emergence of modern human culture. Cambridge western Europe at the Neandertal-to-modern human transition. Science 333: Archaeological Journal 11:5–16. 623–627. Simon, Julian. L. 1977. The economics of population growth. Princeton, NJ: Miles, A. E. W. 1963. The dentition in the assessment of individual age in Princeton University Press. skeletal material. In Dental anthropology. Don Brothwell, ed. Pp. 191–209. Smith, Tanya M., Michel Toussaint, Donald J. Reid, Anthony J. Olejniczak, Oxford: Pergamon. and Jean-Jacques Hublin. 2007. Rapid dental development in a Middle ———. 2001. The Miles method of assessing age from tooth wear revisited. Paleolithic Belgian Neanderthal. Proceedings of the National Academy of Journal of Archeological Science 28:973–982. Sciences of the USA 104(51):20220–20225. Nei, Masatoshi, and Dan Graur. 1984. Extent of protein polymorphism and Sorensen, M. V., and W. R. Leonard. 2001. Neandertal energetics and foraging the neutral mutation theory. Evolutionary Biology 27:73–118. efficiency. Journal of Human Evolution 40:483–495. Neubauer, Simon, and Jean-Jacques Hublin. 2012. The evolution of human Steegmann, A. Theodore, Jr., Frank J. Cerny, and Trenton W. Holliday. 2002. brain development. Evolutionary Biology 39(4):568–586. Neandertal cold adaptation: Physiological and energetic factors. American Noonan, James P., Graham Coop, Sridhar Kudaravalli, Doug Smith, Johannes Journal of Human Biology 14:566–583. Krause, Joe Alessi, Feng Chen, et al. 2006. Sequencing and analysis of Takahata, Naoyuki. 1993. Allelic genealogy and human evolution. Molecular Neanderthal genomic DNA. Science 314(5802):1113–1118. Biology and Evolution 10:2–22. Nunney, Leonard. 1993. The influence of mating system and overlapping Takahata, Naoyuki, Yoko Satta, and Jan Klein. 1995. Divergence time and generations on effective population size. Evolution 47(5):1329–1341. population size in the lineage leading to modem humans. Theoretical Pop- Nunney, Leonard, and Diane. R. Elam. 1994. Estimating the effective popu- ulation Biology 48:198–221. lation size of conserved populations. Conservation Biology (8):175–184. Teleki, Geza, Edward E. Hunt, and John-Henry Pfifferling. 1976. Demographic Ovchinnikov, Iogor V., and Olga I. Kholina. 2010. Genome digging: insight observations (1963–1973) on the chimpanzees of Gombe National Park, into the mitochondrial genome of Homo. PLoS ONE 5(12):e14278. Tanzania. Journal of Human Evolution 5:559–598. Patou-Mathis, Marie He´le`ne. 2000. Neanderthal subsistence behaviours in Trinkaus, Erik. 1995. Neanderthal mortality patterns. Journal of Archaeological Europe. International Journal of Osteoarchaeology 10(5):379–395. Science 22:121–144. Powell Adam, Stephen Shennan, and Mark G. Thomas. 2009. Late Pleistocene ———. 2011. Late Pleistocene adult mortality patterns and modern human demography and the appearance of modern human behavior. Science 324: establishment. Proceedings of the National Academy of Sciences of the USA 1298–1301. 108(4):1267–1271. Ramirez-Rossi, Fernando V., and Jose´Maria Bermudez de Castro. 2004. Sur- Trinkaus, Erik, and Robert L. Tompkins. 1990. The Neandertal life cycle: the prisingly rapid growth in Neanderthals. Nature 428:936–939. possibility, probability, and perceptibility of contrasts with recent humans. Rawlins, Richard G., and Matt J. Kessler. 1987. Demography of the free ranging In Primate life history and Evolution. C. Jean De Rousseau, ed. Pp. 153– Cayo Santiago Macaques (1976–1983). In The Cayo Santiago Macaques. 180. Monographs in Primatology 14. New York: Wiley-Liss. Richard G. Rawlins and Matt J. Kessler, eds. Pp. 48–72. Albany: State van Andel, Tjeerd H., William Davies, and Bernard Weninger. 2003. The University of New York Press. human presence in Europe during the Last Glacial period. 1. Human mi- Ray, Nicolas. 2003. Mode´lisation de la de´mographie des populations humaines grations and the changing climate. In Neanderthals and modern humans in pre´historiques a` l’aide de donne´es environnementales et ge´ne´tiques. PhD the European landscape during the last glaciation: archaeological results of the dissertation, Universite´ de Gene`ve. Stage 3 project. Tjeerd H. van Andel and William Davies, eds. Pp. 31–56. Richard, Alison F. 1985. Primates in nature. New York: W. H. Freeman. Cambridge: McDonald Institute for Archaeological Research. Richards, Michael P., and Eric Trinkaus. 2009. Isotopic evidence for the diets van Andel, Tjeerd H., and P. Chronis Tzedakis. 1996. Palaeolithic landscapes of European Neanderthals and early modern humans. Proceedings of the of Europe and environs: 150,000–25,000 years ago: an overview. Quaternary National Academy of Science of the USA 1064(38):16034–16039. Richerson, Peter J., Robert Boyd, and Robert L. Bettinger. 2009. Cultural Science Reviews 15:481–500. innovations and demographic changes. Human Biology 81(2):211–235. Whiteley, Andrew R., Jason A. Coombs, Mark Hudy, Zachary Robinson, Keith Rogers, Alan R., and Lynn B. Jorde. 1995. Genetic evidence on modem human H. Nislow, and Benjamin H. Letcher. 2012. Sampling strategies for esti- origins. Human Biology 67:1–36. mating brook trout effective population size. Conservation Genetics 13:625– Ruff, Christopher B., Erik Trinkaus, and Trenton W. Holliday. 1997. Body 627. mass and encephalization in Pleistocene Homo. Nature 387:173–176. Wilson, Allan C., Rebecca L. Cann, Steven M. Carr, Matthew George, Ulf B. Sacher, George A. 1959. Relation of lifespan to brain weight and body weight Gyllensten, Kathleen M. Helm-Bychowski, Russell G. Higuchi, et al. 1985. in mammals. In The lifespan of animals. Gordon Ethelbert Ward Wolsten- Mitochondrial DNA and two perspectives on evolutionary genetics. Bio- holme and Cecilia Mary O’Connor, eds. Pp. 115–133. CIBA Foundation logical Journal of the Linnean Society 26:375–400. Colloquia on Ageing, vol. 5. Boston: Little-Brown. Wood, James W. 1987. The genetic demography of the Gainj of Papua New ———. 1975. Maturation and longevity in relation to cranial capacity in Guinea. 2. Determinates of effective population size. American Naturalist hominid evolution. In Primate functional morphology and evolution. Russell 129(2):165–187. Tuttle, ed. Pp. 417–441. The Hague: Mouton. ———. 1998. A theory of pre-industrial population dynamics: demography, Sacher, George A., and Everett F. Staffeldt. 1974. Relation of gestation time economy, and well-being in Malthusian systems. Current Anthropology 39: to brain weight for placental mammals: implications for the theory of 99–135. vertebrate growth. American Naturalist 108:593–615. Wright, Sewall. 1931. Evolution in Mendelian populations. Genetics 16:97– Sa´nchez Gon˜i, Marı´a Fernanda, Amaelle Landais, William J. Fletcher, Filipa 159. Naughton, Ste´phanie Desprat, and Josette Duprat. 2008. Contrasting im- ———. 1938. Size of population and breeding structure in relation to evo- pacts of Dansgaard-Oeschger events over a western European latitudinal lution. Science 87:430–431. S214 Current Anthropology Volume 54, Supplement 8, December 2013

Agreements and Misunderstandings among Three Scientific Fields Paleogenomics, Archaeology, and Human Paleontology

by Carles Lalueza-Fox

The emergence of paleogenomics (the study and analysis of ancient genomes) has provided a new, powerful source of information that can be used to test previous hypotheses regarding human evolution. However, various mis- understandings concerning the interpretation of genetic data in an archaeological and paleontological context and the existence of different scientific goals tend to hinder the fluent and fruitful collaboration between these fields. Here we explore some of the subjects creating confusion, such as the problems associated with molecular clocks, the difference between sequence divergence and species divergence, and the limitations of the uniparental markers. Limited understanding of how the expression of a genome shapes the phenotype (including morphology and cognition) is the main obstacle to linking the genetic and the morphological evidence available. In the case of Neanderthals (and probably Denisovans, too), it is obvious that the conspicuous morphological differences cannot be explained by differences in a list of about 100 genes alone, thus suggesting that regulatory genomic elements must have been involved. A functional analysis of the genes involved as well as a study of the genomic architecture— a complexity level above the simple DNA message—could help us fill this gap. It is hoped that this future work will lead to the emergence of an interrelated and multidisciplinary view of the study of the past based on real collaborative efforts among disciplines.

Introduction or paleoclimatology) was seen as valuable and most-needed additional evidence. This kind of subjective multidisciplinary match was very convenient for increasing the chances of pub- The interaction between archaeologists, paleontologists, and lication of those population genetic papers. However, many researchers from the emerging field of paleogenomics has of these analyses were in fact purely descriptive and provided traditionally been plagued by misunderstandings and a lack very limited insights into the genetic structure of past and of collaborative efforts. Over the last three decades, molecular extant human populations. As such, their real utility would biologists working on population analysis of human samples be in providing a context that with increasingly large numbers have usually tried to fit their results to hypotheses proposed of samples could be used for testing competing explanations previously on the basis of morphological or archaeological in terms of past migrations and population affinities. When studies. These hypotheses were often chosen at random from viewed from the other point of view, the behavior of archae- the available literature by the authors of these population ologists and paleontologists has often been equally biased, genetics studies, who were clearly unfamiliar with the current sometimes ignoring widely accepted genetic results in a rather state of the art in these other fields. Furthermore, the genetic condescending attitude of intellectual isolation. Many of these results themselves—especially with data, such as mitochon- researchers seem to consider geneticists as newcomers to the drial DNA sequences, with limited phylogenetic resolving study of the human past; this may be true, but they are here power—frequently did not allow the favoring of one hy- to stay. pothesis over another. For this reason, having possible support These disagreements between different fields can be par- from another field (paleontology, archaeology, even linguistics tially explained by some limitations associated with the genetic markers typically employed as well as problems in our current understanding of the relationship between genotype and phe- Carles Lalueza-Fox is Director of the Paleogenomics Group at the Institute of Evolutionary Biology (Consejo Superior de notype. Here, I would like to highlight some of these diffi- Investigaciones Cientificas, Universitat Pompeu Fabra, Dr. Aiguader culties and suggest how they can be overcome. In the future, 88, 08003 Barcelona, Spain [[email protected]]). This paper was we can expect that an interrelated and multidisciplinary view submitted 3 VII 13, accepted 7 VIII 13, and electronically published of our study of the past will be possible, and this can only 8XI13. be achieved with direct and real collaboration.

᭧ 2013 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved. 0011-3204/2013/54S8-0005$10.00. DOI: 10.1086/673387 Lalueza-Fox Difficulties among Scientific Fields That Study the Past S215

Molecular Clocks, Sequence Divergence, and to infer from that a Paleolithic colonization of Mars.” There- Species Divergence fore, the smaller the ancestral population size, the closer se- quence divergence times and species divergence time would We will start by discussing some of the problems associated be; but we have to keep in mind that both features do not with the evolutionary interpretation of the genetic data, as need to be coincident. they are usually the subject of misinterpretations by archae- ologists and paleontologists. The molecular clock hypothesis is based in the regularity of the mutation process in neutral Mitochondrial DNA: Limitations of genetic regions along time, thus involving the possibility of Uniparental Markers using it as a time estimator for molecular evolution. There Before the mass availability of genome-wide data, people are, however, some problems with the accuracy of a molecular working on the genetics of human populations had to base clock. First, the current genetic diversity (either a population their interpretations on single genetic loci, mainly uniparental of study or a species) needs to be well characterized; second, markers such as maternally inherited mitochondrial DNA the mutation rate needs to be known; third, we need to have (mtDNA) and the paternally inherited Y chromosome. Al- precise dates to calibrate the clock (they are usually taken though it is frequently stated that mtDNA is just a single, from the fossil record); and fourth, we need to work with uniparental genetic marker, the limitations of the mtDNA for selectively neutral genomic regions. All four factors can have interpreting evolutionary processes are not fully recognized their own limitations; for instance, there is conflicting evi- in the population genetics literature. Because of the stochastic dence for estimating the mutation rates from family pedigrees factors associated with demography, some genetic markers and from evolutionary data (the former rate being usually may reflect population or species history and some may not much faster than the later). Also, because of the existence of (Balloux 2010). This is related to lineage sorting—in other ubiquitous regulatory elements and undetected selective words, the process of gene-lineage fixation along an evolu- sweeps, it is sometimes not so obvious that a particular ge- tionary process. Incomplete lineage sorting occurs when a nomic region is neutrally evolving. Thus, it is not surprising gene tree or genealogy differs from the species phylogeny, a that time estimates can always be subjected to refinements phenomenon that produces conflicting phylogenies and non- and corrections. When we say, for instance, that the origin monophyletic groups for a particular genetic marker. The of the Neanderthal mitochondrial DNA variation can be dated uniparental markers (mtDNA or Y chromosome) are greatly to about 110,000 years ago (Briggs et al. 2009), we are as- affected by these random processes. In this sense, a remarkable suming that the sampled Neanderthals are representative of discrepancy between mtDNA and nuclear DNA phylogenetic the whole Neanderthal variation. If the next mitochondrial trees has been recently described for polar bears (Miller et al. genomes to be sequenced turn out to be more variable than 2012). Of course, this variation in coalescence times along the currently available ones, the “Neanderthal Eve” will be genetic markers would not be a problem if multiple nuclear moved dramatically back in time. Alternatively, if the future genetic markers or even complete genomes could be gener- mitochondrial genomes are quite similar to the previous ones, ated, as is increasingly the case even for extinct hominin spe- the date will not be significantly altered. Therefore, another cies. obvious trait of the molecular clock date is its capacity of In this regard, the estimate divergence times for the sep- being recalibrated depending on the sampling. aration of Eurasian and African populations, generated from In the last decades, a new population genetics approach the observed mtDNA diversity, has yielded dates of !100,000 with a strong mathematical base, know as coalescence, has years ago (almost always around 60,000–80,000 years ago), also been developed. The coalescence theory allows us to go while the time depth for the African populations has never backward in time from the existing genetic variation until been older than 200,000 years ago (the time of the so-called finding its common ancestors, providing inferences on pop- mitochondrial Eve). Nevertheless, nuclear DNA divergence ulation demography and genetic divergence. Another com- times obtained from complete genomes are estimated to be mon misunderstanding with other scientists dealing with the around 600,000–800,000 years (Green et al. 2010; Reich et al. study of the past is the confusion between sequence diver- 2010). It is probably oversimplistic to directly interpret these gence and species (or population) divergence. The coalescence molecular clock dates from the point of view of a simple times obtained always predate the real species divergence sim- speciation event because demographic events such as popu- ply because there is a certain genetic variation in any group lation fluctuations could have greatly affected the mitochon- of individuals at any given time. The Italian geneticist Guido drial as opposed to the nuclear genome diversity. In fact, no Barbujani (Barbujani, Bertorelle, and Chikhi 1998:489) fa- bottleneck is even needed. If the effective population size was mously illustrated this point with the following remark: “Sup- constant at the time of the emergence of modern humans, pose that some Europeans colonize Mars next year: if they the mtDNA would still coalesce at some point in the past successfully establish a population, the common mitochon- (Weaver 2012). If this was the case, anterior demographic drial ancestor of their descendants will be Paleolithic. But it events would have been “erased” and thus would be unde- would not be wise for a population geneticist of the future tectable from the analysis of the mtDNA. The African thermal S216 Current Anthropology Volume 54, Supplement 8, December 2013 conditions, unfavorable to DNA preservation, makes it likely nomes seems to be provided from yet another introgression that no ancient DNA would ever be retrievable from speci- event that took place around 35,000–50,000 years ago by con- mens before the “mitochondrial Eve,” and thus we will need tact with some archaic African lineage now extinct (Hammer to rely more and more on the analysis of complete modern et al. 2011). This archaic population, morphologically un- African genomes. determined, would have split about 700,000 years ago from the lineage leading to the ancestors of modern humans (Ham- Gene Flow from Archaic Hominins mer et al. 2011). Signals of admixture with Neanderthals have also been detected in North African populations, probably One interesting example of a coalescent discrepancy between deriving from a back-to-Africa migration after the contact in an mtDNA and the nuclear genome has been described in the Near East (Sa´nchez-Quinto et al. 2012). Thus, the complex the Denisova hominin (named after Denisova Cave in Siberia, evolutionary events that took place in Africa during the Mid- Russia). The analysis of the complete mtDNA genome showed dle Stone Age are still being unraveled. that Neanderthals were the sister group of modern humans This is a crucial point, as some of the limitations of uni- and that the Denisova lineage diverged from those of Ne- parental markers chiefly arise when they are used to detect anderthals and modern humans about one million years ago gene flow and hybridization events. We therefore need to (Krause et al. 2010). This tree was misinterpreted to represent redefine the use of mtDNA and the Y chromosome in human the evolutionary relationships between these three species. evolution studies because they have clearly failed to detect the However, subsequent analysis of the complete Denisova nu- real evolutionary processes that took place in the out-of-Africa clear genome produced a rather different picture in which expansion of our species (see, e.g., Briggs et al. 2009; Krings Denisova was now the sister group of Neanderthals, and both et al. 1997; Serre et al. 2004). Moreover, thousands of modern lineages shared a common ancestor around 640,000 and with human genomes from a large number of populations as well modern humans around 804,000 years ago (Reich et al. 2010). as new ancient hominin genomes will be available in the near The authors suggested that the Denisovan mtDNA lineage future and will provide clearer answers to questions concern- could represent an archaic mtDNA that was introduced into ing the origin of our species than those obtained from uni- Denisovan ancestors by hybridization with some archaic hom- parental markers. The scientific time of the mtDNA and Y inin and subsequently preserved in the population by incom- chromosome as the main tool to correlate with the fossil plete lineage sorting. An additional individual from the same record is coming to an end. site showed an almost identical mtDNA (Reich et al. 2010), thus demonstrating that this discordance between mtDNA Limitations of the First Neanderthal and nuclear DNA is not restricted to a single Denisovan in- Genome Draft dividual. The authors suggested this kind of discordance is not outside the range of what could be expected within a Some people may think that not much was discovered about population. However, the subsequent Denisovan high-cov- Neanderthals themselves after release of the first genome draft. erage genome (Meyer et al. 2012) showed this individual had In fact, it is easier to understand ourselves by comparing us a remarkably low heterozygosity (only ∼26%–33% of that to Neanderthals than to understand what makes a Neander- seen in modern Eurasians), which seems to indicate a very thal a Neanderthal (or a Denisovan a Denisovan; Lalueza-Fox small population size. and Gilbert 2011). This problem is derived from the low While Denisovans could be descendants of a morpholog- genomic coverage of the first draft (Green et al. 2010). With ically unknown eastern form of hominin that inhabited large a 1.3# coverage, if a particular read has an ancestral nucle- areas of Asia while Homo neanderthalensis was mainly evolving otide in a position where modern humans have a fixed, de- in Europe, the inferences drawn from the mtDNA alone, if rived nucleotide, it is likely that this read is neither the product nuclear data were unavailable, would have been rather dif- of an unknown, chimpanzee-like contamination nor post- ferent, pointing to a recent survival of more primitive hom- mortem damage (fig. 1). However, those positions where inin forms such as Homo erectus. A genomic comparison with modern humans have a fixed ancestral variant and Neander- modern humans also found that modern Melanesians, but thals a derived one are more difficult to validate (fig. 2). In not other non-African modern humans, share about 4.5% of this case, a Neanderthal read may harbor a novel variant, but their genomic regions with Denisova (Reich et al. 2010, 2011). it could also be simply due to damage or sequencing error. If anything, the Denisova study shows that the emerging pic- Of course, the damage tends to be template specific; therefore, ture of human evolution is one in which gene flow between increasing the coverage should make it possible to track those different hominin populations (or species) was common. A genes that have been modified in the Neanderthal evolution- similar result was found previously upon analysis of the Ne- ary lineage only (Lalueza-Fox and Gilbert 2011). The analysis anderthal genome in which non-African modern humans of segregating loci in Neanderthals suffers from a similar share about 2.5% of their genomic regions with the former shortcoming. Thus, with the current low coverage, it is im- (Green et al. 2010). More recently, a genomic analysis in sub- possible to distinguish random damage (Briggs et al. 2007; Saharan populations has detected that about 2% of their ge- Hofreiter et al. 2001) and/or background contamination (see, Lalueza-Fox Difficulties among Scientific Fields That Study the Past S217 e.g., Green et al. 2006; Wall and Kim 2007) from heterozy- chimpanzee genomes (2006) failed to fulfill our expectations gosity. Indeed, this can only be achieved with genome cov- in terms of being able to understand the genetic basis of the erages of around 15–20#, something that is technically pos- conspicuous morphological (and cognitive) differences that sible but exceedingly expensive in most cases (Denisovan and exist between these two species. Indeed, those people who Neanderthal specimens from Denisova cave, by now at 30– assumed that a quick look at the genetic differences would 50# coverage, are a remarkable exception). Alternatively, tar- provide an easy answer to the evolutionary processes in- geted methods can be designed to retrieve a specific genetic volved—for instance, in key hominin adaptations such as marker several times, as was the case for the ABO blood group bipedalism or brain size and complexity—were certainly dis- and bitter taste gene from two Neanderthal specimens (Lal- appointed. The problem resides in both the difficulties in ueza-Fox et al. 2008, 2009, 2011). In any case, our under- understanding gene function and also in the complexity of standing of the genomic diversity of these extinct hominins the genome operating above the simple DNA level. To start will, it is hoped, be improved with a high-coverage genome. with, what was once called “junk DNA” was found to be functional even though these genetic regions do not code for Beyond the Genome any protein. This is partly due to the existence of many reg- ulatory elements that interact with networks of genes, thus Amino acid positions in about 80 genes have been found to shaping the final organism resulting from expression of the differ between Neanderthals and modern humans by com- genome (Carroll 2008). In other words, similar or even iden- paring both genomes with those of chimpanzees (Burbano et tical genomes could produce different phenotypes as a result al. 2010; Green et al. 2010). With an increased coverage this of differences in the regulation of gene transcription (the figure will likely increase to around 100 genes, as in the case process by which DNA makes RNA, the molecule from which of Denisova (Meyer et al. 2012). These positions correspond proteins are subsequently generated). to those where Neanderthals share the ancestral genetic var- iant with the chimpanzees but modern humans display a An Example of a Regulatory Element: fixed, derived variant. This is, of course, suggestive of func- microRNA tional differences in these genes (although it is not always the case because the resulting proteins can have a similar efficiency There are many types of genomic regulatory elements, in- albeit with some amino acid changes in the underlying genes). cluding the so-called microRNAs (miRNA). The miRNAs are This list contains genes whose exact function remains un- small, noncoding RNAs with a length of 19–25 nucleotides known in most cases, and only when malfunction is somehow in their mature form that act as posttranscriptional regulators present (usually in the form of deleterious mutations) does of gene expression by acting on the DNA transcripts. It is a particular disease emerge as a consequence. However, these estimated that miRNAs regulate more than 30% of all protein- 80 preliminary genes include several associated with metab- coding genes, building complex regulatory networks that con- olism, physiology, and cognition and some with more precise trol almost every cellular process. One set of such miRNAs roles, such as being involved in the movement of sperm, the is present only in present-day humans and is thus a good expression and development of follicle hairs in the skin, or candidate for having contributed to human-specific pheno- some olfactory receptors (Green et al. 2010). While this in- types. The discovery of one miRNA, namely miR-1304, that formation offers an exceptional opportunity to create a list differs between two closely related species such as modern of genes shaped by recent selection in modern humans and humans and Neanderthals is of special interest: modern hu- thus genes modeled by the common meaning of humankind, mans carry what seems to be a fixed substitution, whereas it would be a mistake to believe that phenotypical differences Neanderthals present the ancestral allele in a nucleotide that between Neanderthals and modern humans can be explained is located just in the seed region of miRNA-1304 and is there- by this short genetic list alone (Lalueza-Fox and Gilbert 2011). fore likely to alter the spectrum of target genes for miR-1304 Some years ago, the publications of the human (2001) and (Green et al. 2010).

Figure 1. Only read of the Neanderthal genome draft (from Vi33.26) at the MC1R gene between positions 885 and 954. This illustrates the limitations of investigating Neanderthal-specific variants and also the heterozygosity using a low-coverage draft. The Vindija read does not have the Neanderthal-specific guanine substitution at nt 919 described in Monti Lessini and Sidro´n 1252. However, it could have a nondescribed, synonymous substitution at nt 942 because A to G substitutions are not known to be associated with postmortem damage. However, only the genotyping of this position in additional Neanderthal samples or an increased coverage will allow us to confirm this new, Neanderthal-specific genetic variant. S218 Current Anthropology Volume 54, Supplement 8, December 2013

The genomic search for target genes for this ancestral miR- which would be compatible with a beneficial role for the new 1304 has shown an increase of more than 15 times the number derived miR-1304 allele. Because of the relatively recent di- of putative targets (N p 515 ) for the human miRNA, thus vergence dates between Neanderthal and modern human ge- indicating an important functional evolution for miR-1304. nomes (around 800,000 years), it is perhaps unrealistic to The 36 predicted targets for Neanderthal miR-1304 include expect to find many fixed differences between both human two important genes for teeth formation, namely enamelin groups, and even functionally important differences could be and amelotin (Lopez-Valenzuela et al. 2012), and miRNA expected to segregate to some extent in both lineages. It could overexpression experiments using a luciferase-based assay be that the conspicuous phenotypic differences among ancient confirmed that the ancestral version of miR-1304 greatly re- human lineages are due to the summatory effect of a particular duces enamelin- and amelotin-associated reporter gene ex- combination of genetic variants even if some of them seg- pression by 50% (Lopez-Valenzuela et al. 2012). Interestingly, regate at low frequencies. If anything, the miRNA analysis other genes in the Neanderthal miR-1304 list include cog- again shows the complexity involved in unraveling the human nitive genes such as TCF4 (associated with neuropsychiatric evolutionary process. disorders such as schizophrenia and impaired verbal learning) Further functional studies could help our understanding or CD24 (associated with multiple sclerosis). of the link between regulation of the expression of genes Although it is difficult to determine how this down reg- associated with enamel formation and the final teeth mor- ulation would affect the individual phenotype, it is known phology. In any case, this is a nice example of what can be that the volume of coronal dentine in Neanderthal molars is expected in the future in terms of a view of the genomic larger than in modern humans. Because the absolute volume architecture that goes beyond the simple reading of a DNA is similar for both hominin groups, this results in significantly message. thinner cuspal enamel in Neanderthals than in recent humans (Macchiarelli et al. 2006). Thus, although the ameloblast se- Convergent Evolution cretion rates are similar, the enamel cusp forms faster in Ne- anderthals than in modern humans (Smith et al. 2010). An- Another problem associated with our current lack of knowl- other difference is found in the ameloblastic activity as edge regarding the link between genotype and phenotype is reflected in the periodicity of long-period lines in the enamel the analysis of possible convergent evolutionary traits in hom- (Retzius lines or perikymata; Aiello and Dean 1990). Several inin species. This is related to what is known as the evolution studies on dental growth have shown that the ontogeny in of “evolvability,” that is, the limited physical and even chem- most Neanderthal dentitions examined was more rapid than ical possibilities of a body design to create restrictions on the that of Homo sapiens individuals, either recent or fossil (Smith potential evolution of particular lineages. In the case of hom- et al. 2010). As a result, current dental eruption tables sys- inins, this could mean that only one set of adaptive traits can tematically overestimate the age of Neanderthal individuals emerge with time, although it could also mean that similar at death while accurately predicting those of fossil H. sapiens traits are likely to appear independently in different hominin (Smith et al. 2010). lineages. However, the data generated from the 1,000 Genomes Proj- One interesting example of this was described in the MC1R ect (released in 2011) have shown that the derived miR-1304, gene from Neanderthals (Lalueza-Fox et al. 2007). This gene which was previously thought to be fixed in modern humans, codes for a protein in the membrane of melanocytes that is not; intriguingly, about 5%–7% of Asian individuals share regulates the synthesis of two different pigments in the hair the ancestral miR-1304 version with Neanderthals (Lopez- and skin: the dark, brownish eumelanine and the fair, reddish Valenzuela et al. 2012). This distribution schema fits the pheomelanine. A Neanderthal-specific variant was found to model of genetic introgression from archaic to modern hu- produce a loss of function in the MC1R protein, thus resulting mans as proposed in a recent study of certain alleles of HLA in fair skin and red hair in those Neanderthals carrying this genes (Abi-Rached et al. 2011), although it could also be the variant. As in modern humans, it is likely that being hetero- result of selective sweeps within recent human populations, zygous or homozygous for this particular mutation would

Figure 2. Single read of Vi33.25 at position 735 of the RPTN gene (chr1 position 150393996) showing the ancestral (e.g., shared with the chimpanzee) variant C instead of the T fixed in modern humans. Even in light of the problems associated with low coverage, we can be reasonably certain about reads showing ancestral status because no chimpanzee contamination can be expected. Thus, and maybe a bit paradoxically, the low-coverage Neanderthal genome draft is more useful for determining modern human- specific changes than Neanderthal-specific changes. Lalueza-Fox Difficulties among Scientific Fields That Study the Past S219 produce phenotypes ranging from blond-reddish to “flame” pline through interdisciplinary trainings and meetings will red hair (Lalueza-Fox et al. 2007). However, it is worth em- allow different hypotheses to be tested from the available phasizing that the Neanderthal mutation is not found in pres- evidence. More and more, all the disciplines studying the past ent modern humans; therefore, living red-haired people will contribute from their own fields to the building of robust would have a similar phenotype but for different genetic rea- paradigms, and the current misunderstandings will, it is to sons (e.g., different mutations in the same gene). While this be hoped, fade away. is somehow an anecdotic phenotypic trait that is marginally associated to adaptation to high latitudes, it is possible that other traits related to morphology and cognition could also Acknowledgments be subjected to similar convergent processes. For instance, the increased cranial capacity and brain organization that lead to I am grateful to colleagues and to two anonymous reviewers complex human cognitive functions could, to some extent, who provided constructive comments on the manuscript for have evolved in parallel along different hominin lineages. This this article. My research is supported by a grant (BFU2012- could explain the existence of aspects of modern symbolic 34157) from the Ministerio de Economı´a y Competitividad behavior in Neanderthals well before the arrival of modern of Spain. humans in Europe (Peresani et al. 2011; Zilha˜o et al. 2010). It is likely that convergent evolution runs along particular References Cited gene networks that allow recurrent genetic modifications and Abi-Rached, Laurent, Matthew J. Jobin, Subhash Kulkarni, Alasdair Mc- that thus trigger the repeated opportunity for natural selection Whinnie, Klara Dalva, Loren Gragert, Farbod Babrzadeh, et al. 2011. The in the traits involved. This could explain underlying similar- shaping of modern human immune systems by multiregional admixture with archaic humans. Science 334(6052):89–94. ities in the phenotypical traits present in different hominin Aiello, Leslie C., and Christopher M. Dean. 1990. The microanatomy and lineages. These traits, known as “orthologous phenotypes” or development of teeth. In An introduction to human evolutionary anatomy. “phenologs,” are defined as phenotypes related by the or- Pp. 106–132. London: Academic Press. Balloux, Franc¸ois. 2010. Mitochondrial phylogeography: the worm in the fruit thology of the associated genes in two different species of the mitochondrial DNA tree. Heredity 104:419–420. (McGary et al. 2010). Although MC1R is an example of con- Barbujani, Guido, Giorgio Bertorelle, and Loune`s Chikhi. 1998. Evidence for vergent evolution at a protein-function level, it is likely that Paleolithic and Neolithic gene flow in Europe. American Journal of Human Genetics 62(2):488–491. this phenomenon could be more prevalent in regulatory cir- Briggs, Adrian W., Jeffrey M. Good, Richard E. Green, Johannes Krause, cuits and could affect genes that tend to cluster together dur- Tomislav Maricic, Udo Stenzel, Carles Lalueza-Fox, et al. 2009. Targeted ing the evolutionary process. The importance of convergent retrieval and analysis of five Neandertal mtDNA genomes. Science 325:318– 321. evolution in ancient hominins could be further explored when Briggs, Adrian W., Udo Stenzel, Philip L. F. Johnson, Richard E. Green, Janet more is known about the precise genomic basis of specific Kelso, Kay Prufer, Matthias Meyer, et al. 2007. Patterns of damage in ge- human traits that can be observed in the fossil record. nomic DNA sequences from a Neandertal. Proceedings of the National Acad- emy of Sciences of the USA 104(37):14616–14621. Burbano, Herna´n A., Emily Hodges, Richard E. Green, Adrian W. Briggs, Johannes Krause, Matthias Meyer, Jeffrey M. Good, et al. 2010. Targeted Future Directions investigation of the Neandertal Genome by array-based sequence capture. Science 328:723–725. The genetic basis of many of the traits observed in the fossil Carroll, Sean B. 2008. Evo-devo and an expanding evolutionary synthesis: a record is still unknown. This is partly due to the complexity genetic theory of morphological evolution. Cell 134(1):25–36. of these traits but also to the problems associated with work- Green, Richard E., Johannes Krause, Adrian W. Briggs, Tomislav Maricic, Udo Stenzel, Martin Kircher, Nick Patterson, et al. 2010. A draft sequence of ing with functional genomics, the branch that studies the the Neandertal genome. Science 328:710–722. biological function of the genes and their associated proteins. Green, Richard E., Johannes Krause, Susan E. Ptak, Adrian W. Briggs, Michael The study of most of the phenotypic traits will require the T. Ronan, Jan F. Simons, Lei Du, et al. 2006. Analysis of one million base pairs of Neanderthal DNA. Nature 444:330–336. use of animal models (for instance the creation of transgenic Hammer, Michael F., August E. Woerner, Fernando L. Mendez, Joseph C. mice) and molecular techniques not used by paleogeneticists. Watkins, and Jeffrey D. Wall. 2011. Genetic evidence for archaic admixture An even higher level of multidisciplinary effort will therefore in Africa. Proceedings of the National Academy of Sciences of the USA 108(37): 15123–15128. be needed in the future. Hofreiter, Michael, Viviane Jaenicke, David Serre, Arndt von Haeseler, and Once this information becomes available, it will be possible Svante Pa¨a¨bo. 2001. DNA sequences from multiple amplifications reveal to check the genomic regions involved directly in the phe- artifacts induced by cytosine deamination in ancient DNA. Nucleic Acids Research 29:4793–4799. notypic expression of extinct hominin genomes (specially Krause, J., Q. Fu, J. M. Good, B. Viola, M. V. Shunkov, A. P. Derevianko, and those phenotypic traits that could be traced in the fossil rec- S. Pa¨a¨bo. 2010. The complete mitochondrial DNA genome of an unknown ord), and thus we will be able to finally understand some of hominin from southern Siberia. Nature 464(7290):894–897. Krings, Matthias, Anne Stone, Ralf W. Schmitz, Heike Krainitzki, Mark Stone- the key issues of human evolution. king, and Svante Pa¨a¨bo. 1997. Neandertal DNA sequences and the origin However, this complex enterprise can only be achieved with of modern humans. Cell 90:19–30. multidisciplinary teams and real collaborations among ge- Lalueza-Fox, Carles, Elena Gigli, Marco de la Rasilla, Javier Fortea, and An- tonio Rosas. 2009. Bitter-taste perception in Neandertals through the anal- neticists, archaeologists, and paleontologists. A profound un- ysis of TAS2R38 gene. Biology Letters 5:809–811. derstanding of the limitations and advantages of each disci- Lalueza-Fox, Carles, Elena Gigli, Marco de la Rasilla, Javier Fortea, Antonio S220 Current Anthropology Volume 54, Supplement 8, December 2013

Rosas, Jaume Bertranpetit, and Johannes Krause. 2008. Neandertal paleo- Tagliacozzo. 2011. Late Neandertals and the intentional removal of feathers genomics in the ABO blood group gene. BMC Evolutionary Biology 8:342. as evidenced from bird bone taphonomy at Fumane Cave 44 ky B.P., Italy. Lalueza-Fox, Carles, and M. Thomas P. Gilbert. 2011. Paleogenomics of archaic Proceedings of the National Academy of Sciences of the USA 108(10):3888– hominins. Current Biology 21(24):R1002–R1009. 3893. Lalueza-Fox, Carles, Holger Ro¨mpler, David Caramelli, Claudia Sta¨ubert, Giu- Reich, David, Richard E. Green, Martin Kircher, Johannes Krause, Nick Pat- lio Catalano, David Hughes, Nadine Rohland, et al. 2007. A melanocortin terson, Eric Y. Durand, Bence Viola, et al. 2010. Genetic history of an 1 receptor allele suggests varying pigmentation among Neanderthals. Science archaic hominin group from Denisova Cave in Siberia. Nature 468(7327): 318:1453–1455. 1053–1060. Lalueza-Fox, Carles, Antonio Rosas, Almudena Estalrrich, Elena Gigli, Paula Reich, David, Nick Patterson, Martin Kircher, Frederick Delfin, Madhusudan F. Campos, Antonio Garcı´a-Tabernero, Samuel Garcı´a-Vargas, et al. 2011. R. Nandineni, Irina Pugach, Albert Min-Shan Ko, et al. 2011. Denisova Genetic evidence for patrilocal mating behaviour among Neandertal groups. admixture and the first modern human dispersals into southeast Asia and Proceedings of the National Academy of Sciences of the USA 108(1):250–253. Oceania. American Journal of Human Genetics 89(4):516–528. Lopez-Valenzuela, Marı´a, Oscar Ramı´rez, Antonio Rosas, Samuel Garcı´a-Var- Sa´nchez-Quinto, Federico, Laura R. Botigue´, Sergi Civit, Conchita Arenas, gas, Marco de la Rasilla, Carles Lalueza-Fox, and Yolanda Espinosa-Parrilla. Marı´a Carmen Avila-Arcos, Carlos D. Bustamante, David Comas, and Carles 2012. An ancestral allele of miR-1304 present in Neanderthals regulates Lalueza-Fox. 2012. North African populations carry the signature of ad- genes involved in enamel formation and could explain dental differences mixture with Neandertals. PLoS ONE 7(10):e47765. with modern humans. Molecular Biology and Evolution 29(7):1797–1806. Serre, David, Andre´ Langaney, Mario Chech, Maria Teschler-Nicola, Maja Macchiarelli, Roberto, Luca Bondioli, Andre´ Debe´nath, Arnaud Mazurier, Paunovic, Philippe Mennecier, Michael Hofreiter, Go¨ran Possnert, and Jean-Franc¸ois Tournepiche, Wendy Birch, and M. Christopher Dean. 2006. Svante Pa¨a¨bo. 2004. No evidence of Neandertal mtDNA contribution to How Neanderthal molar teeth grew. Nature 444:748–751. early modern humans. PLoS Biology 2(3):1–5. McGary, Kriston L., Tae Joo Park, John O. Woods, Hye Ji Cha, John B. Wallingford, and Edward M. Marcotte. 2010. Systematic discovery of non- Smith, Tanya M., Paul Tafforeau, Donald J. Reid, Joane Pouech, Vincent obvious human disease models through orthologous phenotypes. Proceed- Lazzari, John P. Zermeno, Debbie Guatelli-Steinberg, et al. 2010. Dental ings of the National Academy of Sciences of the USA 107(14):6544–6549. evidence for ontogenetic differences between modern humans and Nean- Meyer, Matthias, Martin Kircher, Marie-Theres Gansauge, Heng Li, Fernando derthals. Proceedings of the National Academy of Sciences of the USA 107: Racimo, Swapan Mallick, Joshua G. Schraiber, et al. 2012. A high-coverage 20923–20928. genome sequence from an archaic Denisovan individual. Science 338:222– Wall, Jeffrey D., and Sung K. Kim. 2007. Inconsistencies in the Neanderthal 226. genomic DNA sequences. PLoS Genetics 3(10):e175. Miller, Webb, Stephan C. Schuster, Andreanna J. Welch, Aakrosh Ratan, Oscar Weaver, Timothy D. 2012. Did a discrete event 200,000–100,000 years ago C. Bedoya-Reina, Fangqing Zhao, Hie Lim Kim, et al. 2012. Polar and produce modern humans? Journal of Human Evolution 63:121–126. brown bear genomes reveal ancient admixture and demographic footprints Zilha˜o, Joa˜o, Diego E. Angelucci, Ernestina Badal-Garcı´a, Francesco d’Errico, of past climate change. Proceedings of the National Academy of Sciences of Flore´al Daniel, Laure Dayet, Katerina Douka, et al. 2010. Symbolic use of the USA 109(36):E2382–E2390. marine shells and mineral pigments by Iberian Neandertals. Proceedings of Peresani, Marco, Ivana Fiore, Monica Gala, Matteo Romandini, and Antonio the National Academy of Sciences of the USA 107(3):1023–1028. Current Anthropology Volume 54, Supplement 8, December 2013 S221

Hominin Evolution in the Middle-Late Pleistocene Fossils, Adaptive Scenarios, and Alternatives

by Osbjorn M. Pearson

Hominins from Europe and Africa shed light on functional adaptations and other aspects of lifeways during the Middle Paleolithic. By the end of that time span, Neanderthals and modern humans clearly differed physically and perhaps behaviorally. Explanations of the anatomical differences have largely focused on adaptation (directional selection) to climate and habitual activity, but it is hard to rule out the alternative of genetic drift. Drift would have accelerated during periods of low population numbers, while selection operates best when populations are large and expanding. Demographic changes almost certainly tracked climatic conditions in both continents. Environmental and genetic data suggest that European hominins were primarily shaped by drift, while both factors operated in Africa.

The period of time between 250 and 35 ka witnessed the 2007, 2011). Both populations diverged from a common an- emergence of Neanderthals in western Eurasia, modern hu- cestor around 350,000 years ago as gauged by both genetic mans in Africa, and, at around 60 ka, the spread of modern differences (Green et al. 2010) and divergence in cranial di- humans into Eurasia, where they replaced archaic humans, mensions modeled as the result of neutral evolution (Weaver, albeit with a small amount of interbreeding. Interpretations Roseman, and Stringer 2008).1 There is less agreement about of these events have tended to focus on different anatomical the deeper phylogeny of these lineages and related forms from and cultural adaptations as the key underlying forces respon- the late Lower through early Middle Pleistocene, but that sible for producing the differences between modern humans period predates the central focus of this paper. Key anatomical and Neanderthals. The alternative, that genetic drift drove differences between Neanderthals and modern humans in- some or perhaps many of the anatomical changes, has long clude both the differential retention of primitive features in been recognized (Howell 1957) but has received less emphasis. each lineage as well as new features (apomorphies) in each. The time is ripe for a reconsideration of scenarios for adap- In Europe, the Neanderthal lineage evolved a series of apo- tive change because of the accumulation of a critical mass of morphies, including midfacial prognathism, a posterior po- new evidence from paleoecology, genetics, anatomy, and chro- sition of the mental foramen, a retromolar gap in the man- nology. Paleoclimatic records provide insights into why at dible, a broad suprainiac fossa that is oval in form, a large least some of the morphological and genetic evolution may juxtamastoid process coupled with a small mastoid process, have occurred. In this paper, I argue that climate and pop- an occipital bun, double-arched browridges that are reduced ulation genetics are linked. Climate affects ecological pro- in absolute volume and vertical thickness compared with ductivity and biomass, which in turn affects human popu- those of Middle Pleistocene hominins, and a substantially lation numbers. Changes in population size have predictable larger brain than those of most Middle Pleistocene hominins consequences for the expected rate of neutral genetic change. The general outline of the evolution of modern humans 1. Recent papers have produced a range of estimates for when the and Neanderthals is well known (Arsuaga 2010; Arsuaga et ancestors of Neanderthals and modern humans split, ranging from ∼835 al. 1997; Hublin 2009; Martı´non-Torres et al. 2012; Stringer ka for the average divergence for autosomal sequences (Green et al. 2010) to estimates of 270–400 ka for the population (rather than DNA se- quence) divergence time (Green et al. 2010) based on the same data. Other authors have calculated additional estimates for the divergence Osbjorn M. Pearson is Associate Professor in the Department of times between Neanderthals, Denisovans, and modern humans (e.g., Har- Anthropology at the University of New Mexico (MSC 01-1040, ris and Nielsen 2013; Li, Mulliken, and Reich 2010; Meyer et al. 2012; Albuquerque, New Mexico 87131, U.S.A. [[email protected]]). This Reich et al. 2010). It is important to note that estimates of DNA diver- paper was submitted 3 VII 13, accepted 20 VIII 13, and electronically gence dates generally precede (often substantially) estimates of population published 22 XI 13. divergence.

᭧ 2013 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved. 0011-3204/2013/54S8-0006$10.00. DOI: 10.1086/673503 S222 Current Anthropology Volume 54, Supplement 8, December 2013

(Hublin 2009; Stringer 2007). Hublin (2009) considers that tures are Omo I, dated to 195 ka (Brown, McDougall, and the full set of Neanderthal features were present by oxygen Fleagle 2012; Day and Stringer 1991; McDougall, Brown, and isotope stage (OIS) 7,2 and 3-D morphometric analyses of the Fleagle 2005), and the Herto crania at 150–160 ka (Clark et face, temporal bone, and posterior cranial vault corroborate al. 2003; White et al. 2003). this view (Harvati, Hublin, and Gunz 2010). This suite of Advances in imaging, especially synchrotron x-rays, which Neanderthal features had become common in European hom- allow researchers to peer inside teeth and count daily incre- inins by OIS 5, including the specimens from Krapina and ments of enamel accretion (Smith and Tafforeau 2008), have Saccopatore, and they became even more frequent in OIS 4– revealed that Neanderthal children matured more rapidly than 3. This gradual increase in similarity to late Neanderthals has modern children (Smith et al. 2010). Neanderthals thus bear been dubbed the “accretion model” and may have unfolded a closer resemblance to the ancestral condition of even more as a single, long trend or perhaps in two pulses (Hublin 2009). rapid dental skeletal maturation of Homo erectus (Dean and A similar pattern applies to the evolution of Neanderthal Smith 2009; Dean et al. 2001; Graves et al. 2010). In contrast, postcranial morphology (Trinkaus 1983, 2006). Scenarios for the mandibular dentition of Jebel Irhoud 3, a juvenile late the evolution of Neanderthal postcranial morphology have archaic hominin from Morocco dating to 160 ka with affinities tended to emphasize adaptation to either the need to produce to modern humans (Hublin 2001; Hublin and Tillier 1981), large amounts of physical force or to preserve heat in a cold preserves evidence of a slower, modern pace of dental de- climate. velopment (Smith et al. 2007). The problem of whether Sima de los Huesos is young (ca. Both African and European Middle Pleistocene hominins 350 ka) or old (500–600 ka) complicates scenarios for the tended to be medium to tall in stature (Carretero et al. 2012) pace of evolutionary change in Europe (Stringer 2012). The and very heavy for height relative to modern hunters and Sima de los Huesos sample shows mosaics of Neanderthal gatherers (Churchill et al. 2012; Kappelman 1996). A more and non-Neanderthal morphology in virtually all aspects of slender physique typified Omo I from Africa (Pearson et al. its morphology (Arsuaga et al. 1997). Recently Martino´n- 2008) and the early modern humans from Skhul and Qafzeh Torres and colleagues (2012) have shown that the sample has in Israel (Carretero et al. 2012; Ruff, Trinkaus, and Holliday very Neanderthal-like teeth; some of the nonmetrical features 1997). Neanderthals remained at least 20% heavier relative to are even more common in the Sima de los Huesos sample modern human foragers of similar height (Kappelman 1996; than in late, “classic” Neanderthals from OIS 4 to OIS 3, Ruff, Trinkaus, and Holliday 1997). Carretero et al. (2012) which casts doubt on simple models of a steady increase in proposed that this reduction in body mass may have been an Neanderthal features over time. Recent alternative hypotheses evolutionary adaptation to a lifestyle that favored energy ef- that accept the greater antiquity for Sima de los Huesos have ficiency. The stature of early modern humans from the proposed the presence of two lineages in Europe until 300– Levalloiso-Mousterian of the Levant and the Gravettian of 400 ka (Garcı´a and Arsuaga 2011) or complicated scenarios Europe is particularly striking relative to Neanderthals and of local extinction, recolonization, and admixture of two or almost all other samples from Europe before the twentieth more populations (Dennell, Martino´n-Torres, and Bermu´dez century (Carretero et al. 2012). de Castro 2011). While Neanderthals evolved in Europe, hominins in Africa Adaptation or Drift? evolved gradually toward a modern form (Bra¨uer 2008; Pear- son 2008; Rightmire 2009). Modern human apomorphies in- clude a larger brain than generally observed in African crania Over the last 50 years, the dominant view of the differences dating to 300 ka or before; a more globular cranium (Lie- between Neanderthals and modern humans has been that the berman 2011; Lieberman, McBratney, and Krovitz 2002) with dissimilarities in anatomy reflected adaptive differences more bossed parietals and an enlarged temporal lobe (Lie- shaped by natural selection to meet specific challenges. Coon berman 2011); an altered trajectory of endocranial growth (1962), for example, argued that the enormous nose of Ne- relative to Neanderthals (Gunz et al. 2012); a vertically short anderthals had evolved to warm glacial air. Trinkaus’s (1983) face tucked beneath the frontal lobe (Lieberman 2011); re- influential analysis of the Shanidar Neanderthals emphasized tention of a canine fossa into adulthood; and the presence of that Neanderthal morphologies met adaptive needs for greater a projecting chin on the mandible (Stringer 2002, 2007). The strength or leverage relative to modern humans. Trinkaus earliest crania that demonstrate the full suite of modern fea- argued that many distinctive facial features of Neanderthals and their relatively large canines and incisors were adaptations 2. Both ice cores from Greenland and Antarctica as well as deep-sea for increased amounts of anterior biting. Some of the adaptive cores (and other data) help to reconstruct long-term patterns of climate hypotheses have not received experimental support. For ex- change. Many authorities prefer the term “marine isotope stage” for this ample, building on previous observations by Hylander (1977) sequence because the marine sequence is the longest and most complete, but in light of the importance of ice cores in illuminating the last 300 about Neanderthal and Inuit noses, Rae, Koppe, and Stringer kyr, I have used the older and more inclusive oxygen isotope stage (OIS) (2011) found no evidence that the Neanderthal face is cold throughout this paper. adapted. Similarly, Clement, Hillson, and Aiello (2012) found Pearson Hominin Evolution S223 no, or at best ambiguous, support for the hypothesis that are exciting and motivate one to take a closer look at some Neanderthal faces were specially shaped to resist anterior den- of the recent genetic advances. tal loading. The evidence that Neanderthal bodies were adapted to a Genetics cold climate lies in their wide hips, shortened distal limb segments, short limbs relative to trunk length, and large ar- Views of the origin of modern humans and our divergence ticular surfaces and thick long bone shafts, all of which char- from Neanderthals have been profoundly and perhaps deci- acterize recent humans whose ancestors have lived in cold sively influenced by genetic data from living humans as well climates for thousands of years (Holliday 1997; Pearson 2000; as ancient DNA (aDNA) from Neanderthals. The completion Ruff 1994). Wide hips and robust long bones were already of a draft of the Neanderthal nuclear genome (Green et al. present in the Sima de los Huesos sample (Arsuaga et al. 2010) and recovery and analysis of nuclear and mitochondrial 1999; Bonmatı´et al. 2010) and may have been the primitive DNA from “Denisovans,” a third lineage that separated from condition for Middle Pleistocene Homo (Arsuaga et al. 1999). modern humans slightly before Neanderthals (Meyer et al. Wide hips may have also been inherited from Homo erectus 2012; Reich et al. 2010), stand out as signal achievements. (Simpson et al. 2008) rather than appearing as an evolutionary These discoveries have decisively answered the question of novelty in Middle Pleistocene hominins. whether interbreeding occurred between modern and archaic Recently, Betti, von Cramon-Taubadel, and Lycett (2012) humans (it did in both cases) and opened new windows on demonstrated that variance within pelvic dimensions of living which genes may have been involved in producing evolu- humans tracked population history (distance from Africa) tionary novelties in both modern humans and Neanderthals. rather than climate while variance in the dimensions of the Analysis of autosomal DNA indicates a divergence time femur and tibia correlated with minimum annual temperature between modern human and Neanderthal populations of rather than population history. The implications of these find- 270–440 ka (Reich et al. 2010). Work on aDNA has also shed ings are that contrary to previous conclusions (Ruff 1994), more light on Neanderthal population history, suggesting a pelvic form appeared to have followed a pattern of largely marked bottleneck among their ancestors sometime before the time of the Mezmaiskaya neonate, 60–70 ka (Reich et al. neutral evolution like most human cranial dimensions (Betti 2010), and another bottleneck after 48 ka (Dale´n et al. 2012). et al. 2009, 2010; Roseman 2004; Weaver, Roseman, and Furthermore, all Neanderthal mtDNAs share a common an- Stringer 2007, 2008). Given congruent estimates for stature cestor approximately 100 ka and a common ancestor with based on femurs and tibiae, the Sima de los Huesos sample modern humans ∼500 ka (Reich et al. 2010). appears to have less shortened distal limb segments than Ne- anderthals (Carretero et al. 2012), which provides some evi- dence that European hominins evolved more cold adapted Modes of Genetic Evolution proportions over time. In a review of 75 distinctive cranial, At the genetic level, two of the fundamental means by which dental, and postcranial features of early modern humans and evolution can occur are natural selection (referred to subse- Neanderthals, Trinkaus (2006) concluded that only one quar- quently simply as “selection”) and genetic drift. The two pro- ter were unique to Neanderthals while twice that many were cesses are not mutually exclusive, and both often act on a unique to modern humans, a finding that means that Ne- population at the same time. Selection generally works on a anderthal morphology had remained fairly primitive while given gene only if different alleles exist and one confers higher early moderns were much more derived. This could provide fitness than another, although epistasis (the interdependence evidence that early modern humans had shifted to different of genes to produce a phenotype) may produce a shifting niches than archaic humans and had experienced a substantial target for selection. The ultimate source of new alleles is mu- pulse of selection that tailored them for their new habits. tation, which occurs rarely. Most mutations are either neutral Genetic drift provides an alternative explanation for mor- (and have no effect on natural selection) or harmful (by in- phological divergence (Howell 1957). Although this hypoth- terfering with gene function and thus causing deleterious ef- esis has been marginalized historically, recent reviews have fects to the organism); only a small number of mutations emphasized its potential importance (e.g., Hublin 2009). Wea- prove to be beneficial. Most selection pressures that have ac- ver, Roseman, and Stringer (2007, 2008) demonstrated that tually been observed in nature are weak in strength; alleles if one applies a model of neutral evolution to expected di- under strong positive selection rapidly move to fixation while vergence in cranial dimensions, the observed morphological alleles under strong negative selection are rapidly removed divergence between humans and Neanderthals could be ex- from a population (Futuyma 1986). plained solely as the result of genetic drift over the last 350 In a larger population, one expects more of the rare, fa- kyr. In addition, some recent approaches to cultural inno- vorable mutations to arise simply because the number of new vations also emphasize the role of chance, especially if change mutations varies with population size (Cochran and Harp- is dependent on population size and density (e.g., Powell, ending 2009; Hawks et al. 2007). Large populations also tend Shennan, and Thomas 2009; Shennan 2001). These results to moderate, often to a great degree, the effects of drift. Thus, S224 Current Anthropology Volume 54, Supplement 8, December 2013 large populations provide favorable conditions for the pro- East Africa and across the Sahel and much of the Sahara from duction of new, beneficial mutations; large and growing pop- a northeasterly counterclockwise direction. Site ODP 659 also ulations provide the most fertile ground for new mutations receives dust from Western Sahara and portions of the north- to arise and increase in frequency. ern Sahara during these months. Under a neutral model of evolution, most new mutations Ample precipitation over East Africa and the southern Sa- are lost to drift (especially in small or numerically stable pop- hara and Sahel promote the growth of vegetation, which de- ulations). In growing populations, new mutations are more creases the amount of dust that winds scour off of the land. likely to be preserved, while in shrinking populations they are Periods of decreased precipitation diminish the amount of more likely to be lost because of drift (Harpending et al. 1993, vegetation and dependent biomass (including humans) and 1998). Drift slows in large populations but accelerates in small produce more dust. The Arabian Sea dust core shows a populations and can override the signal of all but the strongest 100,000-year oscillation between wet and dry with the most selective pressures. As a result, population size emerges as a intense and long-lasting dry periods corresponding to the key variable in both selection and drift. Estimating population major glacial advances in the Northern Hemisphere (fig. 1). size in the past is difficult and invariably requires one to make Major dry phases would have been guaranteed to produce a series of assumptions that are open to criticism. The prob- greatly expanded Sahara and Kalahari deserts and unfavorable lem may not be intractable, however, because during the Mid- conditions for human habitation. This seems to have hap- dle and Upper Pleistocene, recurrent 100,000-year-long glacial pened many times in the past, with conditions in OIS 2 serv- cycles drove climate change and almost certainly affected ing as a case in point (Brooks and Robertshaw 1990; Deacon hominin populations. and Lancaster 1988). In addition, cores and seismology of several of the oldest East African great lakes, especially Lakes Geology and Paleoclimate Malawi and Tanganyika, have shown that substantial portions of tropical Africa south of the equator experienced severe The climate in Europe in the Middle-Upper Pleistocene was droughts over the last 200 kyr that would not have been dominated by a high-amplitude 100,000-year cycle that ap- inferred from the oxygen isotope curve (Burnett et al. 2010; pears to have been determined by the eccentricity cycle in the Cohen et al. 2007; Scholz et al. 2011), although the effects of earth’s orbit around the sun (deMenocal 2004). The glacial these droughts appear to have been mitigated or absent at cycles show up very clearly in oxygen isotope values from the equator and cannot be generalized to the whole of Africa deep-sea cores and ice cores from Greenland and Antarctica (Blome et al. 2012). Some of the reconstructed lake levels (deMenocal 2004). The paleoclimate of Africa presents a more (e.g., for Lake Malawi) do not closely follow the dust curves complicated picture but one that is ultimately related to or- (fig. 2), suggesting yet another layer of complexity in the bital dynamics because of changes in air circulation and rain- climatic record. fall that arose as consequences of the amount of solar radi- In Europe, major glaciations appear to have pushed hom- ation (insolation) that reached the earth (Siddall et al. 2010; inins out of the northern European plain and Britain and Trauth, Larrasoan˜a, and Mudelsee 2009). In Africa, oscilla- into southern refugia along the Mediterranean Sea (Dennell, tions in precipitation were more crucial than temperature, Martino´n-Torres, and Bermu´dez de Castro 2011; Stringer and paleoclimatic records show that precipitation fluctuated 2006). Very wet and warm periods in Europe produced dense dramatically in Africa during the Pleistocene. forests that may have also been unfavorable habitat (Roe- Records of dust flux from deep-sea cores such as ODP 721/ broeks, Conard, and Van Kolfschoten 1992), although inter- 722 in the Arabian Sea and ODP 659 off of the coast of stadial periods seem to have been far more favorable for hom- Mauritania can serve as proxies for precipitation in East Africa inin populations than the coldest periods of glaciations. and the western Sahel and southern Sahara (deMenocal 2004; Trauth, Larrasoan˜a, and Mudelsee 2009). The record stems Predictions from long-standing patterns of atmospheric circulation. In June, July, and August, clockwise-circulation monsoonal The direct influence of climatic conditions on population sizes winds blow moisture onshore in Somalia from the Indian in Europe and Africa allows a series of predictions about the Ocean and carry dust from Somalia into the Arabian Sea. relative ability of selection and drift to produce changes in During the same months the southerly Shamai winds scour hominin populations. Periods of large-scale glacial advance dust off of the Arabian Peninsula and deposit it in the Arabian in Europe should produce periods of stress and low popu- Sea. The dust-flux record from the Arabian Sea, ODP 721/ lation numbers and rapid genetic drift in Neanderthals. In 722, therefore records both of these influences. Site ODP 659, contrast, periods of warmer but not yet heavily forested con- off of the coast of Mauritania and Western Sahara, receives ditions would have supported a higher biomass of large her- a substantial amount of its dust from the southern Sahara bivores and the humans who preyed on them (Roebroeks, and northern Sahel during these months. During December, Conard, and Van Kolfschoten 1992), thus producing an in- January, and February, the direction of air circulation reverses crease in hominin population numbers and a decelerated rate over East Africa, and the Trade Winds blow air onshore over of drift. Figure 1. Key events in genetic evolution, ages of fossil specimens, oxygen isotope stage (OIS) curves, and dust-flux data from the Arabian Sea. OIS curves adapted from Klein (2009); dust-flux curves from Donges et al. (2012). S226 Current Anthropology Volume 54, Supplement 8, December 2013

Figure 2. Water levels in Lake Malawi over the last 145,000 years and 300,000 years of dust flux from different sea cores. Water levels adapted from Scholz et al. (2011); dust-flux data after Donges et al. (2012).

Mellars and French (2011) have argued that Neanderthals Italy, and the Balkans. At the higher end of estimates, Sørensen in southwestern France had population numbers during the (2011) proposed a population of less than 10,000 individuals Wu¨rm Glaciation (OIS 4–3) that totaled approximately one- during the Eemian interglacial (OIS 5e), when Neanderthal tenth (actually 1/9) as many individuals as the later Aurig- numbers and inhabited territory may have been at a peak. nacian occupation, although many of the assumptions that Such low numbers would make sense given the large terri- led to this conclusion have been challenged (Dogandzˇic´and tories required for the Neanderthal specialization on large- McPherron 2013) and defended (Mellars and French 2013). bodied prey (Stiner 2013). If these remarkably low population Mellars and French (2011) refrained from proposing an es- numbers are accurate, Neanderthals may never have been timate of the number of individuals this would have involved, numerous enough to experience conditions in which there but Bocquet-Appel et al.’s (2005) model for human popu- were enough individuals for favorable mutations to arise at lation in Upper Paleolithic Europe produced an estimate of a brisk pace. Rather, they may have been skirting the edge of the population of Europe during the Aurignacian of 4,424 extinction for most of their existence, generally losing genetic people (95% confidence interval [CI]: 1,738–28,359). If one diversity as they did so. Neanderthals may have only rarely accepts Bocquet-Appel et al.’s (2005) estimates for the Au- experienced periods of population growth and range expan- rignacian and extends Mellars and French’s (2011) conclu- sion. sions to the whole of Europe, it would imply that the Ne- New data from the Denisovan genome from Siberia (Meyer anderthal population of Europe only totaled 492 individuals et al. 2012) suggest that this population also had a strikingly (95% CI: 193–3,151). Reviewing previous estimates of Ne- low long-term effective population size of approximately p anderthal population numbers, Dennell, Martino´n-Torres, Ne 1,667 individuals for the period between 400 and 100 and Bermu´ dez de Castro (2011) proposed the Neanderthal ka (Li, Patterson, and Reich 2012). It is important to bear in population of Europe totaled 3,000–5,000 during interstadials mind that effective population size can be different from (and and 1,500–2,500 during the depths of glacial advances, when sometimes lower by an order of magnitude or more) census

Neanderthal populations were confined to refugia in Iberia, size (the actual number of individuals) and that Ne approx- Pearson Hominin Evolution S227 imates the harmonic mean of the number of breeding indi- lineages of Neanderthals or modern humans, then major viduals over time. Nevertheless, for an effective population changes in anatomy in each lineage should emerge during size to shrink from 16,667 before 400 ka to 1,667 after 400 periods that favor large population numbers. On the other ka as the Denisovans did and apparently remained (Li, Pat- hand, if drift was the key force in driving the divergence of terson, and Reich 2012), the population must have crashed Neanderthals and early modern humans, then key evolution- to 1,667 individuals (or fewer) one or more times. ary events and appearance of new morphologies should ap- Lest one think that Neanderthals and Denisovans were fun- pear during or immediately after periods of low population damentally different from modern humans in the face of numbers. As new morphologies may be effectively invisible climatic instability, it is important to realize that some recent in the fossil record during periods of contracted population research to model effective population size in modern human size, they may, in fact, appear in the record only slightly later, populations based on genomic data suggests that both the once population sizes had rebounded. ancestors of living Europeans and Chinese experienced one Any test of these hypotheses faces practical limitations, in- or more severe bottlenecks between 40 and 20 ka such that cluding an incomplete fossil record, poor dating of some fos- the effective population size of each of these populations sils, and inadequate resolution of current methods in pin- shrank to a size of approximately N p 1,200 during this e pointing morphological or genetic changes to exact spots in interval before rebounding to a higher size (to Ne between 11,000 and 50,000) during the Holocene (Li and Durbin the 100,000-year glacial and faster insolation cycles. Margins 2011). This result implies that similar dynamics, likely at- of error for dates for fossils or genetic events may overlap tributable to climatic cycles, affected archaic and modern pop- both favorable and unfavorable periods of climatic cycles. A ulations in Eurasia in very similar ways. further difficulty particular to Africa lies in the variability of Recently, a number of authors have stressed that climatic dust-flux records: different patterns occur in different cores deterioration in Europe and the Near East could have led to around Africa (fig. 2). For the sake of argument, I assume in the local extinction of populations (Dennell, Martino´n-Torres, this paper that the record of dust flux from the Arabian Sea and Bermu´dez de Castro 2011; Hublin and Roebroeks 2009; is the most relevant to the origin of modern humans, but Shea 2011; Stewart and Stringer 2012; Stringer 2006). Britain, this issue is certainly open to debate. As a case in point, Blome in particular, seems to have been abandoned with each major et al. (2012) synthesized paleoclimatic records for the whole glacial advance and then reoccupied, at least as long as a land of Africa using multiple proxies, including terrestrial, lacus- bridge connected it to the continent (Stringer 2006). Heinrich trine, and oceanic data (fig. 3). The resulting synthesis depicts events, short periods of extreme cold followed by rapid warm- a mosaic of wet and dry periods that are frequently asyn- ing, during glaciations may have posed especially difficult chronous between regions and do not correspond in a con- challenges for hominins in Europe (Stewart and Stringer sistent way to the OISs. Their results for East Africa are per- 2012) and perhaps contributed to a contraction in the range haps the most useful for inferences regarding the origin and of Neanderthals in southern Iberia and the spread of modern dispersal of modern humans. Likewise, Rohling et al. (2013) humans bearing Aurignacian technology into France and present multiproxy data for the Mediterranean and Red Sea northern Spain (d’Errico and Sa´nchez Gon˜i 2003). regions, two areas that were crucial for hominin dispersals In Africa, especially in East Africa, biomass and human from (and perhaps into) Africa during the last 500 kyr. The population size were much more dependent on the availability arrows in figure 3 extend the analyses of Blome et al. (2012) (and predictability) of precipitation, and the dust-flux data by indicating possible population expansions within Africa from deep-sea cores provide an accessible gauge of precipi- and possible expansions into Arabia during the windows of tation (deMenocal 2004; Rohling et al. 2013). Periods of low opportunity described by Rohling et al. (2013). In each case, dust flux indicate more precipitation, more vegetation, more animal biomass, and more people. During these times selec- populations can be inferred to have spread from regions with tion would logically have more power to create phenotypic favorable climate and thus presumably comparatively high change, and genetic drift would be less influential. Periods of human population density into regions previously nearly de- high dust flux correspond to less precipitation, less vegetation, void of people but with newly favorable climatic conditions. fewer people, and thus rapid genetic drift. The scale of the effect of climatic changes on human populations is clearly apparent in the dramatic decrease in the number of sites in Results the Last Glacial Maximum in East Africa (Brooks and Rob- ertshaw 1990); difficulties for human populations likely con- Comparison of the time lines of paleoclimate, the fossil rec- tinued even after that, including evidence of the desiccation ord, and genetic divergences and bottlenecks provide a rough of Lake Victoria until ca. 14.5 ka (Williams et al. 2006) and check of whether key events occur in periods favorable for the desiccation of Lake Tana around the same time (Lamb et large population numbers or in periods unfavorable for large al. 2007; Marshall et al. 2011). populations (fig. 1). In many cases, the relationships are highly If selection was the crucial factor driving change in the suggestive, but problems remain. S228 Current Anthropology Volume 54, Supplement 8, December 2013

Figure 3. A, Geographical regions in Africa adapted from Blome et al. (2012, with permission from Elsevier). B, Climate in Africa adapted from Blome et al. (2012, with permission from Elsevier) with two periods of a wet Sahara coinciding with periods of low sea level (following Rohling et al. 2013) indicated. The arrows show possible directions of colonization from regions of higher population density into adjacent areas. Question marks symbolize that the geographic source of the colonizing population is uncertain (and thus, one might assume, largely of local origin).

Europe mtDNA of a Neanderthal from Valdegoba, Spain, dating to 48.5 ka, shows that all Neanderthal mtDNA sequences post- The age of the fossils from Sima de los Huesos is a key dating this time formed a compact, monophyletic group problem for making sense of the tempo and mode of hominin within the known Neanderthal sequences (Dale´n et al. 2012). evolution in Europe over the last 500 kyr (Stringer 2012). If These late, western Neanderthal mtDNA sequences have a the Sima de los Huesos dates to 500–600 ka (Bischoff et al. 2007), then one can conclude the features characteristic of coalescent age of 58 ka (the end of OIS 4) with a 95% CI the later, “classic” Neanderthals dating to OIS 4–3 increased 54–77 ka. A severe population bottleneck (or selection, which in frequency very slowly within the Neanderthal lineage. may be less likely) could produce such a pattern. Dale´n and Hominins with the full suite of Neanderthal cranial traits colleagues (2012) conducted an Approximate Bayesian Com- appear by OIS 7 (Hublin 2009), but a further pulse of change putation Analysis to test demographic models of neutral evo- made western European Neanderthals from OIS 4–3 especially lution that were the most likely to produce this pattern of distinctive. It is difficult to tell whether this apparently late greatly reduced diversity. The results showed that the most inflection in the rate of “Neanderthalization” was the result probable scenario (isolation and drift in the western and east- of selection within a large population during OIS 5 or of ern populations of Neanderthals at 48 ka) would have in- rapid drift in a small population during OIS 4–3. volved an effective population size (Ne) of western Neander- Neanderthal mtDNA sequences provide support for a late thals of only 300 females, a marked reduction from the p bottleneck in their population. A recent analysis of the ancient estimate for the eastern subpopulation (Ne 2,000 females). Pearson Hominin Evolution S229

The effective population size for the autosomal genes in the that the last interval in East Africa was interrupted by a wet entire population is expected to be four times that of mtDNA period around 55–50 ka. The origin of modern humans dates p (i.e., Ne 1,200 and 8,000 in the western and eastern sub- to OIS 6. The first fossils of recognizably modern form date populations, respectively). The usual cautions about the dif- to 195–160 ka (i.e., the end of OIS 7 and into OIS 6). All ference between census size and Ne apply, but one is left with extant mtDNA sequences coalesce to a common ancestor at the strong impression that western European Neanderthals 140–210 ka (Behar et al. 2008), and Y chromosomes coalesce -ka (Cruciani et al. 2011), although an ex 15.6 ע experienced a major population crash during OIS 4. at 141.5 A second possibility for Europe is that the fossils from the tremely rare Y-chromosome haplotype from an African Amer- Sima de los Huesos date to only around 350 ka. Stringer ican man was recently reported that coalesces with other Y (2012) argues that the younger age is supported by the mosaic chromosomes at 338 ka (Mendez et al. 2013). OIS 6 has been presence of many distinctive Neanderthal cranial, dental, and likened to the hyperarid conditions of the Last Glacial Max- postcranial features in the sample. These features are similarly imum in OIS 2 (Deacon and Lancaster 1988), which featured common in European fossils dating to 300–200 ka but much greatly decreased archaeological visibility of human popula- more rare (or absent) in earlier fossils from other sites in tions in much of Africa (Brooks and Robertshaw 1990). Ge- Europe. Stringer (2012) also notes that an age of 600–500 ka netic drift would be expected to be the dominant factor during for the Sima de los Huesos fossils would place them earlier such a period, but it is worth reiterating that African climate than the estimated population divergence times for the an- was a complex and regionally variable mosaic (Blome et al. cestors of modern humans and Neanderthals and that the 2012). dated spelothems may, in fact, have been breached by a flow Indirect evidence from autosomal genes also supports the of younger sediments within the cave so that younger strata hypothesis that the African ancestors of modern humans ex- containing the hominins now underlie an only partially com- perienced a major population bottleneck during this period. plete but older spelothem. However, Spanish researchers pre- Fagundes et al. (2007) simulated several scenarios for the fer the older date, noting that the younger age is contradicted origin of modern humans with a sample of 50 autosomal loci by fossil fauna from the same deposit as the hominins, in- that were subsequently compared with observed patterns of cluding relatively primitive fossils of Ursus deningeri and the variation in human nuclear loci. They found the best cor- vole Clethrionomys acrorhiza (Garcı´a and Arsuaga 2011). Nev- respondence to observed patterns of human genetic variation ertheless, if the hominins from Sima de los Huesos date to in a model that features an origin of modern humans in Africa around 350 ka, the time span for drift would be cut in half, followed by exponential population growth, expansion from implying a more rapid pace of Neanderthalization later in the Africa and replacement of archaic hominins outside of Africa sequence. If this was the case, much of the genetic and mor- (specifically in Asia in their model) followed by exponential phological change may have been concentrated in bursts of population growth in Asia, and finally a migration from Asia drift that corresponded to major contractions in Neanderthal to the Americas followed by a final burst of exponential pop- numbers during OIS 8, 6, and 4. A shortened time span would ulation growth in the New World. The best-fitting model also create a stronger association between expansion in brain produced a series of posterior estimates for demographic and size and adoption of Middle Paleolithic technologies between historical parameters, including the age of the speciation event 300 and 200 ka. that produced modern humans (median: 141,455; 95% CI: 103,535–185,642), the age of the migration from Africa (me- dian: 51,102; 95% CI: 40,135–70,937), the age of the colo- Africa nization of the Americas (median: 10,280; 95% CI: 7,647– The Arabian Sea dust core shows a long relatively wet and 15,945), the size of the archaic African population (median: stable period between 640 and 427 ka. This period is asso- 12,772; 95% CI: 6,604–20,211), the population size during ciated with the first appearance of Homo heidelbergensis (or the bottleneck during speciation (median: 600; 95% CI: 76– Homo rhodesiensis, if this name is to be preferred) in Africa 1,620), the size of the bottleneck when leaving Africa (median: (i.e., the Bodo cranium, dated to 600 ka; Clark et al. 1994; 462; 95% CI: 64–1,224), and the size of the bottleneck when Rightmire 1996) and, intriguingly, marked technological ad- leaving Asia to settle the Americas (median: 452; 95% CI: 71– vances represented by precociously early blade production and 1,280). core technology in the Kapthurin Formation at Lake Baringo Not all of the genetic data supports the conclusion that a (Johnson and McBrearty 2010; Tryon and McBrearty 2006). population bottleneck produced modern humans, and some It is possible that large population sizes in Africa during much of the data strongly contradict that hypothesis. Using data of the Middle Pleistocene drove both cultural innovations and from complete genomes of several modern men comprising anatomical evolution via positive selection on beneficial new two Yoruba, three Europeans, one Chinese, and one Korean, mutations. Li and Durbin (2011) applied population genetics models to The dust core also indicates marked dry periods in East infer changes in human effective population size over the last Africa during OIS 8 (301–242 ka), OIS 6 (186–127 ka), and million years. Intriguingly, their data showed no evidence of OIS 4–2 (71–12 ka), although Blome et al. (2012) suggest a bottleneck between 200 and 100 ka. In fact, their results S230 Current Anthropology Volume 54, Supplement 8, December 2013 show growth in effective population size from ∼450 ka until Conclusions 120–150 ka and very similar histories (and likely shared his- As a result of this comparison of records of paleoclimate, tories in an ancestral source population) of Yoruban, eastern morphological change, and genetic change, it seems apparent Asian, and European population size before 60 ka. After that, that many of the observed changes leading to Neanderthals all three populations experienced bottlenecks, although the were more likely to have been the products of drift than one that affected the ancestors of the Yoruba appears to have selection, whereas both drift and selection may have been been less severe and allowed an earlier recovery. important in the emergence of modern humans. Many of the Likewise, by applying a population genetics model to ex- key events appear to date to periods in which population sizes pectations for (neutral) change in cranial dimensions, Weaver were greatly reduced and genetic drift would have been rapid. (2012) showed that crania that had dimensions that differed The picture that emerges is one of human population history by one standard deviation from modern crania could be ex- that was highly (although almost certainly not exclusively) pected by around 165 ka, which corresponds reasonably well contingent on climatic changes. to when most researchers agree that modern (or nearly mod- One prominent example of this dependence on climate ern) humans appear in the East African fossil record. Weaver’s comes from mtDNA intramatch distributions that show rapid model, however, assumes a constant effective population. A population growth in Africa at ca. 80 ka (Harpending et al. potential explanation for the apparent lack of a contraction 1993, 1998; Sherry et al. 1994); estimates of lake levels from in the effective population size of the ancestors of modern Lakes Malawi and Tangyanika show a return of wet conditions humans in Africa is that if there were in fact bottlenecks within at the same time (Burnett et al. 2010; Scholz et al. 2011), a subdivided population in Africa, following the bottlenecks, although the pattern of wet and dry periods for Africa as a members of dissimilar populations mixed extensively, restor- whole forms a complex mosaic that frequently departs from ing to the resulting population much of the genetic variation the pattern observed in Lakes Malawi and Tangyanika (Blome that existed before each bottleneck. The complex mosaic of et al. 2012). Rohling et al. (2013) demonstrate that favorable favorable climates over time in different parts of Africa re- conditions combining low sea levels with elevated levels of ported by Blome et al. (2012) may have provided the right precipitation to support terrestrial biomass (including hu- conditions for this sort of mechanism (fig. 3). mans) would have facilitated movements across the Bab-el- Given that both climatic and genomic data suggest a bot- Mendab strait between East Africa and Arabia only in narrow tleneck in East Africa and Arabia after 60 ka, it is highly likely windows of time, the latest of which dates to 70–65 ka. These that a substantial amount of genetic drift occurred in the patterns help to illuminate a particularly irksome issue in population of modern humans as they left Africa or for a research on the origin of modern humans: the question of period of time immediately afterward. As a result, outbreeding why modern humans only expanded out of Africa at 50–60 would have been highly favorable if heterozygosity was greatly ka if “anatomically modern” morphology arose between 200 increased by these events, especially for loci such as the major and 150 ka (e.g., Klein 2009). The answer seems to be that histocompatibility complex, in which alleles from archaic Eur- climatic conditions did not favor a large, interconnected pop- asian populations are far more frequent in populations out- ulation in Africa between 125–ca. 80 ka because each region side of Africa than they are in other loci (Abi-Rached et al. experienced one or more dry periods during this interval 2011). (Blome et al. 2012). Many of the famous cultural advances Low lake levels in Lake Malawi and Lake Tanganyika and associated with the Upper Paleolithic and Late Stone Age are high levels of dust flux suggest generally unfavorable condi- also likely to have depended on population size and density tions for human population growth in tropical Africa during (Powell, Shennan, and Thomas 2009; Premo and Kuhn 2010). much of OIS 5 (Blome et al. 2012; Scholz et al. 2011). The result was likely relatively rapid genetic drift and population differentiation among modern humans in Africa. Another population contraction in most of Africa in OIS 2 probably Acknowledgments accounts for late (Holocene or terminal Pleistocene) appear- Many thanks to Erella Hovers and Steve Kuhn for the invi- ance of crania that have cranial metrics that cannot be dis- tation to participate in this stimulating conference and to the tinguished from one or more extant populations (De Villiers Wenner-Gren Foundation for organizing and funding it. In- and Fatti 1982; Habgood 1989). Early Holocene (and likely sightful comments by conference participants, two anony- from the end of OIS 2) human remains show evidence of mous reviewers, and Steve Kuhn helped to refine the ideas strong morphological differentiation among African popu- presented here; any remaining faults are my own. lations. The spread of pastoralism and agricultural popula- tions in Africa has blurred or erased these stark distinctions (Tishkoff et al. 2009). For Africa, then, the dust-core and References Cited genetic data suggest that selection may have been important Abi-Rached, Laurent, Matthew J. Jobin, Subhash Kulkarni, Alasdair Mc- Whinnie, Klara Dalva, Loren Gragert, Farbod Babrzadeh, et al. 2011. The from 600–400 ka, but periods of drift had more potential to shaping of modern human immune systems by multiregional admixture be the dominant influence thereafter. with archaic humans. Science 334:89–94. Pearson Hominin Evolution S231

Arsuaga, J.-L., I. Martı´nez, A. Gracia, and C. Lorenzo. 1997. The Sima de los Clement, Anna F., Simon W. Hillson, and Leslie C. Aiello. 2012. Tooth wear, Huesos crania (Sierra de Atapuerca, Spain): a comparative study. Journal Neanderthal facial morphology and the anterior dental loading hypothesis. of Human Evolution 33:219–281. Journal of Human Evolution 62:367–376. Arsuaga, Juan-Luis. 2010. Terrestrial apes and phylogenetic trees. Proceedings Cochran, Gregory, and Henry Harpending. 2009. The 10,000 year explosion: of the National Academy of Sciences of the USA 107:8910–8917. how civilization accelerated human evolution. New York: Basic. Arsuaga, Juan-Luis, Carlos Lorenzo, Jose´-Miguel Carretero, Ana Gracia, Ig- Cohen, Andrew S., Jeffrey R. Stone, Kristina R. M. Beuning, Lisa E. Park, nacio Martı´nez, Nuria Garcı´a, Jose´-Marı´a Bermu´dez de Castro, and Eudald Peter N. Reinthal, David Dettman, Christopher A. Scholz, et al. 2007. Eco- Carbonell. 1999. A complete human pelvis from the Middle Pleistocene of logical consequences of early Late Pleistocene megadroughts in tropical Spain. Nature 399:255–258. Africa. Proceedings of the National Academy of Sciences of the USA 104: Behar, Doran M., Richard Villems, Himla Soodyall, Jason Blue-Smith, Luı´sa 16422–16427. Pereira, Ene Metspalu, Rosaria Scozzari, et al. 2008. The dawn of human Coon, C. S. 1962. The origin of races. New York: Knopf. matrilineal diversity. American Journal of Human Genetics 82:1130–1140. Cruciani, Fulvio, Beniamino Trombetta, Andrea Massaia, Giovanni Destro- Betti, Lia, Franc¸ois Balloux, William Amos, Tsunehiko Hanihara, and Andrea Bisol, Daniele Sellitto, and Rosaria Scozzari. 2011. A revised root for the Manica. 2009. Distance from Africa, not climate, explains within-population human Y chromosomal phylogenetic tree: the origin of patrilineal diversity phenotypic diversity in humans. Proceedings of the Royal Society B 276:809– in Africa. American Journal of Human Genetics 88:814–818. 814. Dale´n, Love, Ludovic Orlando, Beth Shapiro, Mikael Brandstro¨m Durling, Betti, Lia, Franc¸ois Balloux, Tsunehiko Hanihara, and Andrea Manica. 2010. Rolf Quam, M. Thomas P. Gilbert, J. Carlos Dı´ez Ferna´ndez-Lomana, Eske The relative role of drift and selection in shaping the human skull. American Willerslev, Juan Luis Arsuaga, and Anders Go¨therstro¨m. 2012. Partial ge- Journal of Physical Anthropology 141:76–82. netic turnover in Neandertals: continuity in the East and population re- Betti, Lia, Noreen von Cramon-Taubadel, and Stephen J. Lycett. 2012. Human placement in the West. Molecular Biology and Evolution 29:1893–1897. pelvis and long bones reveal differential preservation of ancient population Day, M. H., and C. B. Stringer. 1991. Les restes craˆniens d’Omo-Kibish et history and migration out of Africa. Human Biology 84:139–152. leur classification a`l’inte´rieur du genre Homo. L’Anthropologie 95:573–594. Bischoff, James L., Ross W. Williams, Robert J. Rosenbauer, Arantza Aram- Deacon, Janette, and N. Lancaster. 1988. Late Quaternary palaeoenvironments buru, Juan Luis Arsuaga, Nuria Garcı´a, and Gloria Cuenca-Besco´s. 2007. of southern Africa. Oxford: Oxford University Press. High-resolution U-series dates from the Sima de los Huesos hominids yields Dean, Christopher, Meave G. Leakey, Donald Reid, Friedemann Schrenk, Gary ϩϱ 600Ϫ66 kyrs: implications for the evolution of the early Neanderthal lineage. T. Schwartz, Christopher Stringer, and Alan Walker. 2001. Growth processes Journal of Archaeological Science 34:763–770. in teeth distinguish modern humans from Homo erectus and earlier hom- Blome, Margaret Whiting, Andrew S. Cohen, Christian A. Tryon, Alison S. inins. Nature 414:628–631. Brooks, and Joellen Russell. 2012. The environmental context for the origins Dean, M. Christopher, and B. Holly Smith. 2009. Growth and development of modern human diversity: a synthesis of regional variability in African of the Nariokotome youth, KNM-WT 15000. In The first humans: origin climate 150,000–30,000 years ago. Journal of Human Evolution 62:563–592. and early evolution of the genus Homo. Frederick E. Grine, John G. Fleagle, Bocquet-Appel, Jean-Pierre, Pierre-Yves Demars, Lorette Noiret, and Dmitry and Richard E. Leakey, eds. Pp. 101–120. New York: Springer. Dobrowsky. 2005. Estimates of Upper Paleolithic meta-population size in deMenocal, Peter B. 2004. African climate change and faunal evolution during Europe from archaeological data. Journal of Archaeological Science 32:1656– the Pliocene-Pleistocene. Earth and Planetary Science Letters 220:3–24. 1668. Dennell, Robin W., Marı´a Martino´n-Torres, and Jose´M. Bermu´dez de Castro. Bonmatı´, Alejandro, Asier Go´mez-Olivencia, Juan-Luis Arsuaga, Jose´ Miguel 2011. Hominin variability, climatic instability and population demography Carretero, Ana Gracia, Ignacio Martı´nez, Carlos Lorenzo, Jose´ Marı´a Be´r- in Middle Pleistocene Europe. Quaternary Science Reviews 30:1511–1524. mudez de Castro, and Eudald Carbonell. 2010. Middle Pleistocene lower d’Errico, Francesco, and Maria Fernanda Sa´nchez Gon˜i. 2003. Neandertal back and pelvis from an aged human individual from the Sima de los extinction and the millennial scale climatic variability of OIS 3. Quaternary Huesos site, Spain. Proceedings of the National Academy of Sciences of the Science Reviews 22:769–788. USA 107:18386–18391. De Villiers, H., and L. P. Fatti. 1982. The antiquity of the Negro. South African Bra¨uer, Gu¨nter. 2008. The origin of modern anatomy: by speciation or in- Journal of Science 78:321–332. traspecific evolution? Evolutionary Anthropology 17:22–37. Dogandzˇic´, Tamara, and Shannon P. McPherron. 2013. Demography and the Brooks, Alison S., and Peter Robertshaw. 1990. The Glacial Maximum in demise of Neandertals: a comment on “Tenfold population increase in tropical Africa: 22,000–12,000 BP. In Low latitudes,vol.2ofThe world at Western Europe at the Neandertal-to-modern human transition.” Journal 18,000 BP. Clive Gamble and Oolda Soffer, eds. Pp. 121–169. London: of Human Evolution 64:311–313. Unwin Hyman. Donges, Jonathan F., Reik V. Donner, Martin H. Trauth, Norbert Marwan, Brown, Francis H., Ian McDougall, and John G. Fleagle. 2012. Correlation of Hans-Joachim Schellnhuber, and Ju¨rgen Kurths. 2012. Nonlinear detection the KHS Tuff of the Kibish Formation to volcanic ash layers at other sites, of paleoclimate-variability transitions possibly related to human evolution. and the age of early Homo sapiens (Omo I and Omo II). Journal of Human Proceedings of the National Academy of Sciences of the USA 108:20422–20427. Evolution 63:577–585. Fagundes, Nelson J. R., Nicholas Ray, Mark Beaumont, Samuel Neuen- Burnett, Allison P., Michael J. Soreghan, Christopher A. Scholz, and Erik T. schwander, Francisco M. Salzano, Sandro L. Bonatto, and Laurent Excoffier. Brown. 2010. Tropical East African climate change and its relation to global 2007. Statistical evaluation of alternative models of human evolution. Pro- climate: a record from Lake Tanganyika, tropical East Africa, over the past ceedings of the National Academy of Sciences of the USA 104:17614–17619. 90 kyr. Palaeogeography, Palaeoclimatology, Palaeoecology 303:155–167. Futuyma, Douglas J. 1986. Evolutionary biology. 2nd edition. Sunderland, MA: Carretero, Jose´-Miguel, Laura Rodrı´guez, Rebeca Garcı´a-Gona´lez, Juan-Luis Sinauer. Arsuaga, Asier Go´mez-Olivencia, Carlos Lorenzo, Alejandro Bonmatı´, Ana Garcı´a, Nuria, and Juan Luis Arsuaga. 2011. The Sima de los Huesos (Burgos, Gracia, Ignacio Martı´nez, and Rolf Quam. 2012. Stature estimation from northern Spain): palaeoenvironment and habitats of Homo heidelbergensis complete long bones in the Middle Pleistocene humans from the Sima de during the Middle Pleistocene. Quaternary Science Reviews 30:1413–1419. los Huesos, Sierra de Atapuerca (Spain). Journal of Human Evolution 62: Graves, Ronda R., Amy C. Lupo, Robert C. McCarthy, Daniel J. Wescott, and 242–255. Deborah L. Cunningham. 2010. Just how strapping was KNM-WT 15000? Churchill, Steven E., Lee R. Berger, Adam Hartstone-Rose, and B. Headman Journal of Human Evolution 59:542–554. Zondo. 2012. Body size in African Middle Pleistocene Homo.InAfrican Green, Richard E., Johannes Krause, Adria W. Briggs, Tomislav Maricic, Udo genesis: perspectives on hominin evolution. S. C. Reynolds and A. Gallagher, Stenzel, Martin Kircher, Nick Patterson, et al. 2010. A draft of the Nean- eds. Pp. 319–346. Cambridge: Cambridge University Press. dertal genome. Science 328:710–722. Clark, J. D., J. de Heinzelin, K. D. Schick, W. K. Hart, T. D. White, G. Gunz, Philipp, Simon Neubauer, Lubov Golovanova, Vladimir Doronichev, WoldeGabriel, R. C. Walter, et al. 1994. African Homo erectus: old radio- Bruno Maureille, and Jean-Jacques Hublin. 2012. A uniquely modern hu- metric ages and young Oldowan assemblages in the Middle Awash Valley, man pattern of endocranial development: insights from a new cranial re- Ethiopia. Science 264:1907–1910. construction of the Neandertal newborn from Mezmaiskaya. Journal of Clark, J. Desmond, Yonas Beyene, Giday WoldeGabriel, William K. Hart, Paul Human Evolution 62:300–313. R. Renne, Henry Gilbert, Alban Defleur, et al. 2003. Stratigraphic, chro- Habgood, P. J. 1989. An examination of regional features on Middle and early nological and behavioral contexts of Pleistocene Homo sapiens from Middle Late Pleistocene sub-Saharan African hominids. South African Archaeological Awash, Ethiopia. Nature 423:747–752. Bulletin 44:17–22. S232 Current Anthropology Volume 54, Supplement 8, December 2013

Harpending, H. C., S. T. Sherry, A. R. Rogers, and M. Stoneking. 1993. The placement and age of modern humans from Kibish, Ethiopia. Nature 433: genetic structure of ancient human populations. Current Anthropology 34: 733–736. 483–496. Mellars, Paul, and Jennifer C. French. 2011. Tenfold population increase in Harpending, Henry C., Mark A. Batzer, Michael Gurven, Lynn B. Jorde, Alan western Europe at the Neandertal-to-modern human transition. Science 333: R. Rogers, and Stephen T. Sherry. 1998. Genetics and traces of ancient 623–627. demography. Proceedings of the National Academy of Sciences of the USA 95: ———. 2013. Population changes across the Neanderthal-to-modern-human 1961–1967. transition in western France: a reply to Dogandzˇic´ and McPherron (2013). Harris, Kelley, and Rasmus Nielsen. 2013. Inferring demographic history from Journal of Human Evolution 65:330–333. a spectrum of shared haplotype lengths. PLoS Genetics 9:e1003521. Mendez, Fernando L., Thomas Krahn, Bonnie Schrack, Astrid-Maria Krahn, Harvati, Katerina, Jean-Jacques Hublin, and Philipp Gunz. 2010. Evolution Krishna R. Veeramah, August E. Woerner, Forka Leypey Mathew Fomine, of Middle-Late Pleistocene human cranio-facial morphology: a 3-D ap- et al. 2013. An African American paternal lineage adds an extremely ancient proach. Journal of Human Evolution 59:445–464. root to the human Y chromosome phylogenetic tree. American Journal of Hawks, John, Eric T. Wang, Gregory M. Cochran, Henry C. Harpending, and Human Genetics 92:454–459. Robert K. Moyzis. 2007. Recent acceleration of human adaptive evolution. Meyer, Matthias, Martin Kircher, Marie-Theres Gansauge, Heng Li, Fernando Proceedings of the National Academy of Sciences of the USA 104:20753–20758. Racimo, Swanpan Mallick, Joshua G. Schraiber, et al. 2012. A high-coverage Holliday, Trenton W. 1997. Body proportions in Late Pleistocene Europe and genome sequence from an archaic Denisovan individual. Science 338:222– modern human origins. Journal of Human Evolution 32:423–447. 226. Howell, F. C. 1957. The evolutionary significance of variation and varieties Pearson, Osbjorn M. 2000. Activity, climate, and postcranial robusticity: im- of “Neanderthal” man. Quarterly Review of Biology 32:330–347. plications for modern human origins and scenarios of adaptive change. Hublin, Jean-Jacques. 2001. Northwestern African Middle Pleistocene hom- Current Anthropology 41:569–607. inids and their bearing on the emergence of Homo sapiens.InHuman roots: ———. 2008. Statistical and biological definitions of “anatomically modern” Africa and Asia in the Middle Pleistocene. L. Barham and K. Robson-Brown, humans: suggestions for an integrated approach to modern morphology. eds. Pp. 99–121. Bristol, UK: Western Academic & Specialist Press. Evolutionary Anthropology 17:38–48. ———. 2009. The origin of Neandertals. Proceedings of the National Academy Pearson, Osbjorn M., Danielle F. Royer, Frederick E. Grine, and John G. of Sciences of the USA 106:16022–16027. Fleagle. 2008. A description of the Omo I postcranial skeleton, including Hublin, Jean-Jacques, and Wil Roebroeks. 2009. Ebb and flow or regional newly discovered fossils. Journal of Human Evolution 55:421–437. extinctions? on the character of Neandertal occupation of northern envi- Powell, Adam, Stephen Shennan, and Mark G. Thomas. 2009. Late Pleistocene ronments Comptes Rendus Palevol 8:503–509. demography and the appearance of modern human behavior. Science 324: Hublin, J.-J., and A. M. Tillier. 1981. The Mousterian juvenile mandible from 1298–1301. Irhoud (Morocco): a phylogenetic interpretation. In Aspects of human evo- Premo, L. S., and Steven L. Kuhn. 2010. Modeling effects of local extinctions lution. Chris Stringer, ed. Pp. 167–185. London: Taylor & Francis. on culture change and diversity in the Paleolithic. PLoS ONE 5:e15582. Hylander, William L. 1977. The adaptive significance of Eskimo craniofacial Rae, Todd C., Thomas Koppe, and Chris B. Stringer. 2011. The Neanderthal morphology. In Orofacial growth and development. A. A. Dahlberg and T. face is not cold adapted. Journal of Human Evolution 60:234–239. M. Graber, eds. Pp. 129–169. Paris: The Hague. Reich, David, Richard E. Green, Martin Kircher, Johannes Krause, Nick Pat- Johnson, Cara Roure, and Sally McBrearty. 2010. 500,000 year old blades from terson, Eric Y. Durand, Bence Viola, et al. 2010. Genetic history of an the Kapthurin Formation, Kenya. Journal of Human Evolution 58:193–200. archaic hominin group from Denisova Cave in Siberia. Nature 468:1053– Kappelman, John. 1996. The evolution of body mass and relative brain size 1060. in fossil hominids. Journal of Human Evolution 30:243–276. Rightmire, G. Philip. 1996. The human cranium from Bodo, Ethiopia: evidence Klein, Richard G. 2009. The human career. 3rd edition. Chicago: University for speciation in the Middle Pleistocene? Journal of Human Evolution 31: of Chicago Press. 21–39. Lamb, Henry F., C. Richard Bates, Paul V. Coombes, Michael H. Marshall, ———. 2009. Middle and Later Pleistocene hominins in Africa and southwest Mohammed Umer, Sarah J. Davies, and Eshete Dejen. 2007. Late Pleistocene Asia. Proceedings of the National Academy of Sciences of the USA 106:16046– desiccation of Lake Tana, source of the Blue Nile. Quaternary Science Reviews 16050. 26:287–299. Roebroeks, Wil, Nicholas J. Conard, and Thijs Van Kolfschoten. 1992. Dense Li, Heng, and Richard Durbin. 2011. Inference of human population history forests, cold steppes, and the Palaeolithic settlement of Europe. Current from individual whole-genome sequences. Nature 475:493–496. Anthropology 33:551–586. Li, Heng, James Mullikin, and David Reich. 2010. Date of population diver- Rohling, Eelco J., Katherine M. Grant, Andrew P. Roberts, and Juan-Cruz gence between Neandertals and modern humans. Supplementary online Larrasoan˜a. 2013. Palaeoclimate variability in the Mediterranean and Red materials 14 to Green, Richard E., Johannes Krause, Adria W. Briggs, Tom- Sea regions during the last 500,000 years: implications for hominin migra- islav Maricic, Udo Stenzel, Martin Kircher, Nick Patterson, et al. 2010. A tions. Current Anthropology 54(suppl. 8):S183–S201. draft of the Neandertal genome. Science 328:710–722. Roseman, Charles C. 2004. Detecting interregionally diversifying natural se- Li, Heng, Nick Patterson, and David Reich. 2012. Inferred population size lection on modern human cranial form by using matched molecular and changes in the history of Denisovans. Supplementary note 17 to Meyer, morphometric data. Proceedings of the National Academy of Sciences of the Matthias, Martin Kircher, Marie-Theres Gansauge, Heng Li, Fernando Ra- USA 101:12824–12829. cimo, Swanpan Mallick, Joshua G. Schraiber, et al. 2012. A high-coverage Ruff, C. B. 1994. Morphological adaptation to climate in modern and fossil genome sequence from an archaic Denisovan individual. Science 338:222– hominids. Yearbook of Physical Anthropology 37:65–107. 226. Ruff, Christopher B., Erik Trinkaus, and Trenton W. Holliday. 1997. Body Lieberman, Daniel E. 2011. The evolution of the human head. Cambridge, MA: mass and encephalization in Pleistocene Homo. Nature 387:173–176. Belknap. Scholz, C. A., A. S. Cohen, T. C. Johnson, J. King, M. R. Talbot, and E. T. Lieberman, Daniel E., Brandeis M. McBratney, and Gail Krovitz. 2002. The Brown. 2011. Scientific drilling in the Great Rift Valley: the 2005 Lake evolution and development of cranial form in Homo sapiens. Proceedings Malawi Scientific Drilling Project: an overview of the past 145,000 years of of the National Academy of Sciences of the USA 99:1134–1139. climate variability in Southern Hemisphere East Africa. Palaeogeography, Marshall, Michael H., Henry F. Lamb, Dei Huws, Sarah J. Davies, Richard Palaeoclimatology, Palaeoecology 303:3–19. Bates, Jan Bloemendal, John Boyle, Melanie J. Leng, Mohammed Umer, Shea, John J. 2011. Transitions or turnovers? climatically-forced extinctions and Charlotte Bryant. 2011. Late Pleistocene and Holocene drought events of Homo sapiens and Neanderthals in the east Mediterranean Levant. Qua- at Lake Tana, the source of the Blue Nile. Global and Planetary Change 78: ternary Science Reviews 27:2253–2270. 147–161. Shennan, Stephen. 2001. Demography and cultural innovation: a model and Martino´n-Torres, Marı´a, Jose´Marı´a Bermu´dez de Castro, Aida Go´mez-Robles, its implications for the emergence of modern human culture. Cambridge Leyre Prado-Simo´n, and Juan Luis Arsuaga. 2012. Morphological descrip- Archaeological Journal 11:5–16. tion and comparison of the dental remains from Atapuerca-Sima de los Sherry, S. T., A. R. Rogers, H. Harpending, H. Soodyall, T. Jenkins, and M. Huesos site (Spain). Journal of Human Evolution 62:7–58. Stoneking. 1994. Mismatch distributions of mtDNA reveal recent human McDougall, Ian, Francis H. Brown, and John G. Fleagle. 2005. Stratigraphic population expansions. Human Biology 66:761–775. Pearson Hominin Evolution S233

Siddall, M., E. J. Rohling, T. Blunier, and R. Spahni. 2010. Patterns of millennial ———. 2011. The origin of our species. London: Allen Lane. variability over the last 500 ka. Climate of the Past 6:295–303. ———. 2012. The status of Homo heidelbergensis (Schoetensack 1908). Evo- Simpson, Scott W., Jay Quade, Naomi E. Levin, Robert Butler, Guillaume lutionary Anthropology 21:101–107. Dupont-Niven, Melanie Everett, and Sileshi Semaw. 2008. A female Homo Tishkoff, Sarah A., Floyd A. Reed, Franc¸oise R. Friedlaender, Christopher erectus pelvis from Gona, Ethiopia. Science 322:1089–1092. Ehret, Alessia Ranciaro, Alain Froment, Jibril B. Hirbo, et al. 2009. The Smith, Tanya M., and Paul Tafforeau. 2008. New visions of dental tissue genetic structure and history of Africans and African Americans. Science research: tooth development, chemistry, and structure. Evolutionary An- 324:1035–1044. thropology 17:213–226. Trauth, Martin H., Juan C. Larrasoan˜a, and Manfred Mudelsee. 2009. Trends, Smith, Tanya M., Paul Tafforeau, Donald J. Reid, Rainer Gru¨n, Stephen Eggins, rhythms and events in Plio-Pleistocene African climate. Quaternary Science Mohamed Boutakiout, and Jean-Jacques Hublin. 2007. Earliest evidence of Reviews 28:399–411. modern life history in North American early Homo sapiens. Proceedings of Trinkaus, Erik. 1983. The Shanidar Neandertals. New York: Academic Press. the National Academy of Sciences of the USA 104:6128–6133. ———. 2006. Modern human versus Neandertal evolutionary distinctiveness. Smith, Tanya M., Paul Tafforeau, Donald J. Reid, Joane Pouech, Vincent Current Anthropology 47:597–620. Lazzari, John P. Zermeno, Debbie Guatelli-Steinberg, et al. 2010. Dental Tryon, Christian A., and Sally McBrearty. 2006. Tephrostratigraphy of the evidence for ontogenetic differences between modern humans and Nean- Bedded Tuff Member (Kapthurin Formation, Kenya) and the nature of derthals. Proceedings of the National Academy of Sciences of the USA 107: archaeological change in the later Middle Pleistocene. Journal of Human 20923–20928. Evolution 65:492–507. Sørensen, Bent. 2011. Demography and the extinction of European Nean- Weaver, Timothy D. 2012. Did a discrete event 200,000–100,000 years ago derthals. Journal of Anthropological Archaeology 30:17–29. produce modern humans? Journal of Human Evolution 63:121–126. Stewart, J. R., and C. B. Stringer. 2012. Human evolution out of Africa: the role of refugia and climate change. Science 335:1317–1320. Weaver, Timothy D., Charles C. Roseman, and Chris B. Stringer. 2007. Were Stiner, Mary C. 2013. An unshakable Middle Paleolithic? trends versus con- Neandertal and modern human cranial differences produced by natural servatism in the predatory niche and their social ramifications. Current selection or genetic drift? Journal of Human Evolution 53:135–145. Anthropology 54(suppl. 8):S288–S304. ———. 2008. Close correspondence between quantitative- and molecular- Stringer, Chris. 2002. Modern human origins: progress and problems. Phil- genetic divergence times for Neandertals and modern humans. Proceedings osophical Transactions of the Royal Society of London B 357:563–579. of the National Academy of Sciences of the USA 105:4645–4649. ———. 2006. Homo britannicus: the incredible story of human life in Britain. White, Tim D., Berhane Asfaw, David DeGusta, Henry Gilbert, Gary D. Rich- London: Penguin. ards, Gen Suwa, and F. Clark Howell. 2003. Pleistocene Homo sapiens from ———. 2007. The origin and dispersal of Homo sapiens: our current state of Middle Awash, Ethiopia. Nature 423:742–747. knowledge. In Rethinking the human revolution: new behavioural and bio- Williams, Martin, Michael Talbot, Paul Aharon, Yassin Abdi Salaam, Frances logical perspectives on the origin and dispersal of modern humans. Paul Mel- Williams, and Knut Inge Brendeland. 2006. Abrupt return of the summer lars, Katie Boyle, Ofer Bar-Yosef, and Chris Stringer, eds. Pp. 15–20. Cam- monsoon 15,000 years ago: new supporting evidence from the lower White bridge: McDonald Institute for Archaeological Research. Nile Valley and Lake Albert. Quaternary Science Reviews 25:2651–2665. S234 Current Anthropology Volume 54, Supplement 8, December 2013

Variability in the Middle Stone Age of Eastern Africa

by Christian A. Tryon and J. Tyler Faith

CAϩ Online-Only Material: Supplement A

Eastern Africa is an important area to study early populations of Homo sapiens because subsets of those populations likely dispersed to Eurasia and subsequently throughout the globe during the Upper Pleistocene. The Middle Stone Age (MSA) archaeology of this region, particularly aspects of stone-tool technology and typology, is highly variable with only rare cases of geographic and temporal patterning. Although there are differences in timing and perhaps frequency of occurrence, those elements that make up the MSA lithic tool kit are also found at contemporaneous sites elsewhere in Africa and Eurasia, making it difficult to identify a unique archaeological signal for hominin dispersals out of eastern Africa. Rather, regional variation appears to be the outcome of possibly long-term interactions between particular physical and social environments experienced by hominin populations.

The archaeological record of eastern Africa has the potential Herto Member of the Bouri Formation (Clark et al. 2003; to play a central role in our understanding of the behavioral White et al. 2003), Aduma (Haile-Selassie, Asfaw, and White evolution of modern human populations. The fossil record 2004; Yellen et al. 2005), and Porc Epic (Assefa 2006; Clark from this region includes the earliest specimens attributed to et al. 1984; Pleurdeau 2004; Vallois 1951) in Ethiopia and Homo sapiens ∼195–154 ka (Clark et al. 2003; McDougall, from the Upper Ngaloba Beds at Laetoli (Day, Leakey, and Brown, and Fleagle 2005; White et al. 2003). Both fossil and Magori 1980), Mumba Rockshelter (Bra¨uer and Mehlman genetic evidence are consistent with this region providing the 1988; Mehlman 1989), and perhaps the Lake Eyasi Beds (cf. source population(s) for subsequent dispersals out of Africa Domı´nguez-Rodrigo et al. 2007, 2008; Mehlman 1987) in and feature prominently in models favoring the “southern Tanzania. We use the MSA archaeological record of eastern route” from Africa to Arabia (reviewed in Beyin 2011), with Africa to characterize the behaviors of hominin populations each dispersing group sampling a portion of the biological that included H. sapiens in addition to other archaic forms. and behavioral variability present in the parent population This allows comparison with neighboring regions to better (Gunz et al. 2009; Prugnolle, Manica, and Balloux 2005; cf. understand the pattern and timing of archaeological diversity Lycett and von Cramon-Taubadel 2008). Whereas the fossil and innovation during the Middle and Upper Pleistocene. and genetic data provide the best insights into past biological Put simply, is the archaeological record of eastern Africa con- variation, it is the archaeological record that provides the sistent with the biological evidence that this region is the richest source of information on the behavioral variability of source area for H. sapiens? fossil hominins. A few points of terminology require clarification. Following All of the early fossils of H. sapiens from eastern Africa are the International Commission on Stratigraphy, we refer to the associated with Middle Stone Age (MSA) artifacts. These in- time interval bounded by the Brunhes-Matuyama paleomag- clude those from the Kibish Formation (Brown and Fuller netic reversal dated to ∼781 ka and the onset of the Last 2008; McDougall, Brown, and Fleagle 2005; Shea 2008), the Interglacial at ∼126 ka as the Middle Pleistocene. The Upper Pleistocene lasts from the Last Interglacial until the Holocene Christian A. Tryon is Assistant Professor in the Department of at ∼11.7 ka. By the MSA, we follow the common usage (e.g., Anthropology at Harvard University (Peabody Museum of Clark 1988; Goodwin and Van Riet Lowe 1929; McBrearty Archaeology and Ethnology, 11 Divinity Avenue, Cambridge, and Brooks 2000) to refer to sites with lithic assemblages that Massachusetts 02138, U.S.A. [[email protected]]). J. are characterized by stone or bone points and the frequent Tyler Faith is Postdoctoral Fellow in the Archaeology Program at the School of Social Science of the University of Queensland use of Levallois methods for flake production. MSA sites lack (Brisbane, Queensland 4072, Australia [[email protected]]). This the large cutting tools such as cleavers and handaxes found paper was submitted 3 VII 13, accepted 20 VIII 13, and electronically in Acheulian (Early Stone Age [ESA]) sites. Backed pieces published 13 XII 13. may be present but are less common than at Later Stone Age

᭧ 2013 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved. 0011-3204/2013/54S8-0007$10.00. DOI: 10.1086/673752 Tryon and Faith Middle Stone Age Variability S235

(LSA) sites. Following McBrearty and Tryon (2006), the term ducted in Uganda and Kenya in the 1920s and 1930s (e.g., “early” MSA (EMSA) refers to sites that predate the Last Leakey 1931, 1936; O’Brien 1939; Wayland 1934; Wayland Interglacial. “Later” MSA (LMSA) sites date to within or after and Burkitt 1932). The Central Rift Valley of Kenya is an the Last Interglacial. Finally, “eastern Africa” refers to the important exception, where multiple MSA sites have been the modern-day countries of Tanzania, Kenya, Uganda, Ethiopia, focus of long-term study by Leakey (1931, 1936) and Isaac Somalia, Djibouti, and Eritrea. Geographically, this region is and his students (Isaac, Merrick, and Nelson 1972; Merrick bounded by the Indian Ocean, the Red Sea and a low elevation 1975), including Ambrose (1986, 2001, 2010) and others (e.g., coastal plain to the east, central highlands (11,000 m asl) Anthony 1972; Waweru 2007). Surprisingly, despite investi- dissected by the eastern arm of the Rift Valley, and the western gation since the 1930s and the presence of relevant archae- arm of the Rift to the west, with vegetation ranging from ological material (Leakey et al. 1972; Mabulla 1990), Olduvai desert scrub to Afro-alpine forest. The area encompasses ∼3.6 Gorge in Tanzania has played little role in our understanding million km2; for comparison this is similar in size to western of MSA sites in eastern Africa largely because of apathy on Europe or India and is a third the size of the Sahara desert the part of Mary Leakey (1984:213), the principal excavator: or the United States. “In Africa, the hand axe culture did eventually give place to Eastern Africa as used here is defined by modern political a surprisingly uninspiring group of industries lumped to- boundaries as a matter of convenience, but as shown in figure gether under the term Middle Stone Age; a stage in prehistoric 1 and discussed below, most sites discussed in this paper also archaeology for which I have never been able to feel any share a common environmental context, occurring within or enthusiasm.” We hope that our paper, together with the others near the boundaries of White’s (1983) Somali-Masai center reported in this volume, will serve to inspire new ideas and of regional endemism. We recognize that eastern Africa thus stimulate discussion concerning a time period that we believe provides a useful but imperfect geographic unit. Alternative is both interesting and important to understanding our evo- approaches could emphasize different boundaries, for ex- lutionary past. ample, comparing Red Sea coastal sites such as Abdur in Eritrea (Walter et al. 2000) with sites farther north and not Fossil Associations: The MSA Is Not included in this review such as Sodmein Cave in Egypt (Mer- Exclusive to Homo sapiens cier et al. 1999; Vermeersch et al. 1994). Sites are irregularly distributed throughout eastern Africa, largely concentrated within the East African Rift Valley system Although all of the early fossils of Homo sapiens are found (fig. 1), as a result of geological exposure. We do not attempt with MSA artifacts, it is unlikely that our species was the to comprehensively review every known eastern African MSA exclusive author of MSA lithic technology. On present evi- archaeological assemblage, but we emphasize those sites that dence, the oldest MSA sites in eastern Africa, at 1276 ka are sufficiently well published to determine the presence and (Morgan and Renne 2008), are at least 70 kyr older than the absence of archaeological attributes and have some degree of oldest known H. sapiens fossil. The early fossils of H. sapiens chronological or stratigraphic control (see also Basell 2008). and the populations they represent are highly variable, and Summaries of these sites and their age estimates are provided there is as yet no consensus on how to partition this variability in table 1, with their locations shown in figure 1. Stratigraphic (Gunz et al. 2009; Pearson 2008; Trinkaus 2005). Given the and chronological control is particularly important because possible presence of ancestral and sister taxa in the region overall site density is very low, with a complete lack of cov- (e.g., Hammer et al. 2011; Lachance et al. 2012), a more erage for many areas; radiometric dates are few, and there is cautious reading of the available evidence would be that the a rarity of caves or other deeply stratified sequences that allow variability among MSA sites likely encapsulates the behavioral ready observation of change through time. In this, eastern outcomes of multiple hominin populations of varying taxo- Africa is different from western Europe, China, the Levant, nomic affinities. Direct linkage between particular hominins or southern and northern Africa, and it relates largely to and specific archaeological entities is beyond the resolution bedrock geology (i.e., abundant lavas and few limestone or of our data, a problem that arises in other regions such as quartzite deposits). As a result, much of the eastern African the Levant (cf. Hovers 2009; Shea 2006a) and western Europe record consists of open-air sites used as living sites, hunting (cf. Slimak et al. 2011, 2012; Zwyns et al. 2012). localities, areas for obtaining stone raw material, and other functions (see, e.g., Tryon et al., forthcoming). Our inability The MSA: Origins and Endings to demonstrate contemporaneity among these sites in most cases diminishes our ability to distinguish variation due to The appearance and disappearance of MSA technologies can age as opposed to site function or environmental setting. both be defined as processes rather than events, typified by The irregular distribution of eastern African MSA sites is gradual, intermittent, and often complex patterns of change also reflected in the discontinuous nature of their investiga- with the loss of diagnostic ESA (Acheulian) or the addition tion (Gabel 1984; Robertshaw 1990). There have been few of LSA elements over time. As reviewed below, this pattern long-term projects in the region since the seminal work con- is consistent with technological change from existing, local Figure 1. Sketch map showing eastern African Middle Stone Age sites discussed in the text and major biogeographic zones of White (1983). Tryon and Faith Middle Stone Age Variability S237 antecedents and is in many ways comparable with the pattern of the desired Levallois flake blanks (Tryon, McBrearty, and seen in western Europe. There, late Acheulian sites contain Texier 2005). Finally, there is an apparent size gradient be- Levallois technology with the number of handaxes declining tween small (Acheulian) handaxes and large (MSA) points over time (Monnier 2006), and some Upper Pleistocene in- (McBrearty and Tryon 2006), consistent with a gradual shift dustries may show the complex, perhaps nonlinear appear- in artifact types (and perhaps functions) over time. Whether ance of backed elements (Bordes and Teyssandier 2012) or this size gradient masks different methods of production re- retouched points (Slimak 2008). mains uninvestigated.

The MSA Developed from Local Acheulian Antecedents The End of the MSA The overlap between age estimates for the earliest MSA and The end of the MSA was apparently a gradual but complex the latest Acheulian sites supports the hypothesis of a pro- process rather than an event, with the emergence of the sub- longed shift to MSA technologies. Gademotta, Ethiopia, is the sequent LSA developing from local MSA roots. At Enkapune oldest securely dated MSA site at 1276 ka (Morgan and Renne ya Muto, Kenya, the sequence from ∼40 to 55 ka shows a 2008; Wendorf and Schild 1974). The youngest reported basal MSA horizon with Levallois and discoidal methods of Acheulian artifacts are surface collected from the Herto Mem- flake production and rare backed pieces. It is overlain by an ber of the Bouri Formation of Ethiopia, dated to ∼154–160 industry attributed to the LSA dominated by the production ka (Clark et al. 2003), and in situ material perhaps as recent of large (∼7 cm) backed blades and microliths, which is in as ∼125 ka from Abdur, Eritrea (Bruggeman et al. 2004; Walter turn overlain by an industry with abundant microliths (∼2– et al. 2000). These young Acheulian sites are not without their 5 cm), MSA-like core reduction strategies, and ostrich eggshell problems. There remains the possibility that the Acheulian beads (Ambrose 1998). artifacts from the Herto Member are older than the dated In contrast, at Mumba Rockshelter, Tanzania, the strati- sediments by an unknown but possibly large interval. The graphic sequence suggests a gradual change in the frequency Acheulian attribution of the material from Abdur is unfor- of typological and technologically important artifacts. Backed tunately not supported by detailed artifact descriptions or elements persist in low numbers across multiple strata, co- illustrations. However, if accurate, these results suggest a incident with a reduction in the frequency of Levallois cores ∼100–150 kyr overlap between Acheulian and MSA technol- and points and an increased use of bipolar percussion for ogies in eastern Africa. flake production from ∼30 to 68 ka (Eren, Diez-Martin, and The Kapthurin Formation, Kenya, currently provides the Domı´nguez-Rodrigo 2013; Gliganic et al. 2011; Marks and best stratigraphic sequence showing the nature of the ap- Conard 2008; Mehlman 1989, 1991). The nature of the change pearance of MSA technologies within a single depositional is such that the MSA or LSA attribution of a number of basin (McBrearty and Tryon 2006; Tryon 2006; Tryon and industries at Mumba is uncertain (Diez-Martı´n et al. 2009). McBrearty 2006; Tryon, McBrearty, and Texier 2005) com- Similar combinations of typically MSA (e.g., points) and LSA plemented by recent and ongoing work at Olorgesailie, Kenya (e.g., backed pieces) artifacts are found at the Mochena Bo- (A. S. Brooks and J. E. Yellen, personal communication). In rago sequence from Ethiopia (Brandt et al. 2012). As these the Kapthurin Formation, sites with points and small Levallois sites show, the apparent continuity of backed pieces among cores are interstratified with those with cleavers, suggesting strata attributed to the MSA and LSA from eastern Africa that Acheulian and MSA technologies overlapped temporally sites is distinct from the discontinuous appearance of backed within the same geographic region (∼150 km2). Further, sev- pieces in southern Africa (cf. Howiesons Poort and Wilton eral elements of lithic technology found at MSA sites find assemblages) or the late appearance of backed pieces in north- their first expression in the Acheulian. These include the pro- ern Africa (e.g., Close 2002; Deacon and Deacon 1999; Villa duction of blades from cylindrical cores and particularly Le- et al. 2010; Wurz 2013). vallois methods of flake production from assemblages with handaxes and cleavers (Johnson and McBrearty 2010; Leakey MSA Lithic Technological Variability et al. 1969; McBrearty 1999; Tryon 2006). In the Kapthurin Formation and elsewhere in eastern Africa (reviewed in Stone tools and their manufacturing debris make up the bulk Sharon 2007; Tryon, McBrearty, and Texier 2005), Levallois of our evidence for studying hominin behavioral variability. technology formed one of several methods of producing Table 1 summarizes this variability for a number of key dated Acheulian large flake blanks that could be transformed into sites in the region. As detailed below and recently emphasized other tools. In each case, large (110 cm) Levallois flakes, often by Shea (2011b), the MSA record of lithic technology is highly with laterally retouched edges, were produced using the pref- variable from its first appearance. We first examine the var- erential method from centripetally prepared cores (fig. 2a). iation within each of the major artifact classes summarized Levallois cores and flakes at younger sites in the Kapthurin in table 1. Moving from particular artifact types to artifact Formation are smaller and show a greater diversity of Levallois aggregates, we then conduct more formal analyses of inter- approaches (detailed below), perhaps linked to size reduction assemblage variation to more rigorously test hypotheses of 8 6.1 5.6 4.1 2.1 2.7 2.0 5.5 4.9 2.4 7.3 7.3 6.2 3 7.8 3.5 1 2.2 4.7 13.8 10.5 10 pieces % retouched Misc. retouched tool Heavy-duty Ϫϩ ϩ large biface Other ? point Retouched Ϫϩϩϩ ϩ Ϫϩϩϩ ϩ ϩϩϩϩ ϩ Ϫϩϩϩ ϩ pieces Backed ? ? ? ? ϪϪϪ ϩ Ϫ Ϫ ϩ ϩϩϩ ϩ Ϫ ϩϩϩϩ ϩ ϩ Ϫ Ϫ ϩ ϩϩϩ ϩ Ϫ ϩϪϩϩ ϩ ϩ Ϫ Ϫ ϩ core Blades Bipolar ? ϪϪϪϪϪϪϩϪϩϪ ϩ ϩϩϪϩϩϩϩϩϩϪ ϩ ϪϩϩϩϩϪϩϩϩϩ ϩ ϩϪ ϩϪ ϩϪ Ϫϩ ϪϪϪϩϩϪϪϪϩϪ ϩ ϩϩϪϪϪϩϩ ϩ ϪϪϪϩϪϪϪϪϪϪ ϩ core point/ Levallois ? ?? ?? ? ? ? ? ? ? ?? ?? ? ? ? Ϫ ϪϩϪϪϩ ϪϪϪϩϩϪϪϪ core Levallois recurrent ? ? core Levallois preferential core Discoidal Ϫ ϩ ϪϩϩϪ ϩ ϩϪ ϩ ϪϩϩϩϩϩϪϪ ϩ ϩϩ ϩ ϪϩϪϩϩϩϪϪ ϩϩ ϩ core Platform ? ? ? ϪϪ ϩ ϩ ϩ ϪϪ ϩ Ϫ ϩ ϪϪ ϩ Ϫ Ϫ ϪϪ ϩ ϩ ϩ ϪϪϪ Ϫ ϩ Ϫ ϩϪϪ ϩϪϪ ϩϪϪ ϩϩϩ ϩ ϩϩϩϩ ϩ ϩ ϩ ϩϩϪϩ ϩ Ϫ ϩ ϩ ϩ ϩ ϩϩϪ ϩ Ϫ ϩ ϩ ϩ ϩ Ϫ ϩ ϩ ϩ Ϫ Ϫ ϩ ϩϪϪ Ϫ ϩ ϩ ϩ ϪϪϪ Ϫ ϩ ϩ ϩ ϩ ϩ Ϫ ϩ Ϫ ϩ Ϫ Ϫ ϩ ϪϪϩ ϩ ϩϪϪϩ ϩ Ϫ ϩ ϩϪϪϩ ϩ Ϫ ϩ ϩ ϩ ϩ ϩ ϩ ϩ ϩ ϩ ϩ Ϫ ϩ ϩ ϪϪϩ Ϫ ϩ ϩϪϪϩ ϩ Ϫ ϩ ϩϪϪϩ ϩ Ϫ ϩ ϩ ϩ ϩ Ϫ ϩ Ϫ ϪϪϪ Ϫ ϩ ϩ ϪϪϪ Ϫ ϩ ϩ ϩ a a a 40 ? ? 100 1 68 125? Ϫ 1 18–26 18–26 26–55 88–132 ! 10–170 10–170 10–170 1 1 1 1 1 1 ≥ 0.48 36.1 30–72 0.42 34.17 38 3.53 35.32 30–40 3.53 35.32 50–60 3.53 35.32 60–68 3.53 35.32 3.52 35.17 2.74 35.36 2.74 35.36 4.08 39.13 2.74 35.36 Ϫ Ϫ Ϫ Ϫ Ϫ Ϫ Ϫ Ϫ Ϫ Ϫ Ϫ EMSA Lat. Long. Age (ka) Ochre Beads Anvil Grindstone LMSA/ LMSA 10.95 47.37 19– LMSA LMSA LMSA LMSA 10.95 47.37 40–90 ? ? LMSA 10.95LMSA 47.37 10.95 40–90 47.37 40–90 ? ? ? ? LMSA a a a 25 (Kisele Ϫ (GrJi11)tion, Member 4, Pundo LMSA Mak- war Industry LMSA? 0.08 34.63 Rusinga Island LMSA S-groupUpper T-group LMSA LMSALower T-group 6.90 6.90 LMSA 37.76 37.76III 6.90 43–45 37.76 43 45–53 (Mumba In- dustry)A (Kisele In- dustry) LMSA (Sanzako In- dustry) tion, Member 2, Ojalla In- dustry EMSA? 0.08 34.63 IV V VI lower (Nasera Industry) 11 (Mumba Industry)12 LMSA Industry) LMSA tion, Member 6, Pundo Mak- war Industry LMSA? 0.08 34.63 ity KY-55 East, Upper Group LMSA? (Nasera Indus- try) Prolonged Drift Muguruk Forma- Wasirya Beds, Mochena Borago Mochena Borago Mochena Borago Midhishi 2 LSU Mumba Bed V Mumba Bed VI- Mumba Bed VI-B Muguruk Forma- Midhishi 2 LSU Midhishi 2 LSU Midhishi 2 LSU Porc Epic LMSA 9.63 41.87 33–79 Eyasi Beds EMSA? Table 1. Artifact variability at eastern African Middle Stone AgeSite/evidence sites Mumba Bed III Nasera levels 8/9– Nasera levels Muguruk Forma- Mtongwe, Local- Nasera levels 6–7

S238 0 0 3.1 1 2 1.8 1 5.8 3.4 4.1 1.6 2.2 3.8 1.8 1.6 ϩϪ Ϫ ϩ ϩϪ Ϫ ϩ ? ? ϩϪϪϩ ϩ ? Ϫ ? ϪϪϩϪϩϪϩϪϩϩ ϩ ϪϩϪϪϪϩϩ ϩ ϪϪϪϩϪϩϩ ϩ ϪϩϪϪϪϩϩ ϩ ϪϩϪϪϪϩϩ ϩ ϩϩϪϪϪϩϩϩϪϪ ϩ ϪϪϪϪϪϪϩϪϩϪ ϩ ϪϪϪϩϪϩϪ ϩ ? ? ? ? ? ? ? 126 ka boundary between the EMSA/LMSA, or where biostratigraphic data differ from radiometric age estimates. ?? ∼ ϩϩϩϩ uncertain. See text for definitions and references. Multiple assemblages from a single site shown in stratigraphic order. Sites are attributed to Early MSA p ? absent; ? p ϪϪ Ϫ Ϫ ϩ ϩ Ϫ ϪϪϪ Ϫ ϩ ϩ ϩ ϩ Ϫ Ϫ Ϫ Ϫ Ϫ Ϫ ϩ Ϫ ϪϪϪ ϩ ϩϪϪϪ ϩ Ϫ ϩ ϪϪϪϪ ϩ Ϫ ϩ ϩ ϩ ϩϪϪϪ ϩ Ϫ ϩ ϩ ϩϩϪϪ ϩ Ϫ ϩ ϩ ϩ ϩ ϩ Ϫ ϩ Ϫ Ϫ Ϫ Ϫ Ϫ ϪϪϪ Ϫ ϩϪϪϪ ϩ Ϫ ϩϪϪ ϩ ϩ ϩ Ϫ Ϫ ϪϪϪ Ϫ ϩ ϩϪϪϪ ϩ Ϫ ϩ Ϫ ϩ ϩϪϪϪ Ϫ Ϫ ϪϪϪϪ ϩ Ϫ ϩ Ϫ ϩ ϩ ϩ ϩ ϩ Ϫ Ϫ ϩ Ϫ Ϫ Ϫ ϩ ϩ ϪϪϪ Ϫ ϩ Ϫ ϩ ϩ ϩ Ϫ ϩ Ϫ ϩ ϩ Ϫ ϩ present; p ϩ 183 183 183 276 125? 125? ! ! 1 ∼ ! ! 4.08 39.13 4.08 39.13 Ϫ Ϫ pressed as decimal degrees. mation (ETH72-5) EMSA 7.94 38.6 mation, site Koimilot Locus 1 EMSA 0.52 35.9 200–250 ity KY-55 East, Middle Group LMSA? uma sites A4, A8, A8A, VP 1/ 1, VP 1/3(base), Aduma site A1 LMSA 10.42tion, Member 40.52II/III, site BNS 80–100 LMSA LMSA 10.42 40.52 5.41 35.9 80–100 tion, Member 98–123 I, site KHSmation, site EMSAKoimilot Locus 2 5.41 35.9 195–204 EMSA 0.52 35.9 200–250 ity KY-55 East, Lower Group LMSA? contact, Ad- uma site A5 LMSA 10.42 40.52 80–100 tion, Member I, site AHS EMSA 5.41 35.9 195–204 ? mation (ETH72-6, ETH72-9) EMSA 7.94 38.6 mation (ETH72-7B, ETH72-1)mation EMSA(ETH72-8B) 7.94 EMSA 38.6 7.94 38.6 Data from 1938 Kohl-Larsen collections rather than those of Mehlman (1989); these are likely overestimates. Gademotta For- Kapthurin For- Mtongwe, Local- Note. Artifact variability at eastern African sites: a Ardu Beds B, Ad- Ardu Beds B Kibish Forma- Kibish Forma- Kapthurin For- Mtongwe, Local- Ardu Beds B/C Kibish Forma- Gademotta For- Kapedo Tuffs EMSA? 1.07 36.08 123–135 Gademotta For- Gademotta For- (EMSA) or Later MSA (LMSA) based on radiometric dates; uncertainties arise for undated sites, those that span the Latitude and longitude ex

S239 Figure 2. Selected cores and flakes from eastern African Middle Stone Age sites. a, Preferential Levallois core from the Acheulian Factory Site, Kapthurin Formation, Kenya. b, Preferential Levallois core from Gademotta, Ethiopia. c, Recurrent centripetal Levallois core from the Kapedo Tuffs, Kenya. d, Nubian core from the Wasiriya Beds, Rusinga Island, Kenya. e, Recurrent bipolar Levallois core from Nyogonyek, Kapthurin Formation, Kenya. f, Levallois flake from K’one, Ethiopia. g, h, Levallois points from Koimilot, Kapthurin Formation, Kenya, and Midhishi 2, Somalia. i, Discoidal core from the Wasiriya Beds, Rusinga Island, Kenya. j, Blade core from the Kapedo Tuffs, Kenya. k, Single-platform core from Nasera Rockshelter, Tanzania. l, Bipolar core from Nasera Rockshelter, Tanzania. Artifact illustrations after Gresham (1984); Kurashina (1978); Mehlman (1989); Tryon (2003); Tryon, McBrearty, and Texier (2005); Tryon, Roach, and Logan (2008); Tryon et al. (2012); and Wendorf and Schild (1974). Note that k and l use the lower scale bar; all others use the upper scale bar. Tryon and Faith Middle Stone Age Variability S241 temporal and geographic variation within the eastern African Nile Valley and to the east in parts of the Arabian Peninsula MSA. (see Rose et al. 2011; Van Peer 1992, 1998) and provide some of the strongest archaeological evidence for connections be- tween Africa and Arabia in the Upper Pleistocene. Nubian Levallois Technology Is Highly Variable Type 1 cores are found in eastern Africa at sites at or near Levallois technology in eastern Africa is as highly variable as the margins of the Nile drainage basin (fig. 2d), including it is in other well-studied regions such as western Europe or K’one and Aduma in Ethiopia (Kurashina 1978; Yellen et al. the Levant (e.g., Delagnes and Meignen 2006; Hovers 2009). 2005) and Rusinga Island in Kenya (Tryon et al. 2012), doc- Although Levallois cores and flakes have been reported from umenting an extensive range for this Levallois variant. Some eastern African sites since at least the 1930s (e.g., Leakey Levallois approaches, such as that used for blade production 1936), few sites in the region have been analyzed using the at some South African sites (Wurz 2002, 2013), are not found reconfigured understanding of the Levallois concept most in eastern Africa. strongly associated with the work of Eric Boe¨da (1994, 1995). Briefly, the Levallois concept is an approach to flake produc- Beyond Levallois: Other Flake Production Methods tion that targets the preparation and subsequent reduction of a single core surface for the removal of relatively large and Although Levallois technology is a critical part of our un- thick flake blanks (Eren and Lycett 2012). From this Levallois derstanding of MSA sites, other forms of flake production flake removal surface, a single Levallois flake, blade, or point persist, and additional, non-Levallois methods were intro- is removed before repreparation of the convexities of the core duced (table 1). Discoidal and single- and multiple-platform (the preferential method), or multiple flakes, blades, or points cores are widespread at MSA sites (fig. 2i,2k). Discoidal cores are removed before repreparation (the recurrent method). result from the alternate flaking of both sides of the periphery The Levallois flake removal surface is shaped by the removal of (typically) an oval cobble, resulting in a bifacially flaked, of “preparatory” flakes that alter core convexities. These con- biconical core. Platform cores (including the “migrating plane vexities control the fracture pattern that in part dictates the cores” of White and Pettitt 1995) result from the use of one form of the Levallois flake(s) removed from that surface (Van or more edges as striking platforms. Discoidal and platform Peer 1992). Levallois flakes and preparatory flakes may be cores are found at ESA (both Oldowan and Acheulian) sites removed using a number of different patterns, including re- and form a technological substrate for the production of movals from one direction (unidirectional), from opposite sharp-edged flakes in some LSA assemblages (e.g., Mehlman ends (bidirectional), about the circumference of the core (cen- 1989). Bipolar cores (fig. 2l) resulting from the production tripetal), or subtle variations on these major themes (e.g., of small flakes using an anvil are also known from ESA (Ol- unidirectional convergent flaking). dowan) sites (e.g., de la Torre 2004). Bipolar cores occur The combination of the particular Levallois method (pref- irregularly at MSA sites in eastern Africa, including Nasera erential or recurrent) and the flake removal patterns (e.g., and Mumba rockshelters in Tanzania (Diez-Martı´n et al. 2009; unidirectional, bidirectional, centripetal) combine to produce Eren, Diez-Martin, and Domı´nguez-Rodrigo 2013; Mehlman substantial variability within the Levallois approach to flake 1989) and Cartwright’s site in Kenya (Waweru 2007). Blade production. Many of these variants are expressed at multiple or bladelet production (fig. 2j) occurs at EMSA assemblages MSA sites in eastern Africa (fig. 2b–2h). Importantly, variable at Gademotta/Kulkuletti (Wendorf and Schild 1974) and approaches to Levallois flake production are present at EMSA LMSA assemblages at Aduma and Porc Epic in Ethiopia sites such as the Kapthurin Formation, Kenya (Tryon 2003, (Pleurdeau 2004; Yellen et al. 2005) and elsewhere. As noted 2006) and Gademotta/Kulkuletti, Ethiopia (Douze 2008; previously, blades also occur in ESA (Acheulian) assemblages Wendorf and Schild 1974), as well as at LMSA sites, including in eastern Africa. Truncated-facetted pieces used for the pro- Aduma (Yellen et al. 2005) and Porc Epic (Pleurdeau 2004) duction of small flakes are common at some Levantine and in Ethiopia and Rusinga Island in Kenya (Tryon et al. 2012). European Middle Paleolithic sites (papers in McPherron Levallois points (fig. 2g,2h) are present at several eastern 2007) and some eastern African LSA sites (e.g., Newcomer African sites where they make up from ∼8% to 62% of re- and Hivernel-Guerre 1974). Although rare and probably un- covered Levallois flakes, such as Koimilot in the Kapthurin derreported from eastern Africa, truncated-facetted pieces are Formation, Kenya (Tryon 2006); the Bird’s Nest Site (BNS) reported from the MSA at Gademotta/Kulkuletti (Wendorf and Awoke’s Hominid Site (AHS) from the Kibish Formation, and Schild 1974:89); K’one Locality 5, Ethiopia (Kurashina Ethiopia (Shea 2008); and Midhishi 2 in Somalia (Brandt and 1978); and perhaps Lukenya Hill (Clark 1988) and Prolonged Gresham 1989; Gresham 1984). This frequency is within the Drift (Merrick 1975), Kenya. range of but is often greater than that found at some Levantine sites (cf. Hauck 2011; Hovers 2009:217). The Nubian Type 1 Points: Functional, Spatial, and Temporal Variability method is a Levallois point variant distinguished by two elon- gated preparatory flakes removed from the distal end of the Along with the frequent use of Levallois technology, points core. Nubian Type 1 cores are prevalent to the north in the are a defining element of the MSA. Point forms at eastern S242 Current Anthropology Volume 54, Supplement 8, December 2013

African sites are highly variable in size and shape (fig. 3a– phologically highly variable and are not attributed to named 3d). “Point” refers to a broad category of artifacts made of larger archaeological entities equivalent to the Lupemban. stone including unretouched, unifacial, and bifacial imple- While the small, subtriangular bifacially flaked forms (fig. 3b– ments made on Levallois and other flake blanks. The term is 3d) are distinct from lanceolate points or tanged pieces from both a morphological description (a pointed artifact) and an Lupemban or Aterian sites, similar forms occur as far west ethnographically based functional inference (as the tip of a as Mali (Soriano et al. 2010) and as far south as Botswana spear or other hunting implement). Studies of point shape, (Coulson, Staurset, and Walker 2011), reducing their utility microwear patterns, and mastic traces from sites in eastern as a unique regional artifact form. Africa and adjacent areas indicate that many points were hafted and probably used to tip spears, darts, or even arrows Other Shaped Tools (e.g., Brooks et al. 2006; Donahue, Murphy, and Robbins 2002–2004; Shea, 2006b; Van Peer, Rots, and Vermeersch Although used to define MSA technology, points and indeed 2008; Waweru 2007). all forms of shaped or retouched tools are rare at MSA sites, We cannot assume that all points were used as armatures typically making up !5%–7% of the flaked artifact total (table or projectiles. Gademotta is the only eastern African MSA site 1). This is also true of many southern African MSA sites (e.g., subjected to two independent analyses of artifact function Thackeray 1989), and the rarity of retouch has made it dif- (Douze 2008; Wendorf and Schild 1993). Both analyses found ficult to directly apply typologies such as that of Bordes (1961) that typologically defined points were used as cutting tools that emphasize shaped or modified tools (see discussion in rather than as spear tips, serving as an important reminder Villa, Delagnes, and Wadley 2005). In some areas, retouch about the potential dangers of inferring stone-tool function frequency is directly linked to the presence of fine-grained from artifact form. Villa and Lenoir (2006; see also Villa, raw materials, with retouch being rare on lava artifacts but Delagnes, and Wadley 2005) have shown almost the opposite more common on those made of chert or similar rocks for the European Middle Paleolithic record, where some “con- (Tryon, Roach, and Logan 2008). The Bordes system has been vergent scrapers” show impact damage consistent with their successfully applied to sites in northern Africa where chert is use as the tips of thrusting spears. Microwear analyses of widespread (e.g., Hublin, Tillier, and Tixier 1987) and in east- Levantine Levallois points emphasize the diversity of cutting ern Africa to sites such as Gademotta/Kulkuletti, Ethiopia tasks served by these tools in addition to their possible use (Wendorf and Schild 1974), where obsidian was used nearly as spear tips (Beyries and Plisson 1998). exclusively. Despite the relative rarity of retouched imple- Whatever their function, for retouched pieces, EMSA av- ments, three tool classes are important to understanding MSA mm,n p 50 ) is significantly lithic technology: heavy-duty tools, scrapers, and backed 16.40 ע erage point length (54.25 .mm,n p 250 ; pieces 14.30 ע larger than LMSA points (42.22 Mann-Whitney U-test:z p 5.222 ,P ! .001 ; table A1 in CAϩ Several MSA sites have “heavy-duty tools” (sensu Clark online supplement A). These size differences may reflect 2001b) such as picks (fig. 3h). These tools are also found in changes in tool function, including the evolution of complex Acheulian or other earlier regional industries or industrial projectile technology, which may appear at LMSA sites ∼40– complexes such as the Sangoan and are likely a retention of 100 ka (Brooks et al. 2006; Shea 2006b). Some stratigraphic characteristic ESA technologies (table 1). These MSA sites sequences such as Aduma and Nasera show a monotonic include Koimilot in the Kapthurin Formation (Tryon 2006) decrease in point size over time, whereas Mumba, Gademotta/ and the Kapedo Tuffs of Kenya (Tryon, Roach, and Logan Kulkuletti, and Gorgora rockshelters (Leakey 1943) do not 2008), assemblages from Kibish Formation of Ethiopia (sur- (table A1). face collected and not from the localities listed in table 1; Clark (1993) and McBrearty and Brooks (2000) have em- Shea 2008), and the 168–130 ka basal Bed VI at Mumba phasized geographic variation among MSA points at the sub- Rockshelter, Tanzania (Gliganic et al. 2011; Mehlman 1989: continental scale, although formal definitions or tests of the 194). Similar tools also occur at the undated sites of Muguruk extent of many of these variants remain to be done. The (McBrearty 1988), FxJi 61 near Koobi Fora (Kelly 1996:159), Lupemban is one of the most distinct MSA regional variants, and the basal MSA levels at Mtongwe in Kenya (Omi 1986, characterized by large (110 cm), thin, bifacially flaked lan- 1988). ceolate points (fig. 3a). Originally defined from sites in central As a tool class, scrapers have been reported from some of Africa, Lupemban lanceolates are found as far east as the Lake the earliest archaeological sites. However, in a qualitative Victoria region of Kenya. Although poorly dated in eastern sense, most scrapers from African Oldowan, Acheulian, and Africa, the large size of Lupemban lanceolates suggests attri- many MSA sites are characterized by rare and irregular re- bution to the EMSA, consistent with U-series age estimates touch. These are very different, for example, from the classic of 170–270 ka for Lupemban assemblages in Zambia (Barham scrapers defined by Bordes. It is only at MSA sites that some 2000) and the 110–170 ka age estimate from sedimentation scraper forms appear that show continuously retouched edges rates published by McBrearty (1988) for western Kenya. Other used to alter the shape of the tool (fig. 3f), a difference in points from eastern African MSA sites are smaller but mor- form that may result from extending the use-life of hafted Tryon and Faith Middle Stone Age Variability S243

Figure 3. Selected tools and beads from eastern African Middle Stone Age sites. a, Lupemban lanceolate point from Muguruk, Kenya. b, Point with basal thinning from Nasera Rockshelter, Tanzania. c, Point from Porc Epic, Ethiopia. d, Point from BNS, Kibish Formation, Ethiopia. e, Point with resharpening flake, Gademotta, Ethiopia. f, Scraper from Gademotta, Ethiopia. g, Grindstone fragment from Mumba Rockshelter, Ethiopia. h, Pick from Kapedo Tuffs, Kenya. i, Ostrich eggshell beads and production fragments from Mumba Rockshelter, Tanzania. j–m, Backed pieces from Mumba Rockshelter, Tanzania (j, k); Mtongwe, Kenya; and Enkapune ya Muto, Kenya. Artifact illustrations after Ambrose (1998); Clark et al. (1984); McBrearty (1986); Mehlman (1989); Omi (1986); Shea (2008); Wendorf and Schild (1974). Note that a through h use the upper scale bar; i through m use the lower scale bar. implements through resharpening (cf. Keeley 1982), suggested Comparisons with historical examples (Clark, Phillips, and in particular by distinctive resharpening flakes found at Gad- Staley 1974), experimental work (Clark and Prince 1978), and emotta/Kulkuletti (Wendorf and Schild 1974; fig. 3e). rare traces of ochre likely used as mastic (Ambrose 1998) Backed pieces first appear at LMSA sites in eastern Africa. suggest that backing is performed to facilitate hafting into a Backed pieces are flakes or blades with one lateral edge made slotted wooden shaft consistent with findings from sites else- steep or blunted (“backed”) by abrasion or direct percussion. where (e.g., Villa and Soriano 2010). In eastern Africa, backed S244 Current Anthropology Volume 54, Supplement 8, December 2013 pieces first appear ∼120–130 ka at the site of Deighton’s Cliff, abundant, direct evidence of eastern African hominin diet is Kenya (Ambrose and Deino 2010) but are a more regular sparse. Plants likely made up the bulk of the diet of any feature at LMSA sites ≤80 ka (fig. 3j–3m; table 1), including hominin population living at or near the equator (Kelly 1995), Porc Epic (Clark et al. 1984; Pleurdeau 2004), Mochena Bo- but at present there is no direct evidence for plant con- rago (Brandt et al. 2012), and Mumba and Nasera rockshelters sumption from eastern African MSA sites. As Marean (1997) (Mehlman 1989, 1991). Backed pieces are also present in small notes, our models of reconstructing past foraging systems in quantities from the !55 ka basal layers at Enkapune ya Muto the region are limited by the lack of modern or historic for- (Ambrose 1998), with undated examples from probable MSA agers (rather than pastoralists) from tropical grasslands, an strata at Mtongwe, Kenya (Omi 1984, 1986, 1988), and Kisese environment that characterizes much of eastern Africa now II Rockshelter, Tanzania (Inskeep 1962; Mehlman 1989:365), and in the Pleistocene. although published details from the latter site are scant. Site location and faunal data provide two alternative means of investigation. Sites such as Porc Epic, Ethiopia, and Nasera, Ochre and Grindstones Tanzania, have been interpreted as overlook sites situated near game pathways (Clark 2001a; Mehlman 1989). Hunters ap- Ochre (or other mineral pigments) and grindstones (fig. 3g) pear to have used natural features such as topographic lows, are two key elements of MSA lithic technology. These two streams, or springs to acquire game at site GvJm46 at Lukenya artifact classes often co-occur (table 1), suggesting a functional Hill (Marean 1990; 1997; Miller 1979) and Rusinga Island association. Ochre staining has been reported from some (Jenkins et al. 2012; Tryon et al. 2010) in Kenya. Ambrose grindstones, including those from the Kapthurin Formation (2001) has argued that MSA populations in the central part and Enkapune ya Muto, in Kenya (Ambrose 1998; McBrearty of the Rift Valley in Kenya positioned themselves at or near and Brooks 2000). The Kapthurin Formation example (from the ecotone between grassland and forest habitats in order to site GnJh-15) is the earliest reported occurrence of grind- best access resources from both environments. The use of ∼ stones and ochre from eastern Africa, dated to 284–500 ka coastal environments is demonstrated by MSA artifacts em- and associated with a lithic assemblage that cannot be con- bedded within an ∼125 ka coastal reef off the coast of Eritrea fidently attributed to the Acheulian or MSA. Otherwise, grind- (Walter et al. 2000) and an undated but well-stratified MSA stones and ochre are found at the LMSA sites (table 1) of artifact sequence in coastal dune sands at Mtongwe, Kenya Aduma (Yellen et al. 2005) and Porc Epic (Clark et al. 1984) (Omi 1984, 1986, 1988, 1991). Despite the importance of in Ethiopia; Mumba and Nasera rockshelters in Tanzania coastal environments for many out-of-Africa dispersal sce- (Mehlman 1989); and Enkapune ya Muto in Kenya (Ambrose narios (e.g., Bulbeck 2007), these two sites provide the only, 1998). albeit sparse, evidence for use of these environments from In addition to working ochre, grindstones may have also eastern African MSA sites. been used to process seeds or other plant material, an activity The faunal assemblages from GvJm46 at Lukenya Hill with considerably less archaeological visibility. Mercader (Marean 1990) and Porc Epic (Assefa 2006) provide the only (2009) reports starch grains from grindstones and other tools large, well-studied, and published archaeofaunal MSA assem- from Ngalue, Mozambique, suggesting grass seed processing blages from eastern Africa. Human exploitation of large mam- ∼ 105 ka. The extent to which these results can be applied to mals is also documented at other sites, including Rusinga grindstones at eastern African MSA sites is unknown but Island, Kenya (Jenkins et al. 2012; Tryon et al. 2010), and should be a focus of future research. As emphasized by Kuhn Loiyangalani, Tanzania (Thompson 2005). Although the sam- and Stiner (2001), the appearance of grindstones for seed ple is small, these studies suggest that at least by the later processing in MSA sites implies a shift toward lower-return parts of the Pleistocene, MSA foragers hunted a variety of foodstuffs that require substantial energy investment and thus large and small ungulates and selectively transported meat- a change in the foraging ecology of hominin populations. rich elements to central places such as caves for further pro- cessing and consumption. Long-distance carcass transport to Other Behaviors central places may distinguish MSA foraging strategies from We synthesize three other attributes of MSA hominin pop- those documented at other ESA sites in East Africa (e.g., Faith, ulations in eastern Africa: foraging behavior, territorial range Domı´nguez-Rodrigo, and Gordon 2009). inferred from raw material treatments, and symbolic behavior. Specifically, symbolic behavior concerns the treatment of the Territory and Movement Inferred from Raw Material dead and the use of ornaments. Transport Data Site-to-source distances for stone raw material provide the Foraging Behavior best empirical estimate of the size of the physical and social As many stone tools served either directly or indirectly in the landscapes familiar to early hominin populations. Compared food quest, we expect shifts in hominin diet to be reflected with ESA hominins, groups making MSA artifacts used finer- in lithic assemblage composition. Although stone tools are grained rocks, particularly obsidian, more frequently (Fe´blot- Tryon and Faith Middle Stone Age Variability S245

Augustins 1990; Merrick, Brown, and Nash, 1994). An ex- Gliganic et al. (2011). Mehlman (1989) considered Bed V and tensive, ongoing program to geochemically characterize Rift lower Bed III to contain industries intermediate between the Valley obsidian sources and artifacts provides the most de- MSA and LSA, whereas Diez-Martı´n et al. (2009) and Eren, tailed raw material transfer data (Ambrose et al. 2012; Mer- Diez-Martin, and Domı´nguez-Rodrigo (2013) ascribe these rick, Brown, and Nash 1994; Negash, Brown, and Nash 2011), layers to the LSA, making the association of these beads with summarized in table A2 in CAϩ online supplement A). ESA the MSA uncertain. Conard (2004) reports directly dated site-to-source distances are !60 km, whereas eastern African ∼29–33 ka ostrich eggshell beads from MSA/LSA strata at MSA site-to-source distances exceed 300 km. This increase Mumba. At Enkapune ya Muto, ostrich eggshell beads date suggests expanded physical and social landscapes through to ∼40 ka from an MSA/LSA stratum that overlies an industry which stone artifacts were carried by highly mobile foragers attributed to the LSA (Ambrose 1998). It is unclear whether and/or transferred through exchange. examples from 130 ka at Kisese II Rockshelter should be MSA hominins apparently regularly transported obsidian attributed to the MSA or LSA (Inskeep 1962; Leakey 1983: cores, flakes, and finished tools ≤30 km, and in the case of 21). Porc Epic, 139 km (table A2). Beyond this and up to a distance In short, there are few demonstrable examples of bead use of 305 km, obsidian frequency declines, and only finished by MSA hominins in eastern Africa, and the behavior is a tools and (resharpening?) flakes are found. These differences relatively late phenomenon. The appearance of beads marks may reflect shifts in raw material procurement strategies one of the few apparent sharp breaks in the MSA record. driven by increased source distance relative to a group’s ter- Whether this is due to the appearance of a neural mutation ritorial range (from provisioning of places to provisioning of that led to the development of language and modern human individuals; Kuhn 2004) or perhaps the trade/exchange of behavior (Klein 2009), a shift to more durable forms of per- finished pieces rather than cores among different groups. Al- sonal expression (as suggested by Kuhn and Stiner 2007), or though sample size is small, sites at a similar distance (∼130– sampling bias due to a small sample of caves or rockshelters 200 km) from the nearest source within the Rift Valley (Porc is unclear. Rare, well-preserved Holocene burials such as Epic) and outside of Rift Valley to the west in the Lake Victoria Njoro River Cave, Kenya (Leakey and Leakey 1950), are also region (Songhor and Muguruk) show very different patterns powerful reminders of the widespread use of seed beads or (table A2). Obsidian is more rare (8% vs. !0.5%) and limited other perishable materials unlikely to preserve at MSA sites, to tools and flakes in the Lake Victoria region. This may result potentially exaggerating the importance of the use of ostrich from the relative difficulty of movement across the steep, often eggshell as a medium for bead production. densely vegetated margins of the Rift rather than along the grassy, open valley floor, suggesting a possible biogeographic Interassemblage Variability control to hominin movement within the region. Similarly, in central Europe, open habitats are associated with greater To explore the nature of interassemblage variability among stone transport distances (≤300 km) than in the topograph- eastern African MSA sites, we use presence/absence data for ically complex region of western Europe (Fe´blot-Augustins artifact classes (listed in table 1). Although variable artifact 1993). typologies in use among researchers can reduce the utility of such approaches (Vermeersch 2001), the categories used here are sufficiently broad to minimize this problem. The data are Symbolic Behavior: Mortuary Practices and Ornaments used to examine (1) temporal variability among EMSA and The 154–167 ka hominin skulls from the Herto Member of LMSA sites (e.g., Shea 2011b), (2) geographic variability across the Bouri Formation, Ethiopia, represent the only example of eastern African sites (Clark 1988; McBrearty and Brooks peri- or postmortem treatment of the dead found at eastern 2000), and (3) local, site-specific sources of variation. In the African MSA sites. Here, cut and polish marks on the skulls following analyses, we conservatively treat all artifact classes have been interpreted as evidence of mortuary practice (Clark whose presence is uncertain (those with a question mark in et al. 2003; White et al. 2003). The defleshed hominin skull table 1) as absent. from the ∼500 ka Acheulian site of Bodo (White 1986) lies A correspondence analysis illustrating the association of only 30 km away, possibly indicating significant time depth different MSA assemblages with different artifact classes (fig. for similar behaviors in the region. 4, top) reveals temporal patterning among EMSA and LMSA Beads are the earliest direct evidence for personal orna- assemblages. The EMSA artifact assemblages overlap in mul- mentation from eastern African sites (fig. 3i). At Porc Epic, tivariate space with many of the LMSA assemblages, but there gastropod opercula were arguably worn as beads, co-occur is a subset of LMSA assemblages that are distinct (Axis 1 with MSA artifacts, and have been directly dated by the AMS scores 1 0). These include the LMSA assemblages dated to radiocarbon method to between ∼33 and 143 ka (Assefa, Lam, !75 ka from Mumba, Nasera Rockshelter, Porc Epic Cave, and Mienis 2008). Ostrich eggshell beads and manufacturing Mochena Borago, and the undated middle and upper assem- debris have been reported from Bed V and lower Bed III at blages from Mtongwe, which differ from EMSA sites by the Mumba Rockshelter, Tanzania, now dated to 30–60 ka by more frequent presence of beads, ochre, backed pieces, bipolar Figure 4. Correspondence analysis of artifact assemblage composition (top; table 1) across MSA localities. Site abbreviations: MuIII p Mumba Bed III; MuV p Mumba Bed V; MuVI-A p Mumba Bed VI-A; MuVI-B p Mumba Bed VI-B; Ey p Eyasi Beds; NaN p Nasera (Nasera Industry); NaM p Nasera (Mumba Industry); NaKi p Nasera (Kisele Industry); Ru p Wasiriya Beds; MiIII p Midhishi 2 LSU III; MiIV p Midhishi 2 LSU IV; MiV p Midhishi 2 LSU V; MiVI p Midhishi 2 LSU VI; PE p Porc Epic; PD p Prolonged Drift; MtU p Mtongwe Upper Group; MtM p Mtongwe Middle Group; MtL p Mtongwe Lower Group; Ar1 p Ardu Beds B/C; Ar2 p Ardu Beds B; Ar3 p Ardu Beds B (base); KiBNS p Kibish site BNS; KiAHS p Kibish site AHS; KiKHS p Kibish site KHS; Ka1 p Kapthurin Formation, Koimilot Locus 1; Ka2 p Kapthurin Formation, Koimilot Locus 2; Ga1 p Gademotta Formation (ETH72-5); Ga2 p Gademotta Formation (ETH72-6, ETH72-9); Ga3 p Gademotta Formation (ETH72-7b, ETH72-1); Ga4 p Gademotta Formation (ETH72-8B); KT p Kapedo Tuffs. Artifact abbreviations: Misc p miscel- laneous retouched piece; P core p platform core; LP core p Levallois preferential core; D core p discoidal core; HD tool p heavy-duty tool; LR core p Levallois recurrent core. The relationship between intersite distance (km) and the Dice-Sorensen coefficient calculated between all MSA assemblages (bottom). Solid line represents least squares regression. Tryon and Faith Middle Stone Age Variability S247 cores, blades, grindstones, and anvils (fig. 4, top; table 1). demism (SMCRE). The SMCRE is the familiar dry savanna Many of these artifact types are also found in the early LSA of eastern Africa characterized by habitats that range from at Enkapune ya Muto (Ambrose 1998), and the LMSA sites Acacia-Commiphora deciduous bushland/thicket to semides- in which they are found have been characterized by some as ert grassland and shrubland with distinctive arid-adapted transitional between the MSA and the LSA (e.g., Diez-Martı´n fauna such as the oryx (Oryx beisa) and Gre´vy’s zebra (Equus et al. 2009; Marks and Conard 2008). Our quantitative analysis grevyi; Grubb et al. 1999). Faunal data from eastern African supports their characterization as distinct from other “typical” MSA sites consistently show hominin occupation of grassland EMSA and LMSA assemblages. habitats broadly similar to those found in the SMCRE (Tryon If geography is a meaningful correlate of assemblage com- et al. 2010, 2012). Other sites occur in the Eastern Forest position, we expect assemblages from sites that are closer transitional zone along the Indian Ocean coast, the Lake Vic- together to be more similar to one another than they are to toria regional mosaic (LVRM), or near the ecotone between more distant sites. To quantify similarity between MSA as- the SMCRE and scattered highland Afromontane areas (fig. semblages, we calculated Dice-Sørensen coefficients for all 1). pairs of assemblages, measuring distance using latitude and We have a poor understanding of the relationship between longitude coordinates (listed in table 1). For any given pair, environmental and behavioral variability in eastern Africa be- the Dice-Sørensen coefficient is calculated as2j/(a ϩ b) , where cause of disparate spatial and temporal scales among paleoen- j is the number of artifact classes that co-occur at assemblages vironmental and archaeological data sets (Blome et al. 2012). A and B, a is the number of artifact classes at site A, and b The composition and boundaries of the biogeographic zones is the number of artifact classes at site B. As illustrated in defined here likely shifted over time, providing different en- figure 4 (bottom), there is a weak but significant inverse re- vironmental conditions for local hominin populations (see lationship between pairwise site distance and the Dice- discussion in Basell 2008), and it is these local conditions that Sørensen coefficients (r p Ϫ0.194 ,P ! .001 ), meaning that our analyses of interassemblage variability suggest are most pairs of assemblages that are nearby are more similar than important in structuring archaeological variability. Coarse- pairs of assemblages that are distant. This correlation is largely grained analyses of hominin demography suggest that com- driven by the tendency for assemblages from the same site pared with other regions of Africa, hominin populations in (distance p 0) to be very similar to each other (fig. 4, top). eastern Africa responded to environmental change by minor Removing these from the analysis results in a much weaker, shifts in settlement location (Blome et al. 2012). On a smaller although still significant, correlation (r p Ϫ0.084 ,P p .035 ). scale, Ambrose (2001) suggested that MSA hominins, partic- To the extent that the presence and absence of particular ularly in the Lake Nakuru/Naivasha basin of Kenya, may have artifact classes can be interpreted as a measure of regional tracked ecotonal boundaries as they shifted in elevation with variation, our results suggest that geography plays a role in environmental change. driving assemblage variability but that its effect is minimal. Evidence from the LVRM provides another example of the Both the correspondence analysis (fig. 4, top) and the re- possible relation between environmental and archaeological lationship between geographic distance and assemblage sim- variability in eastern Africa. Modern distributions of a variety ilarity (fig. 4, bottom) indicate that assemblages from the same of plant and animal taxa consistently indicate eastward dis- locality tend to be more similar to each other than they are persal of forest taxa into and across the LVRM from heavily to other sites. This is confirmed by a Mann-Whitney U-test forested regions in central Africa during humid (i.e., inter- on Dice-Sørensen coefficients calculated for assemblage from glacial) phases (e.g., Kingdon 1981; Rodgers, Owen, and same locality versus assemblages from different localities Homewood 1982; Wronski and Hausdorf 2008). Conversely, (z p Ϫ7.906 ,P ! .001 ). This implies that local factors are a LVRM MSA archaeological sites consistently include arid- dominant force driving interassemblage variability. These lo- adapted fauna such as oryx (O. beisa) and Gre´vy’s zebra (E. cal factors might include site function, stone raw material grevyi) that are characteristic of the SMCRE, suggesting west- quality or abundance, or more speculatively small or restricted ward dispersal during dry (i.e., glacial) conditions (Faith et networks of information exchange within which traditions of al. 2013; Tryon et al. 2012). Hominin populations may well artifact manufacture and use were shared and maintained. have followed a similar environmentally mediated pattern of range shifts. The LVRM conspicuously marks the easternmost Environmental Controls limit of Lupemban MSA sites that are most numerous in the forested regions of central Africa and appear to be associated The environment structures aspects of the material record of with forested paleoenvironments (Barham 2000; Mercader human foragers and environmental change may explain some 2002). In contrast, LVRM MSA sites that co-occur with arid- of the variability among eastern African MSA sites. Archae- adapted fauna have small points like those found to the east ological sites are unevenly distributed among four different in the Rift Valley (Tryon et al. 2012), a connection further environmental or biogeographic zones defined by the distri- demonstrated by rare obsidian artifacts at LVRM sites with butions of endemic flora and fauna (fig. 1). Most sites occur Rift Valley sources (table A1). within White’s (1983) Somali-Masai center of regional en- The extent to which modern environments provide precise S248 Current Anthropology Volume 54, Supplement 8, December 2013 analogues for Pleistocene eastern Africa is uncertain. Fossil are to those from other sites (regardless of geographic dis- fauna from MSA sites include a number of specialized grazers tance), emphasizing the high diversity among these sites and that became extinct by the Holocene, implying important the difficulty of identifying a regional signature using artifact differences in animal communities and grassland composition typology. (Faith et al. 2011, 2012; Marean 1992, 1997). Five extinct While variability poses challenges to identifying a geo- mammals are reported from MSA sites, including an aardvark, graphic signature unique to the eastern African MSA, there Orycteropus crassidens (Lehmann 2009; MacInnes 1956), and is some evidence for temporal change. The identification of four bovids characterized by extreme hypsodonty and/or body temporal patterning among some LMSA (many !75 ka) as- mass: a relative of the wildebeest, Rusingoryx atopocranion semblages on the basis of the more frequent presence of beads, (Faith et al. 2011; Pickford and Thomas 1984); the giant ochre, bipolar cores, anvils, grindstones, and blades (fig. 4, wildebeest Megalotragus sp. (Kelly 1996; Tryon et al. 2012); top) suggests important behavioral changes during the later an extinct blesbok, Damaliscus hypsodon (Faith et al. 2012; Pleistocene. All of these elements characterize sites of the Marean and Gifford-Gonzalez 1991); and the giant longhorn regional LSA, imply a prolonged shift to LSA technologies, buffalo Syncerus antiquus (Marean 1992; Tryon et al. 2012). and suggest that the 70–35 ka interval of major population Although extant species are present, extinct taxa are numer- dispersals within and out of Africa (Soares et al. 2012) was ically dominant at MSA strata from Rusinga Island and one characterized by important technological changes. Some GvJm46 at Lukenya Hill in Kenya (Marean 1992; Tryon et al. of the artifact forms underlying this technological shift have 2012). Their dominance implies that dry grassland or scrub been suggested as markers of out-of-Africa population dis- habitats were more common than the seasonally moist short persals to southern and eastern Asia (Mellars 2006b, 2006c), grasslands found today. although Neanderthal populations in Europe apparently in- As Blome et al. (2012) stress, environmental change across dependently invented similar elements during the same time eastern Africa is asynchronous, with different areas experi- interval (d’Errico and Stringer 2011). encing variable changes in moisture availability and thus hab- Several authors have noted the absence of a clear archae- itat change. The archaeological response to this variability seen ological “out-of-Africa” signature, stressing that there is little among Pleistocene faunas and eastern African MSA sites ap- recognizably (northern or eastern) “African” about the ar- pears to be small-scale movements (e.g., range expansion, chaeological record of early Homo sapiens in Asia, Europe, or topographic shifts) as an adaptation to changing environ- Australia (e.g., Shea, 2011b; Vermeersch 2001). For example, mental conditions. At present, there is no strong evidence for the oldest members of our species outside of Africa are found environmental change as a driver of behavioral innovation at Qafzeh, Israel, in association with artifacts that fit com- among eastern African MSA sites, but this may reflect a lack fortably within the Levantine Mousterian (Hovers 2009). Evi- of stratified MSA sequences associated with detailed paleoen- dence for symbolic behavior from Qafzeh, including beads vironmental data. and ochre-stained burials, predate the oldest comparable evi- dence in Africa by at least 40 kyr (Bar-Yosef Mayer, Vander- Discussion meersch, and Bar-Yosef 2009; Hovers et al. 2003; Taborin 2003; Vanhaeren et al. 2006). Similarly, although the earliest The high degree of variability characteristic of eastern African European fossils of H. sapiens have tropical body proportions MSA lithic technology limits our ability to identify an ar- associated with a recent African origin (Pearson 2000), no chaeological signal linking eastern African MSA human pop- features of the Aurignacian (or later) Upper Paleolithic in- ulations to those that migrated out of Africa. This problem dustries suggest a technological link to Africa. The early record is exacerbated by the fact that many technical elements used from Australasia similarly lacks technological affinities with to manufacture MSA/MP artifacts, such as percussion, shap- the African MSA, consisting largely of simple forms of flake ing (fac¸onnage), retouch, biface manufacture, and wood- production common to all Pleistocene archaeological sites working are also present in older, Acheulian sites. This com- (e.g., Mulvaney and Kamminga 1999). Mellars (2006b, 2006c) mon technological foundation makes likely the independent has suggested backed pieces as a candidate artifact form, but invention of particular artifact forms even among dispersed repeated reinvention (e.g., during the Upper Pleistocene with populations of large-brained hominins (Shea 2006a; see also the Howiesons Poort and the mid-Holocene with the Wilton Lycett 2007). For example, Levallois technology apparently industries in South Africa) argues against its use as a marker developed at Acheulian sites in Africa and Eurasia from mul- of population dispersal. Shea (2011a) suggests that complex tiple independent pathways (Tryon, McBrearty, and Texier projectile technology (i.e., bow and arrow) may have facili- 2005; White, Ashton, and Scott 2011). Perhaps because of tated the spread of H. sapiens out of Africa. However, in the this, the MSA record from eastern Africa consists of lithic absence of similarities in the preserved (i.e., stone) elements types and technologies that are also found at similarly aged of this technology, the hypothesis remains difficult to test and sites in other parts of Africa and Eurasia. Our comparative makes independent evolution impossible to rule out. analyses of the eastern African data suggest that assemblages Should we expect to find an “out of Africa” signal in the from a single site are more similar to one another than they archaeological record at all? The Paleolithic archaeological Tryon and Faith Middle Stone Age Variability S249 record is the outcome of behaviors mediated by particular An emphasis on social and environmental factors shifts our social and physical environments. We expect the archaeolog- expectations in searching for the origins of “modern human ical record to reflect changes in either the social or physical behavior.” If the archaeological signature of early H. sapiens landscape, and dispersals out of Africa were likely associated varies in relation to different social and physical environ- with both. Groups dispersing into the Levant and other parts ments, then we should expect temporally and spatially variable of Eurasia likely encountered territories occupied by Nean- patterning in the expression of those elements linked to be- derthal (and other) hominin populations. While the precise havioral modernity. This is consistent with the irregular tem- nature of this interaction has long been debated, novel social poral-spatial distribution of the archaeological signatures as- environments may have acted as a catalyst for behavioral sociated with modernity (d’Errico 2003; d’Errico and Stringer change, either through innovation or emulation. It may be 2011; Habgood and Franklin 2008; McBrearty and Brooks, that the reason the earliest H. sapiens in the Levant have 2000) and parallels d’Errico and Stringer’s (2011) “cultural artifact assemblages like those of the Neanderthals is that this model” of modern human behavioral origins and Conard’s represents a successful behavioral strategy for that area. Homo (2008) model for Mosaic Polycentric Modernity. By reducing sapiens was the first hominin in Australia, and that continent’s emphasis on the link between biological and behavioral mo- distinctive biota represent a dramatic change in physical en- dernity (e.g., Hovers and Belfer-Cohen 2006; Kuhn and Hov- vironment, perhaps explaining why the archaeological record ers 2006; Lieberman and Bar-Yosef 2005) and emphasizing from that area is quite different from contemporaneous sites the situational nature of the archaeological evidence (see also found in Africa (or Europe or the Levant). We do not find Henshilwood and Marean 2003), these models are consistent an “out-of-Africa” archaeological signature because the east- with several lines of evidence, including an African biological ern African record represents adaptations to that region’s origin for our species but a Eurasian origin for the Aurig- unique setting; with new social and physical environments we nacian (Mellars 2006a), and the presence at Neanderthal sites find new archaeological signatures. The outcome appears to of some behaviors classically linked with modern humans be different behaviors for different regions or environments, (d’Errico and Stringer 2011). Such a perspective has the ad- at least for hominins using MSA and MP technology, a feature vantage of shifting approaches to the eastern African MSA that may be distinct from those using LSA and UP technology record from those that scrutinize it for evidence of “modern (see Kuhn and Stiner 2001; Mercader and Brooks 2001). human behavior” or archaeological signals of dispersal to one From this perspective, evidence for hominin occupation of that emphasizes it for what it is: the behavioral traces of early the Arabian Peninsula is particularly interesting. Rose (2004) populations of H. sapiens and closely related taxa (cf. Shea and Armitage et al. (2011) used the presence of bifacial tools 2011b; comments in Henshilwood and Marean 2003). to link the Arabian and eastern African records. However, the sample size is small, and bifaces have appeared independently multiple times in different areas (e.g., Rose 2007). In what Conclusions we consider the only convincing archaeological evidence link- ing Africa and Arabia, Rose et al. (2011) demonstrated strong From an archaeological perspective, the MSA record of eastern technological similarities in the specific details of core prep- Africa is highly variable and contains no typological or tech- aration and Levallois point production of the Nubian Type 1 nological elements that are uniquely derived relative to other method. These cores are found largely at sites in the Nile regions. Some change may be the result of subtle population Valley and its drainage basin in northeastern Africa and at movements or shifts in relation to environmental change sites in Oman dated to a relatively humid interval during the rather than innovation. This is superimposed on general Last Interglacial ∼106 ka. What is most striking about this is trends of point size decrease and a record beginning with the that multiple lines of evidence demonstrate a contemporary Last Interglacial that occasionally contains backed pieces and dispersal of eastern African flora and fauna (reviewed in Rose beads and the more frequent presence of grindstones and et al. 2011). Initial populations in Arabia may simply reflect ochre, among other artifact classes. Some of the similarities an expansion of “Africa out of Africa.” African-like physical with other regions likely represent analogous behaviors (such environments in Arabia mitigated the need to adapt to novel as the origin and spread of Levallois technology), whereas environments, and with no (known) prior hominin popu- others may indeed be homologous, such as the very particular lations in the Arabian Peninsula, the social environment may behavioral “recipes” that define Nubian Type 1 technology have remained relatively stable. This hypothesis has clear par- found only in northeastern Africa and southeastern Arabia. allels with Dennell and Roebroek’s (2005) concept of “Sa- Our ability to address these issues will certainly increase as vannahstan,” with initial hominins dispersing into Asia re- more African assemblages are studied in ways comparable maining within African-like environments. Later (∼55 ka) with those from other regions and as the number of studied sites from the Arabian peninsula during arid intervals lack and well-published sites increases. The observed variability of Nubian Type 1 cores and suggest instead the development of the eastern African MSA record reduces its utility in identi- regionally distinct variants in Arabia and Africa with envi- fying any sort of archaeological marker for dispersals “out of ronmental change (Delagnes et al. 2012). Africa.” Rather, it represents the long-term outcome of a series S250 Current Anthropology Volume 54, Supplement 8, December 2013 of local adaptations made by Middle and Upper Pleistocene Africa and Excavations at Rambogo Rock Shelter, Kenya. PhD thesis, Uni- versity of Cambridge. populations that included Homo sapiens. ———. 2008. Middle Stone Age (MSA) site distributions in eastern Africa and their relationship to Quaternary environmental change, refugia and the evolution of Homo sapiens. Quaternary Science Reviews 27:2484–2498. Beyin, Ammanuel. 2011. Upper Pleistocene human dispersals out of Africa: Acknowledgments a review of the current state of the debate. International Journal of Evolu- tionary Biology 2011: article ID 615094, doi:10.4061/2011/615094. C. A. Tryon would like to thank Erella Hovers, Steve Kuhn, Beyries, Sylvie, and Hughes Plisson. 1998. Pointes ou outils triangulaires? Leslie Aiello, Laurie Obbink, and all the conference partici- donne´es fontionelles dan le Mouste´rien levantin. Pale´orient 24:5–24. Blome, Margaret Whiting, Andrew S. Cohen, Christian A. Tryon, Alison S. pants for the wonderful week in Sweden. Any synthesis such Brooks, and Joellen Russel. 2012. The environmental context for the origins as this is the outcome of years of reflection and discussion, of modern human diversity: a synthesis of regional variability in African and we would like to acknowledge the following for directly climate 150,000–30,000 years ago. Journal of Human Evolution 62:563–592. Boe¨da, Eric 1994. Le concept Levallois: variabilite´des me´thodes. Paris: Centre or indirectly contributing to the ideas presented here: Sally National de la Recherche Scientifique. McBrearty, Alison Brooks, Erella Hovers, Steve Kuhn, Susan ———. 1995. Levallois: a volumetric construction, methods, a technique. In Anto´n, Terry Harrison, Dan Peppe, Pierre-Jean Texier, John The definition and interpretation of Levallois technology. H. L. Dibble and O. Bar-Yosef, eds. Pp. 41–68. Madison, WI: Prehistory Press. Kingston, Kay Behrensmeyer, Rick Potts, Stanley Ambrose, Bordes, Franc¸ois. 1961. Typologie du pale´olithique ancien et moyen. Publications Curtis Marean, and Nick Conard. We thank Sheila Nightingale de l’Institut de pre´histoire de l’Universite´ de Bordeaux, Me´moire no. 1 for the artifact illustrations. Portions of the research presented Bordeaux: Delmas. Bordes, Jean-Guillaume, and Nicholas Teyssandier. 2012. The Upper Paleo- here were supported by grants from the National Science lithic nature of the Chaˆtelperronian in south-western France: archeo- Foundation (BCS-0118345, BCS-0841530, BCS-1013199), the stratigraphic and lithic evidence. Quaternary International 259:95–101. Wenner-Gren Foundation, the Leakey Foundation, the Na- Brandt, Steven A., Erich C. Fisher, Elisabeth A. Hildebrand, Ralf Vogelsang, Stanley H. Ambrose, Jose´phine Lesur, and Hong Wang. 2012. Early MIS 3 tional Geographic Society, New York University, and the Uni- occupation of Mochena Borago Rockshelter, southwest Ethiopian highlands: versity of Queensland. Finally, none of this would have been implications for Late Pleistocene archaeology, paleoenvironments and mod- possible without the love and support of Rhonda Kauffman, ern human dispersals. Quaternary International 274:38–54. Brandt, Steven A., and Thomas H. Gresham. 1989. L’aˆgedelapierreen Violet Tryon, and Cornel Faith. Somalie. L’Anthropologie 94:459–482. Bra¨uer, Gu¨nter, and Michael Mehlman. 1988. Hominid molars from a Middle Stone Age at the Mumba Rock Shelter, Tanzania. American Journal of Phys- References Cited ical Anthropology 75:69–76. Ambrose, Stanley H. 1986. Hunter-gatherer adaptations to non-marginal en- Brooks, Alison S., John E. Yellen, Lisa Nevell, and Gidon Hartman. 2006. vironments: an ecological and archaeological assessment of the Dorobo Projectile technologies of the African MSA: implications for modern human model. Sprache und Geschichte in Afrika 7(2):11–42. origins. In Transitions before the transition: evolution and stability in the ———. 1998. Chronology of the Later Stone Age and food production in Middle Paleolithic and Middle Stone Age. Erella Hovers and Steven L. Kuhn, East Africa. Journal of Archaeological Science 25:377–392. eds. Pp. 233–256. New York: Springer. ———. 2001. Middle and Later Stone Age settlement patterns in the Central Brown, Francis H., and Chad R. Fuller. 2008. Stratigraphy and tephra of the Rift Valley, Kenya: comparisons and contrasts. In Settlement dynamics in Kibish Formation, southwestern Ethiopia. Journal of Human Evolution 55: the Middle Paleolithic and Middle Stone Age. N. J. Conard, ed. Pp. 21–44. 366–403. Tu¨bingen: Kerns. Bruggeman, J. Henrich, Richard T. Buffler, Mireille M. M. Guillaume, Robert ———. 2010. Coevolution of composite tool technology, constructive mem- C. Walter, Rudo von Cosel, Berhane N. Ghebretensae, and Seife M. Berhe. ory, and language: implications for the evolution of modern human be- 2004. Stratigraphy, palaeoenvironments and model for the deposition of havior. Current Anthropology 51(suppl.):S135–S147. the Abdur Reef Limestone: context for an important archaeological site Ambrose, Stanley H., and Alan Deino. 2010. Technological diversity and geo- from the last interglacial on the Red Sea coast of Eritrea. Palaeogeography, chronology of Middle Stone Age industries in the Central Rift Valley, Kenya: Palaeoclimatology, Palaeoecology 203:179–206. implications for the evolution of modern human behavior. Paleoanthro- Bulbeck, David. 2007. Where river meets sea: a parsimonious model for Homo pology 2010:A1. sapiens colonization of the Indian Ocean rim and Sahul. Current Anthro- Ambrose, Stanley H., Philip Slater, Jeffrey Ferguson, Michael Glascock, and pology 48:315–321. Ian Steele. 2012. Obsidian source survey and Late Quaternary artifact sourc- Clark, J. Desmond. 1988. The Middle Stone Age of East Africa and the be- ing in Kenya: implications for early modern human behavior. Paleoan- ginnings of regional identity. Journal of World Prehistory 2:235–305. thropology 2012:A1–A2. ———. 1993. African and Asian perspectives on the origins of modern hu- Anthony, Barbara. 1972. The Stillbay question. In Congre`s panafricain de pre´- mans. In The origin of modern humans and the impact of chronometric dating. histoire, Dakar 1967: actes de 6e session. H. J. Hugot, ed. Pp. 80–82. Paris: M. J. Aitken, C. B. Stringer, and P. A. Mellars, eds. Pp. 148–178. Princeton, Les Imprimeries Re´unies de Chambe´ry. NJ: Princeton University Press. Armitage, S. J., S. A. Jasim, A. E. Marks, A. G. Parker, V. I. Usik, and H.-P. ———. 2001a. Ecological and behavioral implications of the siting of Middle Uerpmann. 2011. The southern route “out of Africa”: evidence for an early Stone Age rockshelter and cave settlements in Africa. In Settlement dynamics expansion of modern humans into Arabia. Science 331:453–456. of the Middle Paleolithic and Middle Stone Age. N. J. Conard, ed. Pp. 91– Assefa, Zelalem. 2006. Faunal remains from Porc-Epic: paleoecological and 98. Tu¨bingen: Kerns. zooarchaeological investigations from a Middle Stone Age site in south- ———. 2001b. The earlier cultures: Middle and Earlier Stone Age, vol. 3 of eastern Ethiopia. Journal of Human Evolution 51:50–75. Kalambo Falls prehistoric site. Cambridge: Cambridge University Press. Assefa, Zelalem, Y. M. Lam, and Henk K. Mienis. 2008. Symbolic use of Clark, J. Desmond, Yonas Beyene, Giday WoldeGabriel, William K. Hart, Paul terrestrial gastropod opercula during the Middle Stone Age at Porc-Epic R. Renne, Henry Gilbert, Alban Defleur, et al. 2003. Stratigraphic, chro- Cave, Ethiopia. Current Anthropology 49:746–756. nological and behavioural contexts of Pleistocene Homo sapiens from Mid- Barham, Lawrence S. 2000. The Middle Stone Age of Zambia, south-central dle Awash, Ethiopia. Nature 423:747–752. Africa. Bristol: Western Academic & Specialist Press. Clark, J. Desmond, James L. Phillips, and Preston S. Staley. 1974. Interpre- Bar-Yosef Mayer, Daniella E., Bernard Vandermeersch, and Ofer Bar-Yosef. tations of prehistoric technology from ancient Egyptian and other Sources. 2009. Shells and ochre in Middle Paleolithic Qafzeh Cave, Israel: indications 1. Ancient Egyptian bows and arrows and their relevance for African Pre- for modern behavior. Journal of Human Evolution 56:307–314. history. Pale´orient 2:323–388. Basell, Laura S. 2007. An exploration of the Middle Stone Age of Eastern Clark, J. Desmond, and G. R. Prince. 1978. Use-wear on Later Stone Age Tryon and Faith Middle Stone Age Variability S251

microliths from Laga Oda, Haraghi, Ethiopia, and possible functional in- David L. Fox. 2011. Taxonomic status and paleoecology of Rusingoryx ato- terpretations. Azania 13:101–110. pocranion (Mammalia, Artiodactyla), an extinct Pleistocene bovid from Ru- Clark, J. Desmond, Kenneth D. Williams, Joseph W. Michels, and Curtis A. singa Island, Kenya. Quaternary Research 75:697–707. Marean. 1984. A Middle Stone occupation site at Porc Epic Cave, Dire Faith, J. Tyler, Manuel Domı´nguez-Rodrigo, Adam D. Gordon. 2009. Long- Dawa. African Archaeological Review 2:37–72. distance carcass transport at Olduvai Gorge? a quantitative examination of Close, Angela E. 2002. Backed bladelets are a foreign country. In Thinking Bed I skeletal element abundances. Journal of Human Evolution 56:247– small: global perspective on microlithization. S. L. Kuhn and R. G. Elston, 256. eds. Pp. 31–44. Archeological Papers of the American Anthropological As- Faith, J. Tyler, Richard Potts, Thomas W. Plummer, Laura C. Bishop, Curtis sociation, vol. 12. Washington, DC: American Anthropological Association. A. Marean, and Christian A. Tryon. 2012. New perspectives on middle Conard, Nicholas J. 2004. An overview of the patterns of behavioural change Pleistocene change in the large mammal faunas of East Africa: Damaliscus in Africa and Eurasia during the Middle and Late Pleistocene. In From tools hypsodon sp. nov. (Mammalia, Artiodactyla) from Lainyamok, Kenya. Pa- to symbols: from early hominids to modern humans. F. d’Errico and L. Back- laeogeography, Palaeoclimatology, Palaeoecology 361/362:84–93. well, eds. Pp. 294–332. Witswatersrand, South Africa: Witswatersrand Uni- Faith, J. Tyler, Christian A. Tryon, Daniel J. Peppe, and David L. Fox. 2013. versity Press. The fossil history of Gre´vy’s zebra (Equus grevyi) in equatorial East Africa. ———. 2008. A critical view of the evidence for a southern African origin Journal of Biogeography 40:359–369. of behavioural modernity. South African Archaeological Society Goodwin Se- Fe´blot-Augustins, Jehanne. 1990. Exploitation des matie`res premie`res dans ries 10:175–179. l’Acheule´en d’Afrique: perspectives comportementales. Pale´o 2:27–42. Coulson, Sheila, Sigrid Staurset, and Nick Walker. 2011. Ritualized behavior ———. 1993. Mobility strategies in the Late Middle Paleolithic of central in the Middle Stone Age: evidence from Rhino Cave, Tsodilo Hills, Bot- Europe and western Europe: elements of stability and variability. Journal swana. Paleoanthropology 2011:18–61. of Anthropological Archaeology 12:211–265. Day, Michael H., Mary D. Leakey, and Cassian Magori. 1980. A new hominid Gabel, Creighton. 1984. Notes on the history of archaeology in sub-Saharan fossil skull (L.H. 18) from the Ngaloba Beds, Laetoli, northern Tanzania. Africa. Boston University African Studies Center Working Papers, no. 86. Nature 284:55–56. Boston: Boston University African Studies Center. Deacon, Hilary J., and Janette Deacon. 1999. Human beginnings in South Gliganic, Luke A., Zenobia Jacobs, Richard G. Roberts, Manuel Domı´nguez- Africa: uncovering the secrets of the Stone Age. Walnut Creek, CA: Altamira. Rodrigo, and Audax Z. P. Mabulla. 2011. New ages for Middle and Later Delagnes, Anne, and Lillianne Meignen. 2006. Diversity of lithic production Stone Age deposits at Mumba Rockshelter, Tanzania: optically stimulated systems during the Middle Paleolithic in France: are there any chronological luminescence dating of quartz and feldspar grains. Journal of Human Evo- trends? In Transitions before the transition: evolution and stability in the lution 62:533–547. Middle Paleolithic and Middle Stone Age. Erella Hovers and Steven L. Kuhn, Goodwin, Astley J. H., and Clarence van Riet Lowe. 1929. The Stone Age eds. Pp. 85–108. New York: Springer. cultures of South Africa. Annals of the South Africa Museum 27:1–289. Delagnes, Anne, Chantal Tribolo, Pascal Bertran, Michel Brenet, Re´my Cras- Gossa, Tegenu, Yonatan Sahle, and Agazi Negash. 2012. A reassessment of the sard, Jacques Jaubert, Lamya Khalidi, et al. 2012. Inland human settlement Middle and Later Stone Age lithic assemblages from Aladi Springs, southern in southern Arabia 55,000 years ago: new evidence from the Wadi Surdud Afar Rift, Ethiopia. Azania 47:210–222. Middle Paleolithic site complex, western Yemen. Journal of Human Evolution Gresham, Thomas H. 1984. An investigation of an Upper Pleistocene ar- 63:452–474. chaeological site in northern Somalia. MA thesis, University of Georgia, de la Torre, Ignacio. 2004. Omo revisited: evaluating the technological skills Athens. of Pliocene hominids. Current Anthropology 45:439–465. Grubb, Peter, Oliver Sandrock, Ottmar Kullmer, Thomas M. Kaiser, and Fried- Dennell, Robin W., and Wil Roebroeks. 2005. An Asian perspective on early emann Schrenk. 1999. Relationships between eastern and southern African human dispersal from Africa. Nature 438:1099–1104. mammal faunas. In African biogeography, climate change, and early hominid d’Errico, Francesco. 2003. The invisible frontier: a multiple species model for evolution. T. Bromage and F. Schrenk, eds. Pp. 253–267. Oxford: Oxford the origin of behavioral modernity. Evolutionary Anthropology 12:188–202. University Press. d’Errico, Francesco, and Chris B. Stringer. 2011. Evolution, revolution or Gunz, Phillipp, Fred L. Bookstein, Philipp Mitteroecker, Andrea Stadlmayr, saltation scenario for the emergence of modern cultures? Philosophical Horst Seidler, and Gerhard W. Weber. 2009. Early modern human diversity Transactions of the Royal Society B 366:1060–1069. suggests subdivided population structure and a complex out-of-Africa sce- Diez-Martı´n, Fernando, Manuel Domı´nguez-Rodrigo, Policarpo Sa´nchez, Au- nario. Proceedings of the National Academy of Sciences of the USA 106:6094– dax Z. P. Mabulla, Antonio Tarrin˜o, Rebeca Barba, Mary E. Prendergast, 6098. and Luis de Luque. 2009. The Middle to Later Stone Age technological Habgood, Phillip J., and Natalie R. Franklin. 2008. The revolution that didn’t transition in East Africa: new data from Mumba Rockshelter Bed V (Tan- arrive: a review of Pleistocene Sahul. Journal of Human Evolution 55:187– zania) and their implications for the origin of modern human behavior. 222. Journal of African Archaeology 7:147–173. Haile-Selassie, Yohannes, Berhane Asfaw, and Tim D. White. 2004. Hominid Domı´nguez-Rodrigo, Manuel, Fernando Dı´ez-Martin, Audax Mabulla, Luis cranial remains from Upper Pleistocene deposits at Aduma, Middle Awash, Luque, Luis Alcala´, Antonio Tarrin˜o, Jose A. Lo´pez-Sa´ez, Rebecca Barba, Ethiopia. American Journal of Physical Anthropology 123:1–10. and Pastory Bushozi. 2007. The archaeology of the Middle Pleistocene Hammer, Michael F., August E. Woerner, Fernando L. Mendez, Joseph C. deposits of Lake Eyasi, Tanzania. Journal of African Archaeology 5:47–78. Watkins, and Jeffrey D. Wall. 2011. Genetic evidence for archaic admixture Domı´nguez-Rodrigo, Manuel, Audax Mabulla, Luis Luque, Jeroen W. Thomp- in Africa. Proceedings of the National Academy of Sciences of the USA 108: son, William J. Rink, Pastory Bushozi, Fernando Dı´ez-Martin, and Luis 15123–15128. Alcala. 2008. A new archaic Homo sapiens fossil from Lake Eyasi, Tanzania. Hauck, Thomas C. 2011. The Mousterian sequence of Hummal and its ten- Journal of Human Evolution 54:899–903. tative placement in the Levantine Middle Paleolithic. In The Lower and Donahue, Randolph E., Michael L. Murphy, and Lawrency H. Robbins. 2002– Middle Paleolithic in the Middle East and neighboring regions. J.-M. Le Ten- 2004. Lithic microwear analysis of Middle Stone Age artifacts from White sorer, R. Jagher, and M. Otte, eds. Pp. 309–323. E´ tudes et recherches ar- Paintings Rock Shelter, Botswana. Journal of Field Archaeology 29:155–163. che´ologiques de l’Universite´ de Lie`ge 126. Lie`ge: Centre de recherches ar- Douze, Katja. 2008. Le Early Middle Stone Age en Ethiopie: le contexte de che´ologiques de l’Universite´deLie`ge. Gademotta: analyse technologique du site ETH-72-6. Me´moire de Master Henshilwood, Christopher S., and Curtis W. Marean. 2003. The origin of 2, Universite´ de Toulouse II, Le Mirail. modern human behavior: critique of the models and their test implications. Eren, Metin I., Fernando Diez-Martin, and Manuel Domı´nguez-Rodrigo. Current Anthropology 44:627–651. 2013. An empirical test of the relative frequency of bipolar reduction in Hovers, Erella. 2009. The lithic assemblages of Qafzeh Cave. Oxford: Oxford Beds VI, V, and III at Mumba Rockshelter, Tanzania: implications for the University Press. East African Middle to Late Stone Age transition. Journal of Archaeological Hovers, Erella, and Anna Belfer-Cohen. 2006. “Now you see it, now you Science 40:248–256. don’t”: modern human behavior in the Middle Paleolithic. In Transitions Eren, Metin I., and Stephen J. Lycett. 2012. Why Levallois? a morphometric before the transition: evolution and stability in the Middle Paleolithic and comparison of experimental “preferential” Levallois flakes versus debitage Middle Stone Age. Erella Hovers and Steven L. Kuhn, eds. Pp. 295–304. flakes. PLoS ONE 7:29723. New York: Springer. Faith, J. Tyler, Jonah N. Choiniere, Christian A. Tryon, Daniel J. Peppe, and Hovers, Erella, Shimon Ilani, Ofer Bar-Yosef, and Bernard Vandermeersch. S252 Current Anthropology Volume 54, Supplement 8, December 2013

2003. An early case of color symbolism: ochre use by early modern humans Lieberman, Daniel E., and Ofer Bar-Yosef. 2005. Apples and oranges: mor- at Qafzeh Cave. Current Anthropology 44:491–522. phological versus behavioral transitions in the Pleistocene. In Interpreting Hublin, Jean-Jacques, Anee-Marie Tillier, and Jacques Tixier. 1987. L’hume´rus the past: essays on human, primate and mammal evolution. D. E. Lieberman, d’enfant Mouste´rien (Homo 4) du Jeble Irhoud (Maroc) dans son contexte R. J. Smith, and J. Kelley, eds. Pp. 275–296. Boston: Brill Academic. arche´ologique. Bulletin et Me´moire de la Socie´te´ d’Anthropologie de Paris 4: Lycett, Stephen J. 2007. Why is there a lack of Mode 3 Levallois technologies 115–142. in East Asia? a phylogenetic test of the Movius-Schick hypothesis. Journal Inskeep, Raymond R. 1962. The age of the Kondoa rock paintings in the light of Anthropological Archaeology 26:541–575. of recent excavations at Kisese II Rock Shelter. In Actes du IVe Congress Lycett, Stephen J., and N. von Cramon-Taubadel. 2008. Acheulean variability panafricain de pre´histoire et l’e´tude du quaternaire. G. Mortelmans and J. and hominin dispersals: a model-bound approach. Journal of Archaeological Nenquin, eds. Pp. 249–256. Tervuren: Muse´e Royal de l’Afrique Centrale. Science 35:553–562. Isaac, Glynn L., Harry V. Merrick, and Charles M. Nelson. 1972. Stratigraphic Mabulla, Audax Z. P. 1990. Preliminary report on an archaeological survey and archaeological studies in the Lake Nakuru basin, Kenya. Palaeoecology of the Ndutu Beds, Olduvai Gorge, Tanzania. Nyame Akuma 33:20–25. of Africa and the Surrounding Islands 6:225–234. MacInnes, Donald G. 1956. Fossil Tubulidentata from East Africa. Fossil Mam- Jenkins, Kirsten, J. Tyler Faith, Christian A. Tryon, Daniel J. Peppe, Sheila mals of Africa 10:1–38. Nightingale, Julian A. Ogondo, Cara Roure Johnson, and Steve Driese. 2012. Marean, Curtis W. 1990. Late Quaternary paleoenvironments and faunal ex- New excavations of a Late Pleistocene bonebed and associated MSA artifacts, ploitation in East Africa. PhD dissertation, University of California. Rusinga Island, Kenya. Paleoanthropology 2012:A17. ———. 1992. Implications of late Quaternary mammalian fauna from Lu- Johnson, Cara Roure, and Sally McBrearty. 2010. 500,000 year old blades from kenya Hill (south-central Kenya) for paleoenvironmental change and faunal the Kapthurin Formation, Kenya. Journal of Human Evolution 58:193–200. extinctions. Quaternary Research 37:239–255. Keeley, Lawrence H. 1982. Hafting and retooling: effects on the archaeological ———. 1997. Hunter-gatherer foraging strategies in tropical grasslands: record. American Antiquity 47:798–809. model-building and testing in the East African Middle and Later Stone Age. Kelly, Alison J. 1996. Intraregional and interregional variability in the East Journal of Anthropological Archaeology 16:189–225. Turkana (Kenya) and Kenyan Middle Stone Age. PhD thesis, Rutgers Uni- Marean, Curtis W., and Dianne Gifford-Gonzalez. 1991. Late Quaternary versity, New Brunswick, NJ. extinct ungulates of East Africa and palaeoenvironmental implications. Na- Kelly, Robert L. 1995. The foraging spectrum. Washington, DC: Smithsonian ture 350:418–420. Institution Press. Marks, Anthony E., and Nicholas J. Conard. 2008. Technology vs. typology: Kingdon, Jonathon. 1981. Where have the colonists come from? a zoogeo- the case for and against a transition from the MSA to the LSA at Mumba graphical examination of some mammalian isolates in eastern Africa. Af- Cave, Tanzania. In Space and time: which diachronies, which synchronies, rican Journal of Ecology 19:115–124. which scales? typology vs technology. T. Aubry, F. Almeida, A. C. Arau´jo, and Klein, Richard G. 2009. The human career. 3rd edition. Chicago: University M. Tiffagom, eds. Pp. 123–131. BAR International Series 1831. Oxford: of Chicago Press. Archaeopress. Kuhn, Steven L. 2004. Upper Paleolithic raw material economies at U¨ c¸ag˘ızlı McBrearty, Sally. 1981. Songhor: a Middle Stone Age site in western Kenya. cave, Turkey. Journal of Anthropological Archaeology 23:431–448. Quaternaria 23:171–190. Kuhn, Steven L., and Erella Hovers. 2006. General introduction. In Transitions ———. 1986. The archaeology of the Muguruk Site, Western Kenya. PhD before the transition: evolution and stability in the Middle Paleolithic and dissertation, University of Illinois, Urbana-Champaign. Middle Stone Age. Erella Hovers and Steven L. Kuhn, eds. Pp. 1–12. New ———. 1988. The Sangoan-Lupemban and Middle Stone Age sequence at York: Springer. the Muguruk site, western Kenya. World Archaeology 19:379–420. Kuhn, Steven L., and Mary C. Stiner. 2001. The antiquity of hunter-gatherers. ———. 1999. Archaeology of the Kapthurin Formation. In Late Cenozoic In Hunter-gatherers: an interdisciplinary perspective. C. Panter-Brick, R. H. Layton, and P. Rowley-Conwy, eds. Pp. 99–142. Cambridge: Cambridge environments and hominid evolution: a tribute to Bill Bishop. P. Andrews and University Press. P. Banham, eds. Pp. 143–156. London: Geological Society. ———. 2007. Body ornamentation as information technology: towards and McBrearty, Sally, and Alison S. Brooks. 2000. The revolution that wasn’t: a understanding of the significance of early beads. In Rethinking the human new interpretation of the origin of modern human behavior. Journal of revolution: new behavioural and biological perspectives on the origin and dis- Human Evolution 39:453–563. persal of modern humans. Paul Mellars, Katie Boyle, Ofer Bar-Yosef, and McBrearty, Sally, and Christian A. Tryon. 2006. From Acheulian to Middle Chris Stringer, eds. Pp. 45–54. Cambridge: McDonald Institute. Stone Age in the Kapthurin Formation, Kenya. In Transitions before the Kurashina, Hiro. 1978. An examination of prehistoric lithic technology in transition: evolution and stability in the Middle Paleolithic and Middle Stone east-central Ethiopia. PhD dissertation, University of California. Age. E. Hovers and S. L. Kuhn, eds. Pp. 257–277. New York: Springer. Lachance, Joseph, Benjamin Vernot, Clara C. Elbers, Bart Ferwerda, Alain McDougall, Ian, Francis H. Brown, and John G. Fleagle. 2005. Stratigraphic Froment, Jean-Marie Bodo, Godfrey Lema, et al. 2012. Evolutionary history placement and age of modern humans from Kibish, Ethiopia. Nature 433: and adaptation from high-coverage whole-genome sequences of diverse 733–736. African hunter-gatherers. Cell 150:457–469. McPherron, Shannon P., ed. 2007. Tools versus cores: alternative approaches to Leakey, Louis S. B. 1931. The Stone Age cultures of Kenya Colony. Cambridge: stone tool analysis. Newcastle: Cambridge Scholars. Cambridge University Press. Mehlman, Michael J. 1987. Provenience, age and associations of archaic Homo ———. 1936. Stone Age Africa: an outline of prehistory in Africa. Oxford: sapiens crania from Lake Eyasi, Tanzania. Journal of Archaeological Science Oxford University Press. 14:133–162. ———. 1943. The industries of the Gorgora Rock Shelter, Lake Tana. Journal ———. 1989. Late Quaternary archaeological sequences in northern Tanzania. of the East Africa Natural History Society 17:199–203. PhD dissertation, University of Illinois, Urbana-Champaign. Leakey, Margaret, Philip V. Tobias, John E. Martyn, and Richard E. F. Leakey. ———. 1991. Context for the emergence of modern man in eastern Africa: 1969. An Acheulean industry with prepared core technique and the dis- some new Tanzanian evidence. In Cultural beginnings: approaches to un- covery of a contemporary hominid mandible at Lake Baringo, Kenya. Pro- derstanding early hominid lifeways in the African savanna. J. Desmond Clark, ceedings of the Prehistoric Society 3:48–76. ed. Pp. 177–196. Monographien (Romisch-Germanisches Zentralmuseum, Leakey, Mary D. 1983. Africa’s vanishing art: the rock paintings of Tanzania. Forschunginstitut fur Vorund Fruhgeschichte), vol. 19. Bonn: Habelt. New York: Doubleday. Mellars, Paul. 2006a. Archaeology and the dispersal of modern humans in ———. 1984. Disclosing the past: an autobiography. New York: McGraw-Hill. Europe: deconstructing the “Aurignacian.” Evolutionary Anthropology 15: Leakey, Mary D., Richard L. Hay, D. L. Thurber, R. Protsch, and R. Berger. 167–182. 1972. Stratigraphy, archaeology, and age of the Ndutu and Naisiusiu Beds, ———. 2006b. Going east: new genetic and archaeological perspectives on Olduvai Gorge, Tanzania. World Archaeology 3:328–341. the modern human colonization of Eurasia. Science 313:796–800. Leakey, Mary D., and Louis S. B. Leakey. 1950. Excavations at the Njoro River ———. 2006c. Why did modern human populations disperse from Africa Cave. Oxford: Clarendon. ca. 60,000 years ago? a new model. Proceedings of the National Academy of Lehmann, Thomas. 2009. Phylogeny and systematics of the Orycteropodidae Sciences of the USA 103:9381–9386. (Mammalia, Tubulidentata). Zoological Journal of the Linnean Society 155: Mercader, Julio. 2002. Forest people: the role of African rainforests in human 649–702. evolution and dispersal. Evolutionary Anthropology 11:117–124. Tryon and Faith Middle Stone Age Variability S253

———. 2009. Mozambican grass seed consumption during the Middle Stone Shea, John J. 2006a. The Middle Paleolithic of the Levant: recursion and Age. Science 326:1680–1683. convergence. In Transitions before the transition: evolution and stability in Mercader, Julio, and Alison S. Brooks. 2001. Across forests and savannas: Later the Middle Paleolithic and Middle Stone Age. E. Hovers and S. L. Kuhn, eds. Stone Age assemblages from Ituri and Semliki, Democratic Republic of Pp. 189–212. New York: Springer. Congo. Journal of Anthropological Research 57:197–217. ———. 2006b. The origins of lithic projectile point technology: evidence from Mercier, Norbert, He´le`ne Valladas, Laurence Froget, Jean-Louis Joron, Pierre Africa, the Levant, and Europe. Journal of Archaeological Science 33:823– M. Vermeersch, Phillip Van Peer, and Jan Moeyersons. 1999. Thermolu- 846. minescence dating of a Middle Paleolithic occupation at Sodmein Cave, ———. 2008. The Middle Stone Age archaeology of the Lower Omo Valley Red Sea mountains (Egypt). Journal of Archaeological Science 26:1339–1345. Kibish Formation: excavations, lithic assemblages, and inferred patterns of Merrick, Harry V. 1975. Change in Later Pleistocene lithic industries in eastern early Homo sapiens behavior. Journal of Human Evolution 55:448–485. Africa. PhD dissertation, University of California. ———. 2011a. The archaeology of an illusion: the Middle-Upper Paleolithic Merrick, Harry V., Francis H. Brown, and William P. Nash. 1994. Use and transition in the Levant. In The Lower and Middle Palaeolithic in the Middle movement of obsidian in the Early and Middle Stone Ages of Kenya and East and neighboring regions. J.-M. Le Tensorer, R. Jagher, and M. Otte, eds. northern Tanzania. In Society, culture, and technology in Africa,vol.11 Pp. 169–182. E´ tudes et recherches arche´ologiques de l’Universite´deLie`ge (supplement). S. T. Childs, ed. Pp. 29–44. Philadelphia: MASCA. 126. Lie`ge: Centre de recherches arche´ologiques de l’Universite´ de Lie`ge. Miller, Sheryl F. 1979. Lukenya Hill, GvJm 46, excavation report. Nyame ———. 2011b. Homo sapiens is as Homo sapiens was: behavioral variability Akuma 14:31–34. vs. “behavioral modernity” in Paleolithic archaeology. Current Anthropology Monnier, Gilliane. 2006. The Lower/Middle Paleolithic periodization in west- 52:1–35. ern Europe: an evaluation. Current Anthropology 47:709–744. Slimak, Ludovic. 2008. The Neronian and the historical structure of cultural Morgan, Leah E., and Paul R. Renne. 2008. Diachronous dawn of Africa’s shifts from Middle to Upper Paleolithic in Mediterranean France. Journal Middle Stone Age: new 40Ar/39Ar ages from the Ethiopian Rift. Geology 36: of Archaeological Science 35:2204–2214. 967–970. Slimak, Ludovic, John I. Svendsen, Jan Mangerud, Hugues Plisson, Herbjørn Mulvaney, Derek J., and Johan Kamminga. 1999. Prehistory of Australia. Wash- P. Heggen, Alexis Bruge`re, and Pavel Y. Pavlov. 2011. Late Mousterian ington, DC: Smithsonian Institution Press. persistence near the arctic circle. Science 332:841–845. Negash, Agazi, Francis H. Brown, and Barbara Nash. 2011. Varieties and ———. 2012. Response to “Comment on late Mousterian persistence near sources of artefactual obsidian in the Middle Stone Age of the Middle Awash, the arctic circle.” Science 335:167. Ethiopia. Archaeometry 53:661–673. Soares, Pedro, Farida Alshamali, Joana B. Pereira, Vero´nica Fernandes, Nuno Newcomer, Michael H., and Franc¸ois Hivernel-Guerre. 1974. Nucleus sur M. Silva, Carla Alfonso, Marta D. Costa, et al. 2012. The expansion of e´clat: technologie et utilisation par diffe´rentes cultures pre´historiques. Bul- mtDNA haplogroup L3 within and out of Africa. Molecular Biology and letin de la Societe Pre´historique Franc¸aise 71:119–128. Evolution 29:915–927. O’Brien, Terence P. 1939. The prehistory of Uganda protectorate. Cambridge: Soriano, Sylvain, Michel Rasse, Chantal Tribolo, and Eric Huysecom. 2010. Cambridge University Press. Ounjougou: a long Middle Stone Age sequence in the Dogon country Omi, Giichi, ed. 1984. Mtongwe: an interim report of the East and Northeast (Mali). In West African archaeology: new developments, new perspectives.P. African Prehistory Project 1982. Matsumoto, Japan: Shinshu University. Allsworth-Jones, ed. Pp. 1–14. BAR International Series 2164. Oxford: Ar- ———, ed. 1986. Mtongwe: an interim report of the East and Northeast African chaeopress. Prehistory Project 1984. Matsumoto, Japan: Shinshu University. Taborin, Yves. 2003. La mer et les premiers hommes modernes. In E´changes ———, ed. 1988. Mtongwe and Mgonga: an interim report of the East and et diffusion dans la pre´histoire me´diterrane´enne. B. Vandermeersch, ed. Pp. Northeast African Prehistory Project 1986. Matsumoto, Japan: Shinshu Uni- 113–122. Paris: E´ ditions du comite´des travaux historiques et scientifiques. versity. Thackeray, Anne I. 1989. Changing fashions in the Middle Stone Age: the ———, ed. 1991. An interim report of the East and Northeast African Prehistory stone artefact sequence from Klasies River main site, South Africa. African Project 1988/1989. Matsumoto, Japan: Shinshu University. Archaeological Review 7:33–57. Pearson, Osbjorn M. 2000. Postcranial remains and the origin of modern Thompson, Jessica C. 2005. The impact of post-depositional processes on humans. Evolutionary Anthropology 9:229–247. bone surface modification frequencies: a corrective strategy and its appli- ———. 2008. Statistical and biological definitions of “anatomically modern” cation to the Loiyangalani site, Serengeti Plain, Tanzania. Journal of Ta- humans: suggestions for a unified approach to modern morphology. Evo- phonomy 3:67–89. lutionary Anthropology 17:38–48. Trinkaus, Erik. 2005. Early modern humans. Annual Review of Anthropology Pickford, Martin, and Herbert Thomas. 1984. An aberrant new bovid (Mam- 34:207–230. malia) in subrecent deposits from Rusinga Island, Kenya. Proceedings of the Tryon, Christian A. 2003. The Acheulian to Middle Stone Age transition: Koninklijke Akademie des Wetenschappen B 87:441–452. tephrostratigraphic context for archaeological change in the Kapthurin For- Pleurdeau, David. 2004. Gestion des matie`res premie`res et comportements tech- mation, Kenya. PhD dissertation, University of Connecticut. niques dans le Middle Stone Age africain: les assemblages lithiques de la grotte ———. 2006. “Early” Middle Stone Age lithic technology of the Kapthurin du Porc E´pic (Dire Dawa, Ethiopie). BAR International Series 1317. Oxford: Formation (Kenya). Current Anthropology 47:367–375. Archaeopress. Tryon, Christian A., J. Tyler Faith, Daniel J. Peppe, David L. Fox, Kieran P. Prugnolle, Franck, Andrea Manica, and Franc¸ois Balloux. 2005. Geography McNulty, Kirsten Jenkins, Holly Dunsworth, and William Harcourt-Smith. predicts neutral genetic diversity of human populations. Current Biology 2010. The Pleistocene archaeology and environments of the Wasiriya Beds, 15:159–160. Rusinga Island, Kenya. Journal of Human Evolution 59:657–671. Robertshaw, Peter, ed. 1990. A history of African archaeology. London: James Tryon, Christian A., J. Tyler Faith, Daniel J. Peppe, William F. Keegan, Kristen Currey. N. Keegan, Kirsten H. Jenkins, Sheila Nightingale, et al. Forthcoming. Sites Rodgers, William A., Charles F. Owen, and Katherine M. Homewood. 1982. on the landscape: paleoenvironmental context of late Pleistocene archae- Biogeography of East African forest mammals. Journal of Biogeography 9: ological sites from the Lake Victoria Basin, equatorial East Africa. Quater- 41–54. nary International. Rose, Jeffrey. 2004. The question of Upper Pleistocene connections between Tryon, Christian A., and Sally McBrearty. 2006. Tephrostratigraphy of the East Africa and South Arabia. Current Anthropology 45:551–555. Bedded Tuff Member (Kapthurin Formation, Kenya) and the nature of ———. 2007. The Arabian Corridor Migration Model: archaeological evi- archaeological change in the later Middle Pleistocene. Quaternary Research dence for hominin dispersals into Oman during the Middle and Upper 65:492–507. Pleistocene. Proceedings of the Seminar for Arabian Studies 37:219–237. Tryon, Christian A., Sally McBrearty, and Pierre-Jean Texier. 2005. Levallois Rose, Jeffrey I., Vitaly I. Usik, Anthony E. Marks, Yamandu H. Hilbert, Chris- lithic technology from the Kapthurin Formation, Kenya: Acheulian origin topher S. Galletti, Ash Parton, Jean Marie Geiling, Victor Cˇ erny´, Mike W. and Middle Stone Age diversity. African Archaeological Review 22:199–229. Morley, and Richard G. Roberts. 2011. The Nubian Complex of Dhofar, Tryon, Christian A., Daniel J. Peppe, J. Tyler Faith, Alex Van Plantinga, Sheila Oman: an African Middle Stone Age industry in southern Arabia. PLoS Nightingale, Julian A. Ogondo, and David L. Fox. 2012. Middle Stone Age ONE 6:e28239. artifacts and associated fauna from Rusinga and Mfangano islands, Lake Sharon, Gonen, 2007. Acheulian large flake industries: technology, chronology, Victoria, Kenya. Azania 47:14–38. and significance. Oxford: Archaeopress. Tryon, Christian A., Neil T. Roach, and M. Amelia V. Logan. 2008. The Middle S254 Current Anthropology Volume 54, Supplement 8, December 2013

Stone Age of the northern Kenyan Rift: age and context of new archaeo- Wayland, E. J., and M. C. Burkitt. 1932. The Magosian culture of Uganda. logical sites from the Kapedo Tuffs. Journal of Human Evolution 55:652– Journal of the Royal Anthropological Institute of Great Britain and Ireland 664. 62:369–390. Vallois, Henri. V. 1951. La mandibule humaine fossile de la Grotte du Porc- Wendorf, Fred, and Romauld Schild. 1974. A Middle Stone Age sequence from Epic pre`s Dire´-Daoua (Abyssinie). L’Anthropologie 55:231–238. the Central Rift Valley, Ethiopia. Warsaw: Institute of the History of Material Vanhaeren, Marian, Francesco d’Errico, Chris Stringer, Sarah L. James, Jon- Culture, Polish Academy of Sciences. athan A. Todd, and Henk K. Mienis. 2006. Middle Paleolithic shell beads ———. 1993. Probable functions of Mousterian points and convergent in Israel and Algeria. Science 1785–1788. sidescrapers in the Middle Stone Age of Ethiopia. Quaternaria Nova 3:39– Van Peer, Philip. 1992. The Levallois reduction strategy. Madison, WI: Prehis- 51. tory Press. White, Frank. 1983. The vegetation of Africa, a descriptive memoir to accompany ———. 1998. The Nile Corridor and the out-of-Africa model: an examination the UNESCO/AETFAT/UNSO vegetation map of Africa. Paris: UNESCO. of the archaeological record. Current Anthropology 39(suppl.):S115–S140. Van Peer, Philip, Veerle Rots, and Pierre M. Vermeersch. 2008. A wasted effort White, Mark, and Paul Pettitt. 1995. Technology of early Palaeolithic western at the quarry: wear analysis and interpretation of an MSA lanceolate point Europe: innovation, variability, and a unified framework. Lithics 16:27–40. from Taramsa-8, Egypt. Paleoanthropology 2008:234–250. White, Mark J., Nick M. Ashton, and Beccy Scott. 2011. The emergence, Vermeersch, Pierre M. 2001. “Out of Africa” from an Egyptian point of view. diversity, and significance of the Mode 3 (prepared core) technologies. In Quaternary International 75:103–112. The ancient human occupation of Britain. N. M. Ashton, S. G. Lewis, and Vermeersch, Pierre M., Philip Van Peer, Jan Moeyersons, and Wim Van Neer. C. B. Stringer, eds. Pp. 53–66. Amsterdam: Elsevier. 1994. Sodmein Cave site, Red Sea mountains (Egypt). Sahara 6:31–40. White, Tim D. 1986. Cut marks on the Bodo cranium: a case of prehistoric Villa, Paola, Anne Delagnes, and Lyn Wadley. 2005. A Late Middle Stone Age defleshing. American Journal of Physical Anthropology 69:503–509. artifact assemblage from Sibudu (KwaZulu-Natal): comparisons with the White, Tim D., Berhane Asfaw, David DeGusta, Henry Gilbert, Gary D. Rich- European Middle Paleolithic. Journal of Archaeological Science 32:399–422. ards, Gen Suwa, and F. Clark Howell. 2003. Pleistocene Homo sapiens from Villa, Paola, and Michel Lenoir. 2006. Hunting weapons of the Middle Stone Middle Awash, Ethiopia. Nature 423:742–747. Age and Middle Paleolithic: spear points from Sibudu, Rose Cottage, and Wronski, Torsten, and Bernhard Hausdorf. 2008. Distribution patterns of land Bouheben. South African Humanities 18:89–122. snails in Ugandan rain forests support the existence of Pleistocene forest Villa, Paola, and Sylvain Soriano. 2010. Hunting weapons of Neanderthals refugia. Journal of Biogeography 35:1759–1768. and early modern humans in South Africa: similarities and differences. Wurz, Sarah. 2002. Variability in the Middle Stone Age lithic sequence, Journal of Anthropological Research 66:5–38. 115,000–60,000 years ago at Klasies River, South Africa. Journal of Archae- Villa, Paola, Sylvain Soriano, Nicholas Teyssandier, and Sarah Wurz. 2010. ological Science 29:1001–1015. The Howiesons Poort and MSA III at Klasies River main site, Cave 1A. ———. 2013. Technological trends in the Middle Stone Age of South Africa Journal of Archaeological Science 37:630–655. Walter, Robert, Richard T. Buffler, J. Henrich Bruggemann, Mireille M. M. between MIS 7 and MIS 3. Current Anthropology 54(suppl. 8):S305–S319. Guillaume, Seife M. Berhe, Berhane Negassi, Yoseph Libeskal, et al. 2000. Yellen, John E., Alison S. Brooks, David M. Helgren, Martha Tappen, Stanley Early human occupation of the Red Sea coast of Eritrea during the last H. Ambrose, Raymond Bonnefille, James K. Feathers, et al. 2005. The interglacial. Nature 405:65–69. archaeology of Aduma Middle Stone Age sites in the Awash Valley, Ethiopia. Waweru, Veronica N. 2007. Middle Stone Age technology at Cartwright’s Site, Paleoanthropology 10:25–100. Kenya. PhD dissertation, University of Connecticut. Zwyns, Nicolas, Wil Roebroeks, Shannon P. McPherron, Adam Jagich, and Wayland, E. J. 1934. Rifts, rivers, rain and early man in Uganda. Journal of Jean-Jacques Hublin. 2012. Comment on “A Late Mousterian persistence the Royal Anthropological Institute of Great Britain and Ireland 64:333–352. near the arctic circle.” Science 335:167. Current Anthropology Volume 54, Supplement 8, December 2013 S255

Roots of the Middle Paleolithic in Eurasia

by Steven L. Kuhn

In this paper I aim to define basic characteristics of the Eurasian Middle Paleolithic (MP) and trace their development. I first outline some generalized features of MP material culture and behavior. A key characteristic of the MP is diversity in lithic technology, manifest at various temporal and spatial scales. Looking more closely, this diversity is composed of repeated recurrences of a restricted range of alternatives. Many of the varieties of lithic production common in the MP first appeared during the early MP and some even earlier. Other typical behaviors include frequent use of fire, ochre, and hafting. These are first seen in Eurasia during the later MP (marine isotope stage [MIS] 6 and 7), the period when the entire suite of MP behaviors seems to coalesce. Many shared patterns of MP behavior reflect a set of adaptations shaped by Eurasian habitats and climates. Other aspects of spatial and temporal variation are more likely to reflect demographic conditions or patterns of cultural transmission among and within groups.

The Eurasian Middle Paleolithic (MP), like the hominins that time depth, first appearing during the early Middle Pleisto- created it, was a highly evolved entity, the product of hundreds cene, if not earlier. Other features, such as consistent use of of thousands of years of cultural and biological evolution. The fire and ochre, are first evident in Eurasia during the later MP was also very long lived and extraordinarily widespread. MP (marine isotope stage [MIS] 6 and 7). It is during this It is undeniably the record of a very successful set of adap- interval at the end of the MP when the entire MP behavioral tations. In the end, the MP ceded place to other ways of life “package” seems to come together (see also Richter 2011). and other ways of doing things, but this is not an especially noteworthy characteristic: to date that has been the ultimate What Are the Key Shared Features of the MP fate of all human cultural endeavors. and How Old Are They? Within the context of the conference “Alternative Pathways to Complexity: Evolutionary Trajectories in the Middle Pa- leolithic and Middle Stone Age,” the goals of this paper are In identifying the principal shared features of the MP, I focus to define key features of the MP in Eurasia and to trace their on the later, “classic” Mousterian of the late Pleistocene. This development in time and space. Below I describe some central period, from roughly 120 to 35 ka (MIS 5e through mid-MIS characters of MP material culture and behavior as evidence 3), saw the most-evolved—and by virtue of their age, the for an evolved set of adaptations and then assess their time best-preserved—examples of MP behavior. Some compari- depth. A salient feature of the MP record is technological sons with other times and places are unavoidable: derived diversity, which is manifest at a variety of scales, spatial as features of behavior only become apparent through compar- well as temporal. Even though technology varies, however, ison with earlier or later periods (i.e., the Lower and Upper MP hominins seem to have drawn on a limited and consistent Paleolithic) or with contemporaneous cultural developments array of alternatives. Much local variability appears to reflect elsewhere (the African Middle Stone Age [MSA]). I begin by tactical and strategic responses to pressures from mobility, discussing simple features, ones that may be apparent from foraging, and raw material availability. Variation within MP a single assemblage. I subsequently discuss complex features technology at a larger scale may stem from demographic con- that emerge only from larger-scale comparisons among sites ditions and their effects on cultural transmission. Other novel or data types. characteristics of the MP record include well-developed py- We have a reasonable understanding of the origins and time rotechnology, hearth-focused activities within sites, and lim- depth of some typical MP features. One can only speculate ited use of material culture (pigments) for signaling. Many about the history of other forms of behavior. Systematic con- common forms of MP lithic technology show considerable sideration of the history of Paleolithic cultural behaviors, even for a phenomenon as well studied as lithic technology, is Steven L. Kuhn is Professor at the School of Anthropology of the limited by several factors. The widespread application of the University of Arizona (Building 30, Tucson, Arizona 85721-0030, Bordes typology as the principal instrument for analyzing MP U.S.A. [[email protected]]). This paper was submitted 3 VII lithic assemblages during the mid- and late twentieth century 13, accepted 14 VIII 13, and electronically published 21 XI 13. greatly constrained perceptions of variability within the MP.

᭧ 2013 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved. 0011-3204/2013/54S8-0008$10.00. DOI: 10.1086/673529 S256 Current Anthropology Volume 54, Supplement 8, December 2013

Archaeologists’ tendency to describe what they consider the 2000; Ruebens 2006) are also fundamentally different from most salient or important characteristics of technologies and Acheulean bifacial handaxes in size, function, and methods to minimize variability (Bar-Yosef and Van Peer 2009) also of production. makes it difficult to compare the material findings. For ex- Flake-tool-dominated assemblages without handaxes date ample, we know a good deal about handaxes in the Lower back to ca. 400 ka in many parts of Eurasia (e.g., Burdukiewicz Paleolithic and methods of flake production in the MP but and Ronen 2003; Fluck 2011; Olle´ et al. 2013; Peris 2007).1 comparatively little about flake production in the Acheulean. In some early assemblages the absence of large core tools is A third issue concerns variation in the density of useful data an effect of raw materials. In areas such as central Italy and points. Because of unavoidable taphonomic and geologic fac- parts of central Europe, knappable stone most often occurs tors (Surovell and Brantingham 2007; Surovell et al. 2009), in small pebble form, making it impractical to manufacture the number of well-preserved localities inevitably declines as large handaxes and cleavers. However, absence of large core a function of age. Moreover, for ecological or geological rea- tools is not inevitably a reflection of raw material constraints. sons, entire intervals are missing or poorly represented in the In northern Europe, an area known for its rich sources of records of some regions. These gaps in knowledge, along with excellent flint, assemblages with and without handaxes were uneven application of various dating methods, make it dif- produced during roughly the same intervals of the Middle ficult to tell equally detailed stories about all features of be- Pleistocene (Ashton and McNabb 1992; White 2000). The havior and material culture. scarcity of large Lower Paleolithic handaxes east of Italy and western Germany (Santoja and Villa 2006:429) is also certainly Simple Characters not a matter of there being no suitable stone available. In Europe, intra-assemblage diversity of flake-tool-pro- Diverse Flake-Production Systems duction systems seems to be most evident in late- or post- Acheulean assemblages, where it is associated with a declining The MP is conventionally distinguished from the Lower Pa- use of general purpose core tools and greater emphasis on leolithic based on the overwhelming numerical dominance of the production of more specialized flake tools. White and small flake tools over large core tools. Over the past 25 years Ashton (2003) characterize much of earlier Acheulean flake it has become clear that MP toolmakers used an extraordi- manufacture in northern Europe as being somewhat unstan- narily diverse set of methods to produce blanks for these flake dardized, with “moving” core platforms producing amor- tools. These include multiple variants of Levallois (Boe¨da phous or polyhedral cores. Evidence for intra-assemblage var- 1994, 1995) but also a variety of non-Levallois systems, in- iation is limited, but this may be because the number of cluding discoid, prismatic blade production, and the “Quina” well-described assemblages is small. Olle´ et al. (2013) argue method (Bar-Yosef and Kuhn 1999; Boe¨da 1991; Boe¨da, that multiple knapping methods are typical of most of the Geneste, and Meignen 1990; Bourguignon 1996, 1997; Del- long Middle Pleistocene sequence at Sierra de Attapuerca. The agnes, Jaubert, and Meignen 2007; Hiscock et al. 2009; Per- site of Yarımburgaz in Turkey, dating to MIS 7, also shows a esani 2003) and a range of less widely documented techniques varied repertoire of methods for producing flakes (Kuhn, Ar- (e.g., Delagnes 1993; Faivre 2012; Geneste and Plisson 1996; sebu¨k, and Howell 1996). In the Levant, assemblages known Slimak 1999). Although a single “modal” system of produc- as Mugharan or Acheulo-Yabrudian are characterized by tion may dominate a given assemblage, more than one system Quina-like production, blade manufacture and bifacial shap- is nearly always represented. In some instances these coex- ing existing side by side in the same assemblages (Jelinek isting systems yielded blanks with very different functional 1990). Yabrudian and allied assemblages date back to at least properties (Boe¨da 1986; Kuhn, Arsebu¨k, and Howell 1996; Meignen, Delagnes, and Bourguignon 2009, table 2.1; Shim- 250–400 ka (Gopher et al. 2010; Mercier and Valladas 2003) elmitz, Barkai, and Gopher 2011), whereas in other instances and possibly earlier. Although most investigators assign them they seem to represent alternative means of obtaining similar as late Lower Paleolithic, these flake-tool-dominated assem- forms (Meignen 2007a). blages bear a strong resemblance to later Mousterian assem- Core tools such as handaxes and cleavers, while generally blages from Europe (Dibble 1991; Jelinek 1982; Le Tensorer associated with the Lower Paleolithic Acheulean, are not ab- et al. 2007). sent from the MP. However, MP handaxes often differ in Individual technological variants have quite different his- character from Acheulean artifacts with similar forms (Villa tories. Some methods of blank production, such as discoid, 2009). Sometimes (as in the Micoquian and Mousterian of extend back to the Oldowan (reviewed by Barsky 2009; see Acheulean Tradition) bifaces are treated as large flake tools: also papers in Peresani 2003). Others have more recent be- the bifacial shaping is essentially preparation of a blank for ginnings. The origins of the Levallois method deserve special additional marginal retouch and resharpening (Boe¨da, 1. Note that I am confining the discussion to late- and post-Acheulean Geneste, and Meignen 1990; Soressi 2002; Soriano 2000). The industries. The very earliest assemblages in Eurasia are all dominated by small bifaces and leaf points of central and eastern European small flake tools (e.g., Carbonell and Rodrique´z 2006), but these are Mousterian industries (Bosinski 1967; Conard and Fischer qualitatively different sorts of assemblages and technologies from the MP. Kuhn Roots of the Middle Paleolithic in Eurasia S257 treatment because the method is so strongly identified with the manufacture of large, thick flakes, often with cortical or the Mousterian (White and Ashton 2003; White, Ashton, and otherwise blunt margins opposing a sharp edge (Al Qadi 2011; Scott 2011). Very ancient beginnings are claimed for Levallois, Bourguignon 1996, 1997; Turq 2000). Shaped tools, mainly though in some cases the putative ancestral technologies are scrapers, typically show very characteristic scaled “Quina” re- very different from better known Mousterian versions. Key touch produced by soft-hammer percussion. The resulting aspects of the Levallois method, including the strategy of “pre- tools have the potential for extensive resharpening, although determining” flake forms by shaping cores extensively, can be this potential is not always realized in discarded tools. The found in Acheulean industries dating to near the Lower/Mid- Yabrudian appears first in the Levant more than 400,000 years dle Pleistocene boundary (e.g., Goren-Inbar 2011; Sharon and ago. Shortly thereafter, similar strategies of artifact production Beaumont 2006; White, Ashton, and Scott 2011). Cores made and use can be found in Anatolia (the “proto-Charentian” of on handaxes, fitting most criteria of Boe¨da’s (1994, 1995) Karain Cave; Otte et al. 1995, 1998) and south-central Europe definition of Levallois, appear to be an integral part of late- (e.g., Velika and Mala Balanica [Mihailovic and Mihailovic Acheulean artifact production systems in both the Levant 2009; D. Mihailovic, personal communication, 2012]). Even- (DeBono and Goren-Inbar 2001) and parts of northern Eu- tually the forerunners of Quina industries appear in western rope (Tuffreau 1995; White and Ashton 2003; White, Ashton, Europe by MIS 7 (Ashton and McNabb 1992; Geneste and and Scott 2011). Researchers have also commented more gen- Plisson 1996; Moncel 2008). On one hand, the complexity erally on potential ties between Levallois and handaxe man- and number of associated traits making up the Quina/Yabru- ufacture, which seem to associate in time and space (Lycett dian pattern suggest that it is unlikely to be a result of repeated 2007; Schick 1994; Villa 2009). independent invention. Dates are consistent with the notion It is very likely that the Levallois method has multiple or- that it spread from east to west, either by diffusion or pop- igins in Eurasia and Africa (Otte 1995; Sharon 2007; Villa ulation movement. At the same time it is difficult to reconcile 2001; White and Ashton 2003:605). It is present discontin- the notion of maintaining a coherent set of associated be- uously in European late-Acheulean and early MP assemblages haviors over 200,000 years and thousands of kilometers with dating as far back as MIS 11 (Santoja and Pe´rez Gonza´lez any known process of cultural transmission. 2010; Villa 2009:266; White, Ashton, and Scott 2011), al- It is worth mentioning the time depth of blade production, though its frequency varies geographically. “Proto-Levallois” given its importance to the definition of the Upper Paleolithic artifacts are reported from many Middle Pleistocene sites (e.g., (e.g., Clark’s [1969] Mode 4). As has been widely docu- Barroso Ruı´z et al. 2011; Copeland 1998; Slimak et al. 2008; mented, prismatic blade production is not exclusive to the White and Ashton 2003; White, Ashton, and Scott 2011). Upper Paleolithic but exhibits considerable time depth in the Local sequences, such as that at Orgnac 3, may show the local MP within Eurasia (Bar-Yosef and Kuhn 1999; Meignen refinement of Levallois production during the latter part of 2007a, 2007b;Re´villion and Tuffreau 1994). A few fairly recent the Middle Pleistocene (Moncel, Moigne, and Combier 2005; MP assemblages even show limited production of small, Moncel et al. 2011), but these cases are unlikely to represent bladelet-sized blanks (Faivre 2012; Slimak 1999). Systematic the sole origin of Levallois in Europe. Rather, the Levallois production of blade blanks in Eurasia can be traced back to method probably crystalized gradually because of changing the Amudian in the Levant, dated to as early as 350–400 ka functional and strategic roles of flake production in hominin (Gopher et al. 2010). However, prismatic or non-Levallois adaptations, channeled by the fracture mechanics of isotropic blade production is also a broader category than Levallois, stone (White, Ashton, and Scott 2011:57). discoid, or Quina production. Because it encompasses a wide Even though Levallois did not originate in the MP, the range of variation, many local manifestations of blade pro- technology did become more widespread, though not ubiq- duction probably represent independent developments. uitous. It may also have become more diversified, as did other methods of flake production (White, Ashton, and Scott 2011). Habitual Transport of Artifacts Many new variants on the basic theme of Levallois appeared, or at least have been described, in later Mousterian assem- MP hominins regularly carried artifacts with them as they blages. When exactly this diversification began is uncertain. moved across the landscapes on which they foraged. This is There is an impression that pre-Mousterian Levallois is com- shown most clearly by the presence of artifacts made of raw paratively homogeneous—mainly of the centripetal recurrent materials obtained from sources located at some distance from or preferential variety (White, Ashton, and Scott 2011)—but the eventual locus of discard. Artifacts moved more than 5– this could simply be a function of restricted numbers of early 20 km almost always make up a minority of MP assemblages, cases. but at least a few are normally present. Maximum distances Another less-well-studied but equally coherent technolog- of transport are normally between 20 and 50 km, but they ical variant is called here, for want of a better term, the Quina/ may exceed 100 km in unusual cases (Feblot-Augustins 1997, Yabrudian “pattern.” Here we are dealing with a linked set 2009; Gamble 1999; Geneste 1988, 1990; Roebroeks, Kolen, of strategies for artifact production, retouch, and use. Meth- and Rensink 1988; Slimak and Giraud 2007). Flake blanks ods of blank production (which may be varied) emphasize and retouched tools were most often carried long distances, S258 Current Anthropology Volume 54, Supplement 8, December 2013 but occasionally core tools or even cores were moved as well. irregularly distributed (see Boe¨da et al. 2008; Villa and Lenoir Habitual transport of artifacts demonstrates that MP homi- 2009; Villa et al. 2009), probably because it is not always nins could and regularly did anticipate needs for certain kinds preserved or recognized. More generally, while there were of artifacts in advance, and therefore (1) activities involving hafted spear points in the Mousterian, not all pointed objects production and use of tools were staged across the landscape, were tips of weapons (Plisson and Beyries 1998), and points and (2) tool use was planned or at least regular enough to were not the only things hafted (Pawlik and Thissen 2011). demand habitual accommodation. Artifact transport may The significance of composite tools has been discussed in even provide a rough index of scales of foraging ranges (Bran- a number of contexts. The manufacture of a simple spear tingham 2006). with a stone point involves several distinct chaıˆnes ope´ratoires The regular transport of tools is a very old pattern, ex- (for shaping the wooden haft, knapping the stone point, and tending at least back to the early Acheulean or the Oldowan. preparing the binder). The integration of multiple elements, However, the distances over which artifacts were transported each formed by a different process, shows a degree of staged, do appear to have increased over time through the late Lower hierarchical organization to activities. At the very least this Paleolithic and MP (Fe´blot-Augustins 1997, 2009; Gamble implies more complex cognitive processes than needed to 1999; Roebroeks, Kolen, and Rensink 1988). At some point, produce simpler, one-part tools. Some have drawn connec- hominins began to carry artifacts over distances longer than tions between the ordered assembly of composite tools and they could have covered in a single bout of foraging, showing grammatically structured speech (Ambrose 2010). The assem- that they were planning beyond immediate needs. Certainly, bly of composite tools also implies greater investment of time the displacement of artifacts more than 50 km in some MP and energy in the production of some items of technology. sites is evidence for thinking about needs for tools several The earliest purported evidence of hafting in Europe comes days in advance. This expanded scale of technological plan- from Scho¨ningen (Germany), where simple wooden handles ning could reflect changes in cognition, for example, an in- reportedly slotted to accommodate flake inserts date to ca. crease in the capacity to project future possibilities. Alter- 400 ka (Thieme and Maier 1995). Use-wear traces, resin on natively, it may be due to expansions in ranging patterns stone tools, and impact fractures on stone points are more associated with colonization of northern latitudes and gen- common in MP assemblages dating to the later Middle and erally deteriorating climate conditions throughout MIS 4. early Upper Pleistocene (Boe¨da et al. 1996, 1999; Mazza et al. 2006; Rotts 2009; Villa and Lenoir 2009; Villa et al. 2009). In the Near East the best direct evidence for hafting is bitumen Hafting and (Simple) Composite Tools traces on late Mousterian artifacts from Syria (Boe¨daetal. Hafting is one of the few real novelties in Eurasian MP tech- 1996). Fractures consistent with impact damage are reported nologies. Evidence for hafting comes from modifications to on early Mousterian pointed tools dating back to 200–250 ka shaped tools, use-wear studies, and more rarely, from visual (Shea 1988; Villa and Lenoir 2009:71), although the sizes and or chemical detection of mastics on stone artifacts. Because shapes of these artifacts suggest that they were better suited hafting-related modification is not necessarily obvious, and to use as knives than spearheads (Shea 2006). Much earlier because use-wear and residue evidence are fugitive, we do not assemblages generally contain few pointed objects or other have a very clear idea of how often things were hafted. How- likely candidates for hafting. ever, even if they are not very common, what appear to be hafting-related modifications or wear on tools have been doc- Pyrotechnology umented in many places (e.g., Rotts 2009). Natural bitumen was used as a mastic by MP hominins in the northern Levant Fire was an important part of the adaptations of MP homi- (Boe¨da et al. 1996, 2008) as well as in Europe (Caˆrciumaru nins. Where conditions of preservation are right, MP sites et al. 2011). Reports of birchbark pitch, which supposedly generally contain evidence of fire, whether as well-defined requires a more elaborate manufacture process (Gru¨nberg hearths, dispersed ash in sediments, or burned bones and 2002; Mazza et al. 2006; Pawlik and Thissen 2011), are es- artifacts (e.g., Berna and Goldberg 2007; Goldberg et al. 2012; pecially impressive. The recognition that MP tool makers Karkanas 2002; Vallverdu´et al. 2012). Moreover, traces of fire sometimes intentionally produced very small flakes (!1 cm; in sites are found throughout the Mousterian range, from Dibble and McPherron 2006; Hovers 2007) could also be northern Europe to the southern Levant. It is difficult to say indirect evidence for hafting, although it may have other ex- whether or not there is a correlation between pyrotechnology planations. and climatic conditions, but even sites in the relatively warm, Points or pointed pieces are one commonly hafted class of dry Levant contain abundant evidence for the use of fire. In artifact, but other forms also show evidence for having been early Mousterian deposits at Hayonim cave (ca. 220 ka), for attached to handles. Hafted MP points tend to be large, and example, wood ash residue makes up a large part of the sed- it is suggested that they would have been attached to large iment even though the cultural deposits are comparatively thrust or thrown spears or used as knives (Churchill 1993; low density (Schiegel et al. 1996; Stiner et al. 2005). Shea 2006). Direct evidence for use as spear tips is rather There is convincing evidence for repeated burning in a Kuhn Roots of the Middle Paleolithic in Eurasia S259 limited area at Gesher Benot Ya’acov (Israel) dating to ca. 800 widespread among MP hominins except possibly for a specific ka (e.g., Alperson-Afil 2008), but not all sites of similar age period in the Levant when early H. sapiens was present in the contain such clear traces of fire. Burning is largely absent in region (see Hovers and Belfer-Cohen 2013). Perishable ma- the earlier Middle Pleistocene deposits at Atapuerca, for ex- terials may possibly have been used for body decoration in ample (Roebroeks and Villa 2011). Fire becomes a regular some instances (Morin and Laroulandie 2012; Peresani et al part of the record starting about 350–400 ka and is virtually 2011), but evidence is currently spotty, as would be expected. ubiquitous after 200 ka in both the Levant and Europe. There MP assemblages contain few signs that bone, antler, or ivory is some evidence for an increase in the frequency of burning were regularly used as raw material for shaped tools, and at Qesem cave (Karkanas et al. 2007) and Tabun (R. Shim- grinding and polishing were seldom employed to modify these elmitz, personal communication 2012) around 300 ka. In their materials. In marked contrast to later periods, there is also synthetic analysis of the European evidence, Roebroeks and little apparent variation across space or environments in the Villa also record two stepwise increases in the frequency of level of investment in or elaboration of technology related to evidence for fire jumps, one during MIS 7 and the second food procurement (Kuhn and Stiner 2001). during MIS 5 (Roebroeks and Villa 2011:5211). However, It is important that forms of material culture such as or- Sandgathe et al. (2011) argue that full control of fire, specif- naments or bone tools occasionally do appear in MP contexts. ically the ability to kindle fires at will, may have come later, Although these are often included in the list of features a contention that will be very difficult to disprove. thought to define “modern human behavior” (e.g., McBrearty and Brooks 2000; Shea 2011), they are behavioral variables, not fixed markers of the “modern human condition.” Beads Use of Pigments and bone artifacts are not ubiquitous within the Upper Pa- The use of pigments is another aspect of material culture that leolithic or later periods: the presence or absence of all of in Eurasia first appears in the MP. It has long been known these traits are functions of a range of influences operating that mineral pigments in the form of ochre and other metallic at both individual and group levels (cf. Shea 2011). If MP oxides were present in MP sites (reviewed by d’Errico 2008; hominins occasionally made ornaments or used antler as a Soressi and d’Errico 2007). We have few direct clues as to raw material, we can assume that they were capable of rec- how these materials were used—whether they were employed ognizing and exploiting the potential of these materials. The to color bodies, clothing, and/or objects or whether they question thus becomes not one of capability but of why they played other roles as binders or preservatives. It is safe to did not regularly take advantage of that potential. assume that they were used to decorate and to otherwise change the appearance of things. Pigments do not seem to be strongly associated with burials in the MP. Complex/Emergent Characters As of this writing, the earliest well-dated use of pigments Geographic and Temporal Diversity in Europe comes from the early Mousterian site of Maastricht Belvedere, dated to 200–250 ka (Roebroeks et al. 2012). Pig- Compared with earlier periods and with some later ones, too, ments are a regular though not ubiquitous element of the MP the Mousterian as a whole shows a striking degree of variation archaeological record dating to the Upper Pleistocene in methods used for making stone artifacts. This diversity is (d’Errico 2007, 2008; Soressi and d’Errico 2007). Interestingly, detectable both within and among assemblages. A large part there is little evidence for the use of pigments in the Near of the interassemblage diversity is apparent within a geo- East except during MIS 5, when they are associated with an- graphic frame of reference, but some of it is also chronolog- atomically modern Homo sapiens fossils (Bar-Yosef Mayer, ical. It is notable that not all areas show comparable levels of Vandermeersch, and Bar-Yosef 2009; Hovers and Belfer- variation. How much, if any, technical variation is linked to Cohen 2013). environmental factors remains uncertain. From a macroscopic perspective, Eurasian MP material cul- ture, which we know mainly through lithic technology, forms Absent Characteristics distinctive regional or geographic groupings (e.g., Clark 2009; It is not especially useful to use negative evidence to define Conard and Fischer 2000; Delagnes, Jaubert, and Meignen the MP or anything else, but it is worth recalling that some 2007; Mellars 1996:334–335; Ruebens 2006). Although certain classes of technology and material culture that are common technological elements, such as Levallois or discoid technol- later on in Eurasia and that appear in some contemporaneous ogy, are widely diffused, there are regional differences in spe- assemblages from southern Africa are very rare if not com- cific aspects of production and resulting artifact forms. Geo- pletely absent from the Mousterian. Generally, there is very graphic and temporal variation in MP lithic technology do little obvious use of durable material culture as signaling me- not involve rapid appearance and turnover of novel methods. dia in MP assemblages. Possible ornaments have been re- Instead, MP toolmakers “cycled through” a number of com- ported from a few late MP sites (e.g., d’Errico 2007; Zilha˜o mon themes (Levallois, discoid, laminar production, Quina, et al. 2010), but they never seem to have become regular and etc.). Technological procedures that appear as novel additions S260 Current Anthropology Volume 54, Supplement 8, December 2013 to a local sequence were often used previously (and later) in Contemporaneous open-air kill/butchery sites contain more other places. There are also notable interregional contrasts in typical Acheulean assemblages with handaxes and limited the amount and patterning of technological variation. For flake production (Porat et al. 2002; a similar pattern of “co- example, the late Mousterian of southern France shows a existence” may obtain in western Europe; Santoja and Villa notable (perhaps extreme) level of variation in lithic tech- 2006). However, this technological diversity is truncated nology. This variability resolves into temporal trends as well around 200–250 ka with the appearance of the early Levantine as functional/strategic associations (Delagnes, Jaubert, and Mousterian. There is some intersite diversity in lithic tech- Meignen 2007; Delagnes and Meignen 2005; Meignen, Del- nology in the early Mousterian that incorporates both Le- agnes, and Bourguignon 2009). On the other side of the Pyr- vallois and non-Levallois laminar production (Meignen enees (Casanova I Martı´et al. 2009; de la Torre 2013), tech- 2007a, 2007b), but the basic blank and artifact forms are fairly nological variation is much more limited, and there are fewer similar throughout the region (Meignen 1994; Monigal 2002). obvious temporal trends. Some of the contrasts in diversity This tendency toward broad technological homogeneity ap- among regions may be a function of differing sample sizes, pears to persist throughout the Mousterian (250–50 ka), al- but the differences in evidence for change over time among though the nature of the shared technology changes over time. regions are unlikely to be due simply to the numbers of well- The greatest level of “intra-assemblage” technical diversity characterized assemblages. may be found in the “middle” Levantine Mousterian asso- Regionalization in western Eurasia becomes apparent rel- ciated with archaic Homo sapiens fossils (Hovers 2009). atively early, ca. 400 ka. Around this time major differences in lithic assemblages are apparent between northwest Europe Diversity of Artifact Life Histories (Acheulean, Clactonian), Iberia (Acheulean) and central Eu- rope (core and flake tool/small tool industries), and the Levant Paleolithic artifacts are seldom recovered in mint condition. (Acheulean, Acheulo-Yabrudian). There may be more subtle The great majority have entered the archaeological record after differences as well, such as contrasts in late-Acheulean flake surviving one or more, sometimes many, cycles of use and technology or handaxe production, but these are difficult to rejuvenation. Part of the diversity of technological procedures isolate because of the low density of well-dated and well- within and among Mousterian assemblages can be grouped reported sites/assemblages. Researchers have long recognized into a series of alternative “life-history strategies” for flake variation across Europe in the shapes of Acheulean handaxes, tools, combining principles in the production of blanks with but strong arguments have been mounted that handaxe shapes subsequent patterns of use and maintenance. Multiple strat- are more representative of raw material differences (Santoja egies may be represented in the same assemblage. At least and Villa 2006) or artifact life histories (McPherron 2006). three distinct life-history strategies have been identified. Regional diversity in technologies of lithic production and 1. A strategy based on exerting control over blank forms manufacture continues to increase over the course of the MP. (“predetermination”) with comparatively limited subsequent Impressionistically it appears that the highest level of geo- shaping (e.g., Levallois, blade production). Artifacts may be graphic differentiation in artifact production and shaping transported, resharpened, and reduced, but the main strategy comes with the late Mousterian (ca. 70–45 ka). Interestingly, for producing fresh edges is the manufacture of new blanks however, regionally distinctive projectile point types are not (Meignen, Delagnes, and Bourguignon 2009). readily detectable until the terminal Mousterian or “transi- 2. A strategy involving minimal control over blank form tional” industries (e.g., Flas 2008; Shea 2006; Slimak 1999), but much subsequent shaping though retouch (Quina, Mi- and even then such artifacts are not universal. coquian, etc.; Meignen, Delagnes, and Bourguignon 2009). Trajectories of change between 400 and 200 ka, leading to The main means for producing fresh edges is through re- the appearance of the full MP “package,” also differ regionally. sharpening, extending the useful lives of tools. Artifacts may In Europe as a whole, a certain level of technological diversity be transported, but this is not always the case. is maintained though the late Middle and early Upper Pleis- 3. A third strategy encompassing both low levels of in- tocene. In fact, total diversity appears to increase over time. vestment in predetermining blank form and limited subse- The number of variants of Levallois method may have ex- quent shaping of blanks (discoid/denticulate). Fresh edges are panded (White, Ashton, and Scott 2011), and some distinctive obtained by extending the useful lives of cores. This might modes of blank production, such as systematic manufacture be referred to as an opportunistic or expedient strategy. of small bladelets (Faivre 2012), also develop in the later Researchers have identified other, more specific organiza- Mousterian. In marked contrast, technological diversity ac- tional characteristics in some Mousterian assemblages. In tually declines over time in the Levant. Diversity is very high what are described as “ramified production systems,” a single during the second part of the Middle Pleistocene. The flake- artifact may be treated alternatively as a tool or core over the tool-dominated assemblages making up Jelinek’s (1990) Mug- course of its use-life (Bourgouignon, Faivre, and Turq 2004). haran complex (Yabrudian, Amudian, and Acheulean facies) A strategically ramified production system, exemplified by the contain a wide range of methods for flake production. These Quina Mousterian, is thought to be distinguished from simple assemblages are confined largely to cave sites and springs. lateral recycling or scavenging of previously discarded material Kuhn Roots of the Middle Paleolithic in Eurasia S261 because the initial design of the blank facilitates the diverse also affect find densities. However, the relationship still ap- functional roles that the object might fill. pears across a wide range of cases and areas. The most par- This diversity in artifact life histories testifies to expansion simonious explanation for this association involves simple in artifact functions and to the further integration of tool use strategies that mobile populations use in keeping supplied into all aspects of hominin life. Most flake tools of the MP with usable artifacts. It is clear that Mousterian hominins were not directly involved in the food quest; instead, they regularly carried artifacts with them as they moved from place were used to make artifacts out of other materials such as to place. If they remained in a particular place only for a very wood or hide (e.g., Anderson-Gerfaud 1990; Lemorini 2000; short period of time, they might rely exclusively on those Plisson and Beyries 1998; Rotts 2009). The need to cope with transported artifacts, a few of which might then be deposited a multitude of functional requirements and to maintain a in the site. However, as the length of an occupation increased, ready supply of artifacts for both predictable and unforeseen both needs and opportunities would encourage hominins to exigencies resulted in more complex cycles of artifact pro- collect local raw materials and produce new tools on the spot. duction and use than characterized previous periods. Varia- A series of short occupations would thus produce relatively tion in artifact life histories is also tied to the mobility of low-density deposits and relatively high frequencies of trans- hominin individuals and groups, at least during the later ported and modified tools. Longer occupations would gen- Mousterian (after MIS 5; Delagnes and Rendu 2011; Kuhn erate both higher densities of artifacts and more unmodified 1995). Certain artifact forms were routinely selected for trans- debris from in situ manufacture (see Surovell et al. 2008 for port based either on high edge/mass ratios or potential for a more extensive and rigorous discussion). Contra some au- resharpening (Eren and Lycett 2011). Others were produced thors (Barton et al. 2012; Riel-Salvatore and Barton 2004), most often for local (in situ) use and discard. this variation does not necessarily describe radically different The current state of knowledge is inadequate for evaluating settlement strategies. Instead, it reflects continuous variation the time depth of these more abstract strategic features of in the duration of occupations. Mousterian technological behavior. The Acheulean and earlier It is not always possible to evaluate the relationship between industries do show a certain level of redundancy or rigidity artifact density and retouch frequency across multiple layers. in artifact production and use. Flake tools tended to be ex- Nonetheless, while the relationship is common or ubiquitous pediently or opportunistically used, whereas large core tools in Mousterian sites dating back to MIS 7, it has not yet been had longer and more diversified life histories. Moreover, tra- demonstrated in earlier periods. At Tabun cave, for example, jectories of manufacture and use were frequently linked to the expected relationship obtains in the Mousterian but not particular raw materials (Goren-Inbar and Belfer-Cohen in the Acheulo-Yabrudian layers (Clark 2008; Kuhn and Clark, 1998). Acheulo-Yabrudian assemblages in the Near East do forthcoming). Bolomor Cave, a deeply stratified Middle Pleis- show more diverse trajectories of flake-tool production and tocene site in Spain (Peris, Calatayud, and Valle 1997), also use, containing, for example, extensively curated and reduced does not show the expected pattern despite excellent pres- scrapers alongside blade tools used with little or no modifi- ervation and meticulous excavation. It is possible that ta- cation (e.g., Shimelmitz, Barkai, and Gopher 2011). Data are phonomy plays a role here, that accumulated geological dis- too few to tie technological strategies to land use and foraging turbances over time disguise the patterning. However, the behavior in the earlier periods. However, what we do know crystallization of this tendency in the MP may also reflect a suggests that whereas the roots of many typical MP techno- shift in land use and in the organization of lithic raw material logical practices are quite ancient, it is the associations and economies to service mobile populations. contexts of these behaviors that varied. For example, in the late MP of southern France, the Quina Mousterian is con- Hearth-Focused Activities sistently associated with cold conditions and exploitation of migratory game—reindeer in particular (Delagnes and Rendu Where geological conditions are amenable to the preservation 2011). However, that is certainly not the case for the Yabru- of thermal features and spatial patterning in artifacts and dian of the Levant. The same is true of discoid production, other debris, Mousterian sites often show evidence for a range Levallois, or other procedures: they appear again and again of activities centered around hearths (e.g., Henry et al. 2004; but in widely varying contexts. Mellars 1996:269–314; Vallverdu´ et al. 2012). This evidence consists of more or less well-defined fireplaces surrounded by a range of debris from toolmaking and food preparation and Density/Retouch Relationship consumption. Different zones of deposition/disposal may be In many sites of the Middle and Upper Paleolithic, the fre- evident, including toss zones for the largest/most intrusive quency of retouched tools is negatively correlated with the artifacts (Speth et al. 2012). Ashes may be removed from density of lithic artifacts in sediments (pieces per cubic meter; hearths (rake out), indicating use over an extended period. Barton et al. 2012; Kuhn 2004; Riel-Salvatore and Barton Our ability to detect recurrent patterns in the use of space is 2004). The correlation is seldom very strong; many other directly limited by the number of well-preserved and well- factors, most notably varying rates of sediment accumulation, excavated sites, so the record inevitably becomes more rarefied S262 Current Anthropology Volume 54, Supplement 8, December 2013 with age. Stiner and colleagues argue that Middle Pleistocene expressed in choices among various raw materials (stone, os- inhabitants of Qesem cave, first occupied as early as 400 ka, seous materials, etc.) as well as in the design and assembly carried food to the cave and processed and consumed it of composite artifacts encompassing multiple materials. In around fires in the same spaces they made tools (Stiner, other words, while technological strategies of Upper Paleo- Barkai, and Gopher 2011; Stiner, Gopher, and Barkai 2009). lithic hominins were at least as complex as those of MP hom- At the same time, butchery patterns may indicate that meat inins, lithic technology tells a smaller part of the story. The was partitioned and shared out in a different manner than it widespread shift to prismatic blade technology with the Upper was among Mousterian or Upper Paleolithic groups. Paleolithic, which appears to mark a reduction in technolog- ical diversity, may be in part a response to a need for regu- Discussion: When and Why Did the MP larization of components of composite artifacts (Bar-Yosef Come Together? and Kuhn 1999). There also is emerging evidence for strategic flexibility in The common characterization of the MP as a period of tech- land use during the MP, though it is more difficult to tease nological stasis and monotony is misleading. In fact, there is out. Associations between lithic technology and choices of a striking level of diversity, especially in methods for flake migratory or territorial game strongly suggest varying mo- production, within and among assemblages as well as within bility regimes organized around the distribution of important and across regions. On the other hand, while there is certainly food resources (e.g., Delagnes and Rendu 2011). But variation evidence for refinement and diversification in manufacture in mobility is as much about the assembly and disassembly technology, few truly new ways of doing things appeared dur- of groups of people as it is about mapping onto food. Some ing the period between 400 and 40 ka. The novel technological researchers (Chazan 2009; Foley and Gamble 2009; Rolland developments of the MP are hafting and use of mastics to 2004) believe that domestic spaces and true “campsites” first produce composite tools, expansion of pyrotechnology, and become recognizable in Eurasia during the later Middle Pleis- the use of pigments. Evidence of these phenomena is some- tocene (but see Villa 2009 for a different view). Evidence cited what discontinuous because of both preservation and typical includes increased use of caves and rockshelters, more fre- analytical approaches. quent presence of burning and combustion features in shelter In addition to diverse methods of flake production, MP sites, regular transport of hunted meat to shelters for pro- technologies embody a varied range of artifact life histories, cessing and consumption, and evidence that other activities from long and complicated to short and simple. MP hominins (e.g., toolmaking and use) were conducted in the same spaces. used a variety of strategies to maintain a steady supply of However, these various features almost certainly coalesced technological aids not just for foraging but for working other gradually, and early MP “campsites” may not be identical to materials into tools and clothing. In some cases, artifact life more recent and more familiar ones. The recurrent density/ histories have shown to be associated with specific patterns retouch relationship described for many MP and Upper Pa- of foraging or mobility. Recurrent correlations between re- leolithic sites may be another symptom of this development. touch frequencies and artifact densities in deeply stratified The contrast between low- and high-density assemblages that sequences and also indicate widely shared strategies for keep- first becomes apparent sometime after 250 ka shows variable ing supplied with artifacts and raw materials, strategies that and complex use of landscape with changing mixes of long- continued into the Upper Paleolithic. Along with the presence and short-term occupations involving different subsets of so- of multiple reduction sequences and life histories within as- cial groups conducting different activities. semblages, these facts testify to a high level of technological Diversity expressed over larger spatial and temporal scales flexibility at both short (behavioral) and long (evolutionary) requires a different sort of explanation. The geographic variety timescales. This flexibility in the manufacture of artifacts and in European MP technologies is undoubtedly tied closely to management of raw materials is in fact a defining feature of variation in topography and climatic conditions across the the MP. Although hard data are difficult to come by, it is European landmass. By the end of MIS 5, MP hominin pop- likely that the MP shows greater procedural and tactical di- ulations were distributed from north-central Europe to the versity in lithic technology than both earlier and later cultural southern Levant and from the Atlantic to the Altai Mountains. periods. The radically different habitats that hominins encountered in At a local scale, the variety in MP lithic technologies and different places as well as the severe fluctuations in climate raw material economies attests to a significant degree of stra- (summarized in Andel and Davies 2003; Rohling et al. 2013) tegic complexity. MP hominins were heavily dependent on would inevitably have led to divergent adaptations among stone artifacts as well as products made using stone tools. local populations. A part of the spatial variability in methods They were prepared to invest more time and energy than their of flake production also probably reflects neutral variation. forebears in preparing multipart composite tools. What may Small populations dispersed across Europe and western Asia, distinguish the MP from the Upper Paleolithic is the fact that isolated from one another by distance and environmental these strategies are expressed entirely in lithic technology. In barriers, would have provided an ideal setting for drift-like the Upper Paleolithic, more tactical and strategic variation is processes and random loss and retention of certain behaviors. Kuhn Roots of the Middle Paleolithic in Eurasia S263

The periodic contraction of local populations into refugia resembles cultural developments in the Eurasian Upper Pa- during harsh climatic intervals and their subsequent reex- leolithic. Moreover, is fits expectations for cumulative cultural pansion during times when conditions were more favorable evolution or “cultural ratcheting” (Tennie, Call, and Toma- (a source/sink model; Dennel 2009) could help maintain or sello 2009; Tomasello 1999). However, the apparent absence even amplify regional variation. Contrasts among regions in of such patterns within the MP does not necessarily indicate patterns of chronological change and diversity might reflect that MP hominins were somehow unable to sustain cultural variation in population sizes and connectivity as well as eco- development over long periods. We can question the evidence logical factors (reviewed in Kuhn 2012). for cumulative cultural evolution in the Upper Paleolithic and The suite of traits identified here as typical of the MP began MSA. The Stillbay and Howieson’s Poort, like the Aurignacian to appear in the second half of the Middle Pleistocene but and Gravettian, appear more like equivalent (though differ- continued to develop and coalesce throughout the Upper ent) cultural developments than examples of stepwise increase Pleistocene (see also d’Errico 2007 for a discussion of other in the complexity or variety of human achievement. What is elements of behavior). Many methods of flake production really remarkable about these particular developments is the typical of the Mousterian first appear around 400–500 ka if geographic scale over which specific cultural variants are not long before that. Likewise, regular use of fire and perhaps found. As such, they may speak more to qualities of con- something recognizable as “domestic spaces” may appear be- nectivity among subpopulations and the potential for culture tween 300 and 400 ka. Pigments and hafting only become traits to spread than to “cultural ratcheting.” evident after 250 ka. There may be an expansion and diver- The appearance of pigments in the MP marks a new role sification of flake production around this same period or for material culture, that of signaling. The use of ochre by slightly later. It should not be surprising that the gradual MP hominins certainly shows evidence for body decoration coalescence of the characters that define the MP in western as a component in social interaction. We might also expect Eurasia resembles what has been documented for the African some of the variation across space in MP artifact forms to MSA (d’Errico 2007; McBrearty and Brooks 2000). Both pat- be evidence for conscious identity formation, what is some- terns are best understood as the gradual evolution of a broad times termed “emblemic style” (see Hegmon 1992; Wiessner suite of cultural adaptations in response to shifting regional 1983). As discussed above, Mousterian assemblages do show environmental and demographic conditions. As we might also considerable variation in modes of production of stone tools. expect, the specific adaptive patterns and sets of behaviors However, lithic technology is an inherently stationary practice differ across the two regions. Nonetheless, the trajectories of that requires close proximity to observe. For this reason, var- development are similar in structure. Resemblances extend to iation in the fine details of technological procedures is likely the discontinuous histories of many elements. In both the to represent simply repetition of different learned patterns MP and MSA, specific forms of behavior—whether methods rather than an active signaling of identity (Tostevin 2007). for working stone or approaches to ornamenting the body— On the other hand, there is a widespread impression that the appear, disappear, and sometimes reappear over time (Hovers MP record contains little expressive “stylistic” variation, par- and Belfer-Cohen 2006). ticularly in artifacts such as points. This impression rests Specific behaviors and archaeological evidence aside, one largely on the observation that in later periods, spear, dart, obvious difference between the MSA record of southern Africa and arrow points tend to show stronger geographic and tem- and the MP record of Eurasia concerns the spatial scale at poral patterning than other stone-tool forms, which is further which directional change can be detected. It currently appears presumed to represent intentional assertion of group identity that a single sequence of industrial succession and techno- style. To a certain extent, appreciation of fine-scale stylistic logical change characterizes South Africa, Namibia, and variation in artifact forms may have been inhibited by the neighboring countries between MIS 5 and 3 (Jacobs and Rob- decades-long hegemony of the Bordes typological system and erts 2008; Jacobs et al. 2008; Wurz 2013). This contrasts later by the recognition that a good deal of variation in tool sharply with Europe, which shows a diversified patchwork of forms reflects use-lives rather than design (e.g., Dibble 1987, independent regional sequences. In one area the Mousterian 1995; McPherron 2006). Even so, regional differences in point may exhibit a definite directional trend, while in a neighboring forms or in the shapes of other retouched tools, for that area there may be a very different trend, or none at all. The matter, are not as easily identified as they are, for example, Levant shows a general pattern similar to southern Africa, in the sub-Saharan African MSA (McBrearty and Brooks with a single, fairly consistent set of cultural developments 2000). The bifacial points found in some central, southern, between 225 and 40 ka (Hovers and Belfer-Cohen 2013). and eastern European assemblages are a possible exception, However, the Levant actually covers a much smaller area than but even these are too widespread to nominate as a regional southern Africa, similar in size to one of the subregions within or local style. In combination with the absence of consistent Europe. systems of body ornamentation in durable materials, this ob- A number of researches have observed that the progression servation leads to the conclusion that intergroup signaling of of late MSA assemblages in southern Africa—in particular identity was not an important function of MP material culture the sequence of MSA 2, Stillbay, and Howieson’s Poort— in general (Kuhn and Stiner 2007). S264 Current Anthropology Volume 54, Supplement 8, December 2013

In the end, two factors than can best account for many of climatic constraints in contrast with the African MSA, for the emergent features of the MP are population size and example (Powell, Shennan, and Thomas 2009)? If so, why did population structure. There is an emerging view that MP Mousterian hominins seldom transcend these limitations, by hominins existed at very low population densities compared diversifying the subsistence base, for example? Were cognitive with later Paleolithic or recent hunter-gatherers and that their factors responsible for this apparent lack of flexibility (e.g., populations were highly fragmented. In addition to genetic Wynn and Coolidge 2004), and if so, what features of cog- evidence for small effective population sizes (Bocquet-Appel nition are implicated and how can this be reconciled with the and Degioanni 2013; Lalueza-Fox 2013), there are ecological observed technological variability and ability to colonize di- reasons to expect thinly distributed populations. Although verse habitats? An alternative view is that because of their they consumed a wide variety of foods (e.g., Henry, Brooks, particular behavioral and biological evolutionary histories, and Piperno 2010), the isotopic and faunal evidence suggest MP populations were “locked into” a particular set of re- that MP hominins tended to feed at a high trophic level, sponses such that immediate changes in behavior would have focusing their efforts on large-game animals (Bocherens and had the immediate consequence of lower fitness (Andersson, Drucker 2003; Gaudzinski-Windheuser and Niven 2009; To¨rnberg, and To¨rnberg 2014). In other words, the particular Richards and Trinkaus 2009; Stiner 2013; Stiner and Kuhn fitness landscapes on which MP populations operated may 2009; Stiner, Munro, and Surovell 2000; Stiner et al. 1999; simply have made certain evolutionary changes more difficult Villa and Lenoir 2009). Because the collective biomass of large than others (Brantingham, Kuhn, and Kerry 2004) indepen- herbivores is lower than small game or plants in terrestrial dent of the essential characteristics of the hominins them- environments, the combination of large-game-focused for- selves. Channeled into certain trajectories by its evolutionary aging and the comparatively high individual energy require- history, the particular experiment that was the MP played out ments of Neanderthal anatomy (Froelhe and Churchill 2009; in a different way than contemporary experiments in other Snodgrass and Leonard 2009; Sorenson and Leonard 2001) parts of the world. leads to the inference that populations were sparse and co- resident groups very small, even in comparison with recent foragers. In aggregate, these diverse lines of evidence point to References Cited partitioning of MP populations into many small, demograph- Alperson-Afil, Nira. 2008. Continual fire-making by hominins at Gesher Benot ically fragile local demes. Because it would be difficult for Ya’aqov, Israel. Quaternary Science Reviews 27:1733–1739. such groups to maintain contact over large areas, social net- Al Qadi, Amjad. 2011. Le Yabroudien en Syrie: e´tat de la question et enjeux works would have been limited in scale and highly com- de la recherche´. In The Lower and Middle Palaeolithic in the Middle East and neighbouring regions. Marcel Otte, Reto Jagher, and Jean-Marie Le Ten- partmentalized (Gamble 1999). sorer, eds. Pp. 77–84. E´ tudes et Recherches Arche´ologiques de l’Universite´ Small absolute population sizes along with frequent local de Lie`ge, no. 126. Lie`ge: E´ tudes et recherches arche´ologiques de l’Universite´ extinction of local demes could account for a range of typical de Lie`ge. Ambrose, Stanley. 2010. Coevolution of composite-tool technology, construc- features of the Eurasian MP. As has been widely discussed, tive memory, and language: implications for the evolution of modern hu- small populations both limit the rates of invention and in- man behavior. Current Anthropology 51(suppl.):S135–S147. crease the likelihood that beneficial but rare novelties will be Andel, Thies van, and William Davies, eds. 2003. Neanderthals and modern humans in the European landscape during the last glaciation. Cambridge: lost (Bentley, Hahn, and Shennan 2004; Henrich 2004; Hovers McDonald Institute for Archaeological Research. and Belfer-Cohen 2006; Powell, Shennan, and Thomas 2009). Anderson-Gerfaud, Patricia. 1990. Aspects of behaviour in the Middle Pa- Frequent extinction of local demes could also constrain rates laeolithic: functional analysis of stone tools from southwest France. In The emergence of modern humans. Paul Mellars, ed. Pp. 389–418. Edinburgh: of drift and lead to recycling of old technological themes Edinbugh University Press. (Premo and Kuhn 2010). The limited use of technology for Andersson, Claes, Anton To¨rnberg, and Petter To¨rnberg. 2014. An evolu- signaling in the MP is also likely a consequence of small tionary developmental approach to cultural evolution. Current Anthropol- ogy, forthcoming. groups that seldom came into contact with and hence seldom Ashton, Nick, and John McNabb. 1992. The interpretation and context of the needed to identify themselves to strangers (Kuhn and Stiner High Lodge industries. In High Lodge: excavations by G. de G. Sieveking 2007). Likewise, the existence of many independent regional 1962–68 and J. Cook 1988. Nick Ashton, Jill Cook, S. G. Lewis, and J. Rose, eds. Pp. 164–168. London: British Museum Press. trajectories in technological change is most parsimoniously Barroso Ruı´z, Cecelio, Daniel Botella Ortega, Miguel Caparro´s, Anne Marie understood as a consequence of poorly interconnected social Moigne, Vincenzo Celiberti, Agne`s Testu, Deborah Barsky, et al. 2011. The networks that tended to keep cultural information local rather Cueva del Angel (Lucena, Spain): an Acheulean hunters habitat in the south of the Iberian Peninsula. Quaternary International 243:105–126. than potentiating its rapid diffusion (Kuhn 2012). Barsky, Deborah. 2009. An overview of some African and Eurasian Oldowan This proposition that MP populations were thinly distrib- sites: evaluation of hominin cognition levels, technological advancement uted and partitioned into small, disconnected local demes and adaptive skills. In Interdisciplinary approaches to the Oldowan. Erella Hovers and David Braun, eds. Pp. 39–47. Vertebrate Paleobiology and might account for many features of the MP record, but this Paleoanthropology Series. New York: Springer. hypothesis in turn begs important questions. The most fun- Barton, C. Michael, Julien Riel-Salvatore, John M. Anderies, and Gabriel damental is why hominins might have maintained such high Popescu. 2011. Modeling human ecodynamics and biocultural interactions in the Late Pleistocene of western Eurasia. Human Ecology 39(6):705–725. trophic levels and low demographic potential for so long. Can Bar-Yosef, Ofer, and Steven Kuhn. 1999. The big deal about blades: laminar the demographic patterns be attributed to environmental or technologies and human evolution. American Anthropologist 101:322–338. Kuhn Roots of the Middle Paleolithic in Eurasia S265

Bar-Yosef, Ofer, and Philip Van Peer. 2009. The chaıˆne ope´ratoire approach in Chazan, Michael. 2009. Assessing the Lower to Middle Paleolithic transition. Middle Paleolithic archaeology. Current Anthropology 50(1):103–131. In Sourcebook of Paleolithic transitions. Marta Camps and Parth Chuahan, Bar-Yosef Mayer, Daniella, Bernard Vandermeersch, and Ofer Bar-Yosef. 2009. eds. Pp. 237–243. New York: Springer. Shells and ochre in Middle Paleolithic Qafzeh Cave, Israel: indications for Churchill, Steven, E. 1993. Weapon technology, prey size selection, and hunt- modern behavior. Journal of Human Evolution 56(3):307–314. ing methods in modern hunter-gatherers: implications for hunting in the Bentley, R. Alexander, Matthew W. Hahn, and Stephen Shennan. 2004. Ran- Palaeolithic and Mesolithic. In Hunting and animal exploitation in the later dom drift and culture change. Proceedings of the Royal Society of London B Paleolithic and Mesolithic of Eurasia. Gail Peterkin, Harvey Bricker, and Paul 271:1443–1450. Mellars, eds. Pp. 11–24. Archaeological Papers of the American Anthro- Berna, Francesco, and Paul Goldberg. 2007. Assessing Palaeolithic fire tech- pological Association no. 4. Washington, DC: American Anthropological nology and associated hominin behavior in Israel. Israel Journal of Earth Association. Sciences 56:107–121. Clark, Amy E. 2008. Changes in occupation intensity during the Lower and Bocherens, Herve´, and D. E. Drucker. 2003. Reconstructing Neandertal diet Middle Paleolithic at Tabun Cave, Israel. MA thesis, University of Arizona, from 120,000 to 30,000 BP using carbon and nitrogen stable isotopic abun- Tucson. dances. In Le roˆle de l’environnement dans le comportements des chasseurs- Clark, Geoffery. 2009. Accidents of history: conceptual frameworks in pa- cueilleurs pre´historiques. Marilyn Patou-Mathis and Herve´Bocherens, eds. leoarchaeology. In Sourcebook of Paleolithic transitions. Marta Camps and Pp. 1–8. BAR International Series 1105. Oxford: British Archaeological Parth Chuahan, eds. Pp. 19–43. New York: Springer. Reports. Clark, Grahame. 1969. World prehistory: a new outline. 2nd edition. Cambridge: Bocquet-Appel, Jean-Pierre, and Anna Degioanni. 2013. Neanderthal demo- Cambridge University Press. graphic estimates. Current Anthropology 54(suppl. 8):S202–S213. Conard, Nicholas J., and B. Fischer. 2000. Are there recognizable cultural Boe¨da, E. 1986. La debitage Levallois de Biache St-Vaast (Pas-de-Calais): pre- entities in the German Middle Palaeolithic? In Toward modern humans: miere e´tude technologique. In Chronostratigraphie et facie´s culturels du pa- Yabrudian and Micoquian, 400–50k years ago. Avraham Ronen and Wein- le´olithique inferieur et moyen dans l’europe du nord-ouest. Alain Tuffreau and stein-Evron, eds. Pp. 7–24. BAR International Series 850. Oxford: British A. Somme´, eds. Pp. 209–218. Bulletin de l’Association Franc¸aise pour Archaeological Reports. l’E´ tude du Quaternaire, no. 26, supple´ment. Paris: L’Association Franc¸aise Copeland, Lorraine. 1998. The Lower Paleolithic of Jordan. In The Prehistoric pour l’E´ tude du Quaternaire. archeology of Jordan. Donald Henry, ed. Pp. 5–22. BAR International Series ———. 1991. Approche de la variabilite´des syste`mes de production lithique 705. Oxford: British Archaeological Reports. des industries du pale´olithique infe´rieur et moyen: chronique d’une var- DeBono, Hila, and Na’ama Goren-Inbar. 2001. Note on a link between Acheu- iabilite´ attendee. Techniques et Culture 17/18:37–79. lian handaxes and the Levallois method. Journal of the Israel Prehistoric ———. 1994. Le concept Levallois: variabilite´ des me´thodes. Paris: CNRS. Society 31:9–23. ———. 1995. Levallois: a volumetric construction, methods, a technique. In Delagnes, Anne. 1993. Un mode de production ine´dit au pale´olithique moyen The definition and interpretation of Levallois technology. Harold Dibble and dans l’industrie du niveau 6e du Pucheuil (Seine-Maritime). Pale´o 5:111– Ofer Bar-Yosef, eds. Pp. 41–69. Madison, WI: Prehistory Press. 120. Boe¨da, Eric, S. Bonilauri, J. Connan, D. Jarvie, Norbert Mercier, M. Tobey, Delagnes, Anne, Jacques Jaubert, and Liliane Meignen. 2007. Les technocom- He´le`ne Valladas, H. al Sakhel, and Sultan Muhesen. 2008. Middle Palaeo- plexes du pale´olithique moyen en Europe occidental dans leur cadre dia- lithic bitumen use at Umm el Tlel around 70,000 BP. Antiquity 82:853– chronique et ge´ographique. In Ne´andertaliens: biologie et cultures. Bernard 886. Vandermeersch, Bruno Maureille, and Yves Coppens, eds. Pp. 213–229. Boe¨da, Eric, Jacques Connan, Daniel Dessort, Sultan Muhesen, Norbert Mer- Documents Pre´historiques 23. Paris: CTHS. cier, He´le`ne Valladas, and Nadine Tisne´rat. 1996. Bitumen as a hafting Delagnes, Anne, and Liliane Meignen. 2005. Diversity of lithic production material on Middle Palaeolithic artefacts. Nature 380(6572):336–338. systems in the Middle Paleolithic in France: are there any chronological Boe¨da, Eric, Jean-Michel Geneste, C. Griggo, Norbert Mercier, Sultan Mu- trends? In Transitions before the transition: evolution and stability in the hesen, J.-L. Reyss, A. Taha, and He´le`ne Valladas. 1999. A Levallois point Middle Paleolithic and Middle Stone Age. Erella Hovers and Steven Kuhn, embedded in the vertebra of a wild ass (Equus africanus): hafting, projectiles eds. Pp. 85–107. New York: Springer. and Mousterian hunting weapons. Antiquity 73(280):394–402. Delagnes, Anne, and William Rendu. 2011. Shifts in Neandertal mobility, Boe¨da, Eric, Jean-Michel Geneste, and Liliane Meignen. 1990. Identification technology and subsistence strategies in western France. Journal of Archae- de chaıˆnes ope´ratoires lithiques du pale´olithique ancient et moye´n. Pale´o 2: ological Science 38(8):1771–1783. 43–80. de la Torre, Ignacio, Jorge Martı´nez-Moreno, and Rafael Mora. 2013. Change Bosinski, Gerhard. 1967. Die mittelpala¨olithischen Funde im westlichen Mit- and stasis in the Iberian Middle Paleolithic: considerations on the signifi- teleuropa. Cologne: Bo¨hlau. cance of Mousterian technological variability. Current Anthropology Bourguignon, Laurence. 1996. La conception de de´bitage Quina. Quaternaria 54(suppl. 8):S320–S336. Nova 6:149–166. Dennell, Robin. 2009. The Paleolithic settlement of Asia. Cambridge: Cambridge ———. 1997. Le Mouste´rien de type Quina: nouvelle de´finition d’une entite´ University Press. technique. Doctoral thesis, Universite´ Paris X Nanterre. d’Errico, Francesco. 2007. The origin of humanity and modern cultures: ar- Bourguignon, Laurence, Jean-Philippe Faivre, and Alain Turq. 2004. Rami- chaeology’s view. Diogenes 54(2):122–133. fication des chaıˆnes ope´ratoires: une spe´cificite´du Mouste´rien? Pale´o 16:37– ———. 2008. The rouge et le noir: implications of early pigment use in 48. Africa, the Near East and Europe for the origin of cultural modernity. In Brantingham, P. Jeffery. 2006. Measuring forager mobility. Current Anthro- Current themes in Middle Stone Age research. Marlize Lombard, Christine pology 47(3):435–459. Sievers, and V. Ward, eds. Pp. 168–174. South African Archaeological Society Brantingham, P. Jeffery, Steven Kuhn, and Kristopher W. Kerry. 2004. On the Goodwin Series, vol. 10. Vlaeberg: South African Archaeological Society. difficulty of the Middle-Upper Paleolithic transition. In The Early Upper Dibble, Harold. 1987. The interpretation of Middle Paleolithic scraper mor- Paleolithic beyond western Europe. P. Jeffery Brantingham, Steven Kuhn, and phology. American Antiquity 52:109–117. Kristopher W. Kerry, eds. Pp. 1–13. Berkeley: University of California Press. ———. 1991. Mousterian assemblage variability on an interregional scale. Burdukiewicz, Jan Michal, and Avraham Ronen, eds. 2003. Lower Palaeolithic Journal of Anthropological Research 47(2):239–257. small tools in Europe and the Levant. BAR International Series 1115. Oxford: ———. 1995. Middle Paleolithic scraper reduction: background, clarification, British Archaeological Reports. and review of the evidence to date. Journal of Archaeological Method and Carbonell, Eudald, and Xose´Pedro Rodrı´quez. 2006. The first human settle- Theory 2:299–368. ment of Mediterranean Europe. Comptes Rendus Palevol 5(1/2):291–298. Dibble, Harold, and Shannon McPherron. 2006. The missing Mousterian. Caˆrciumaru, Marin, Radica-Mariana Ion, Elena Cristina Nit¸u, and Radu S¸te- Current Anthropology 47(5):777–803. fa˘nescu. 2012. New evidence of adhesive as hafting material on Middle and Eren, Metin I., and Stephen J. Lycett. 2011. Why Levallois? a morphometric Upper Palaeolithic artefacts from Gura Cheii-Raˆs¸nov Cave (Romania). Jour- comparison of experimental “preferential” Levallois flakes versus debitage nal of Archaeological Science 39(7):1942–1950. flakes. PLoS ONE 7(1):e29273, doi:10.1371/journal.pone.002927. Casanova I Martı´, Joel, Jorge Martı´nez Moreno, Rafael Mora Turcal, and Faivre, Jean-Philippe. 2012. A material anecdote but technical reality: bladelet Ignacio de la Torre. 2009. Strate´gies techniques dans le pale´olithique moyen and small blade production during the recent Middle Paleolithic at the du sud-est des Pyre´ne´es. L’Anthropologie 113:313–340. Combe-Grenal rockshelter. Lithic Technology 37:5–24. S266 Current Anthropology Volume 54, Supplement 8, December 2013

Fe´blot-Augustins, Jehanne. 1997. La circulation des matie`res premie`res au pa- Paleolithic societies. Brian Adams and Brooke Blades, eds. Pp. 232–246. New le´olithique.E´ tudes et Recherches Arche´ologiques de l’Universite´ de Lie`ge, York: Wiley-Blackwell. no. 75. Lie`ge: Service de pre´histoire, Universite´deLie`ge. Hovers, Erella. 2007. The many faces of cores-on-flakes: a perspective from ———. 2009. Revisiting European Upper Palaeolithic raw material transfers: the Levantine Mousterian. In Cores or tools? alternative approaches to stone the demise of the cultural ecological paradigm? In Lithic materials and tool analysis. Shannon McPherron, ed. Pp. 42–74. Cambridge: Cambridge Paleolithic societies. Brian Adams and Brooke S. Blades, eds. Pp. 25–46. New Scholars Press. York: Wiley-Blackwell. ———. 2009. The lithic assemblages of Qafzeh cave. Oxford: Oxford University Flas, Damien. 2008. La transition du pale´olithique moyen au supe´rieur dans la Press. plaine septentrionale de l’Europe. Anthropologica et Praehistorica 119. Brus- Hovers, Erella, and Anna Belfer-Cohen. 2006. “Now you see it, now you sels: Royal Belgian Society of Anthropology and Prehistory. don’t”: modern human behavior in the Middle Paleolithic. In Transitions Fluck, H. 2011. Culture in the Lower Palaeolithic: technological variability in before the transition: evolution and stability in the Middle Paleolithic and Middle Pleistocene Europe. In Investigating archaeological cultures: material Middle Stone Age. Erella Hovers and Steven Kuhn, eds. Pp. 295–304. New culture, variability and transmission. Benjamin W. Roberts and Marc Vander York: Springer. Lindern, eds. Pp. 77–96. New York: Springer. ———. 2013. On variability and complexity: lessons from the Levantine Foley, Robert, and Clive Gamble. 2009. The ecology of social transitions in Middle Paleolithic record. Current Anthropology 54(suppl. 8):S337–S357. human evolution. Philosophical Transactions of the Royal Society B 364:3267– Jacobs, Zenobia, and Richard G. Roberts. 2008. Testing times: old and new 3279. chronologies for the Howiesons Poort and Still Bay Industries in environ- Froelhe, Andrew, and Steven Churchill. 2009. Energetic competition between mental context. South African Archaeological Society Goodwin Series 10:9– Neandertals and anatomically modern humans. Paleoanthropology 2009:96– 34. 116. Jacobs, Zenobia, Richard G. Roberts, Rex F. Galbraith, Hillary Deacon, Rainer Gamble, Clive. 1999. Palaeolithic societies of Europe. Cambridge: Cambridge Gru¨n, Alex Mackay, Peter Mitchell, Ralf Vogelsang, and Lynn Wadley. 2008. University Press. Ages for the Middle Stone Age of Southern Africa: implications for human Gaudzinski-Windheuser, Sabine, and Laura Niven. 2009. Hominid subsistence behavior and dispersal. Science 322:733–735. patterns during the Middle and Late Paleolithic in northwestern Europe. Jelinek, Arthur J. 1982. The Middle Paleolithic in the Southern Levant, with In The evolution of hominin diets: integrating approaches to the study of comments on the appearance of modern Homo sapiens.InThe transition Paleolithic subsistence. Jean-Jacques Hublin and Michael Richards, eds. Pp. from Lower to Middle Paleolithic and the origin of modern man. Avraham 99–111. Dordrecht: Springer. Ronen, ed. Pp. 57–101. BAR International Series 151. Oxford: British Ar- Geneste, Jean-Michel. 1988. Syste`mes d’approvisionnement en matie`res pre- chaeological Reports. mie`res au pale´olithique moyen et au pale´olithique supe´rieur en Aquitaine. ———. 1990. The Amudian in the context of the Mugharan tradition at the In La mutation, vol. 8 of L’homme de Ne´andertal. Marcel Otte and Janusz Tabun Cave (Mount Carmel), Israel. In The emergence of modern humans. K. Kozlowski, eds. Pp. 61–70. E´ tudes et Recherches Arche´ologiques de Paul Mellars, ed. Pp. 81–90. Ithaca, NY: Cornell University Press. l’Universite´ de Lie`ge, no. 35. Lie`ge: Universite´deLie`ge. Karkanas, Panagiotis. 2002. Ash, bones and guano: a study of the minerals ———. 1990. De´velopment des syste`mes de production lithique au cours du and phytoliths in the sediments of Grotte XVI, Dordogne, France. Journal pale´olithique moyen en Aquitaine septentrionale. In Pale´olithique moyen of Archaeological Science 29(7):721–732. recent et pale´olithique supe´rieur ancien en Europe: ruptures et transitions. Karkanas, Panagiotis, Ruthi Shahack-Gross, Avner Ayalon, Miryam Bar-Mat- Catherine Farizy, ed. Pp. 203–123. Me´moires du Muse´e de Pre´histoire d’Ile- thews, Ran Barkai, Aamos Frumkin, Avi Gopher, and Mary C. Stiner. 2007. de-France, no. 3. Nemours: Association pour la promotion de la recherche Evidence for habitual use of fire at the end of the Lower Paleolithic: site- arche´ologique en I˚le-de-France. formation processes at Qesem Cave, Israel. Journal of Human Evolution Geneste, Jean-Michel, and Hugues Plisson. 1996. Production et utilisation de 53(2):197–212. l’outillage lithique dans le Mouste´rien du sud-ouest de la France: les Tares Kuhn, Steven. 1995. Mousterian lithic technology: an ecological approach. a` Sourzac, Valle´e de l’Isle, Dordogne. Quaternaria Nova 6:343–368. Princeton, NJ: Princeton University Press. Goldberg, Paul, Harold Dibble, Francisco Berna, Dennis Sandgathe, Shannon ———. 2004. Middle Paleolithic assemblage formation at Riparo Mochi. In McPherron, and Alain Turq. 2012. New evidence on Neandertal use of fire: Processual archaeology. A. Johnson, ed. Pp. 31–60. London: Praeger. examples from Roc de Marsal and Pech de l’Aze´IV.Quaternary International ———. 2012. Emergent patterns of creativity and innovation in early tech- 247:325–340. nologies. In Creativity, innovation and human evolution. S. Elias, ed. Pp. Gopher, Avi, Avner Ayalon, Miryam Bar-Matthews, Ran Barkai, Amos Frum- 69–87. Amsterdam: Elsevier. kin, Panagiotis Karkanas, and Ruthi Shahack-Gross. 2010. The chronology Kuhn, Steven, Gu¨ven Arsebu¨k, and F. Clark Howell. 1996. The Middle Pleis- of the Late Lower Paleolithic in the Levant based on U-Th ages of speleo- tocene lithic assemblage from Yarimburgaz Cave, Turkey. Pale´orient 22(1): thems from Qesem cave, Israel. Quaternary Geochronology 5(6):644–656. 31–49. Goren-Inbar, Na’ama, and Anna Belfer-Cohen. 1998. The technological abil- Kuhn, Steven, and Amy E. Clark. Forthcoming. Artifact densities and assem- ities of the Levantine Mousterian: cultural and mental capacities. In Nean- blage formation in the Paleolithic: evidence from Tabun Cave. Journal of dertals and modern humans in western Asia. Takeru Akazawa, Kenichi Akoi, Anthropological Archaeology. and Ofer Bar-Yosef, eds. Pp. 205–221. New York: Plenum. Kuhn, Steven, and Mary C. Stiner. 2001. The antiquity of hunter-gatherers. ———. 2011. Culture and cognition in the Acheulian industry: a case study In Hunter-gatherers, an interdisciplinary perspective. Catherine Panter-Brick, from Gesher Benot Ya’aqov. Philosophical Transactions of the Royal Society Robert H. Layton, and Peter Rowley-Conwy, eds. Pp 99–142. Cambridge: B 366:1038–1049. Cambridge University Press. Gru¨nberg, Judith. 2002. Middle Palaeolithic birch-bark pitch. Antiquity 76: ———. 2007. Body ornamentation as information technology: towards an 15–16. understanding of the significance of early beads. In Rethinking the human Hegmon, Michelle. 1992. Archaeological research on style. Annual Review of revolution: new behavioural and biological perspectives on the origins and Anthropology 21:517–536. dispersal of modern humans. Paul Mellars, Katie Boyle, Ofer Bar-Yosef, and Henrich, Joe. 2004. Demography and cultural evolution: how adaptive cultural C. Stringer, eds. Pp. 45–54. Cambridge: McDonald Institute of Archaeo- processes can produce maladaptive losses: the Tasmanian case. American logical Research. Antiquity 69:197–214. Lalueza-Fox, Carles. 2013. Agreements and misunderstandings among three Henry, Amanda, Alison Brooks, and Delores Piperno. 2010. Microfossils in scientific fields: paleogenomics, archaeology, and human paleontology. Cur- calculus demonstrate consumption of plants and cooked foods in Nean- rent Anthropology 54(suppl. 8):S214–S220. derthal diets (Shanidar III, Iraq; Spy I and II, Belgium). Proceedings of the Lemorini, Cristina. 2000. Reconnaitre des tactiques d’exploitation du milieu au National Academy of Sciences of the USA 108(2):486–491. pale´olithique moyen: e´tude fonctionnelle des industries lithiques de Grotta Henry, Donald, Harold J. Hietala, Arlene Rosen, Yuri Demidenko, Vitali Usik, Breuil (Latium) et de La Combette (Bonnieux, Vaucluse, France). BAR In- and T. L. Armagan. 2004. Human behavioral organization in the Middle ternational Series 858. Oxford: British Archaeological Reports. Paleolithic: were Neanderthals different? American Anthropologist 106:17– Le Tensorer, Jean-Marie, Reto Jagher, Phillipe Rentzel, Thomas Hauck, Kristin 31. Ismail-Meyer, Christine Pu¨mpin, and Dorota Wojtczak. 2007. Long-term Hiscock, Peter, Alain Turq, Jean-Philippe Faivre, and Laurence Bourguignon. formation processes at the natural springs Nadaouiyeh and Hummal in the 2009. Quina procurement and tool production. In Lithic materials and El-Kowm oasis, central Syria. Geoarchaeology 22(6):621–639. Kuhn Roots of the Middle Paleolithic in Eurasia S267

Lycett, Steven. 2007. Why is there a lack of Mode 3 Levallois technologies in kiran, and Piere Noiret. 1995. E´ volution technique au pale´olithique ancien East Asia? a phylogenetic test of the Movius-Schick hypothesis. Journal of de Karain. L’Anthropologie 90:529–561. Anthropological Archaeology 26(4):541–575. Pawlik, Alfred, and Ju¨rgen Thissen. 2011. Hafted armatures and multi-com- Mazza, Paul Peter A., Fabio Martini, Benedetto Sala, Maurizio Magi, Maria ponent tool design at the Micoquian site of Inden-Altdorf, Germany. Journal Perla Colombini, Gianna Giachi, Francesco Landucci, Cristina Lemorini, of Archaeological Science 38:1699–1708. Francesca Modugno, and Erica Ribechini. 2006. A new Palaeolithic dis- Peresani, Marco, ed. 2003. Discoid lithic technology: advances and implications. covery: tar-hafted stone tools in a European Mid-Pleistocene bone-bearing BAR International Series 1120. Oxford: Archaeopress. bed. Journal of Archaeological Science 33:1310–1318. Peresani, Marco, Ivana Fiore, Monica Gala, Matteo Romandini, and Antonio McBrearty, Sally, and Alison Brooks. 2000. The revolution that wasn’t: a new Tagliacozzo. 2011. Late Neandertals and the intentional removal of feathers interpretation of the origin of modern humans. Journal of Human Evolution as evidenced from bird bone taphonomy at Fumane Cave 44 ky B.P., Italy. 39:453–463. Proceedings of the National Academy of Sciences of the USA, doi:10.1073 McPherron, Shannon. 2006. What typology can tell us about Acheulian hand- /pnas.1016212108. axe production. In Axe age: Acheulian tool-making from quarry to discard. Peris, Josep Ferna´ndez. 2007. La Cova del Bolomor: las industrias lı´ticas del Na’ama Goren-Inbar and Gonen Sharon, eds. Pp. 267–285. London: Equi- Pleistoceno Medio en el a´mbito del Mediterra´neo peninsular. Valencia: J. Agui- nox. lar. Meignen, Liliane. 1994. Pale´olithique moyen au proche-orient: le phe´nome´ne Peris, Josep Ferna´ndez, P. M. Calatayud, and R. M. Valle. 1997. Cova del laminaire. In Les industries laminaires au pale´olithique moyen.Ste´phane Bolomor: los primeros habitantes de las tierras valencianas. Valencia: Dipu- Re´villion and Alain Tuffreau, eds. Pp. 125–161. Paris: CNRS. tacio´n de Valencia. ———. 2007a. Middle Paleolithic blady assemblages in the Near East: a Plisson, Hugues, and Sylvie Beyries. 1998. Pointes ou outils triangulaires? reassessment. In Caucasus and the initial dispersals in the Old World.Pp. donne´es fonctionnelles dans le Mouste´rien Levantin. Pale´orient 24(1):5–16. 133–148. Russian Academy of Sciences, Institute of the History of Material Porat, Naomi, Michael Chazan, Henry P. Schwarcz, and Liora Kolska Horwitz. Culture, vol. 21. St. Petersburg: PV. 2002. Timing the Lower to Middle Paleolithic boundary: new dates from ———. 2007b. Le phe´nome`ne laminaire au proche-orient, du pale´olithique the Levant. Journal of Human Evolution 43:107–122. infe´rieur aux de´buts du pale´olithique supe´rieur. In Un sie`cle de construction Powell, Adam, Stephen Shennan, and Mark Thomas. 2009. Late Pleistocene du discours scientifique en pre´histoire: congre`s du centenaire de la Socie´te´ demography and the appearance of modern human behavior. Science 324: pre´historique franc¸aise. J. Evin, ed. Pp. 79–94. Paris: Socie´te´ pre´historique 1298–1301. franc¸aise. Premo, Luke, and Steven Kuhn. 2010. Modeling effects of local extinctions Meignen, Liliane, Anne Delagnes, and Laurence Bourguignon. 2009. Patterns on culture change and diversity in the Paleolithic. PLoS ONE 5(12):e15582. of lithic raw material procurement and transformation during the Middle Re´villion, Ste´phane, and Alain Tuffreau, eds. 1994. Les industries laminaires Paleolithic in western Europe. In Lithic materials and Paleolithic societies. au pale´olithique moyen. Paris: CNRS. Brian Adams and Brooke Blades, eds. Pp. 15–24. New York: Wiley-Blackwell. Richards, Michael, and Erik Trinkaus. 2009. Isotopic evidence for the diets of European Neanderthals and early modern humans. Proceedings of the Na- Mellars, Paul. 1996. The Neanderthal legacy. Princeton, NJ: Princeton Uni- tional Academy of Sciences of the USA 106(38):16034–16039. versity Press. Richter, Ju¨rgen. 2011. When did the Middle Paleolithic begin? In Neanderthal Mercier, Norbert, and He´le`ne Valladas. 2003. Reassessment of TL age estimates lifeways, subsistence and technology: 150 years of Neanderthal studies. Nicholas of burnt flints from the Paleolithic site of Tabun Cave, Israel. Journal of Conard and Ju¨rgen Richter, eds. Pp. 99–110. New York, Springer. Human Evolution 45:401–409. Riel-Salvatore, Julien, and C. Michael Barton. 2004. Late Pleistocene tech- Mihailovic, Dusˇan, and Bojiana Mihailovic. 2009. Cultural regionalization in nology, economic behavior, and land-use dynamics in southern Italy. Amer- the Palaeolithic of the Middle Danube basin and western Balkans. In Le ican Antiquity 69(2):257–274. concept de territoires dans le pale´olithique supe´rieur europe´en. Franc¸ois Djind- Roebroeks, Wil, Jan Kolen, and Eelco Rensink. 1988. Planning depth, antic- jian, Jansz Koizlowski, and Nuno Bicho, eds. Pp. 93–101. BAR International ipation and the organization of Middle Palaeolithic technology: the archaic Series 1938. Oxford: Archaeopress. natives meet Eve’s descendents. Helenium 28:17–34. Moncel, Marie-He´le`ne, ed. 2008. Le site de Payre: occupations humaines dans Roebroeks, Wil, Mark J. Siera, Trine Kellberg Nielsen, Dimitri De Loecker, la valle´e du Rhoˆne a`lafinduple´istoce`ne moyen et au de´but du ple´istoce`ne Josep Maria Pare´s, Charles E. S. Arps, and Herman J. Mu¨chere. 2012. Use supe´rieur.Me´moire de la Societe´ Pre´historique Franc¸ais 46. Paris: Societe´ of red ochre by early Neandertals. Proceedings of the National Academy of pre´historique franc¸ais. Sciences of the USA 109:1889–1894. Moncel, Marie-He´le`ne, Anne-Marie Moigne, and Jean Combier. 2005. Pre- Roebroeks, Wil, and Paola Villa. 2011. On the earliest evidence for habitual Neandertal behaviour during isotopic stage 9 and the beginning of stage use of fire in Europe. Proceedings of the National Academy of Sciences of the 8: new data concerning fauna and lithics in the different occupation levels USA 108(13):5209–5214. of Orgnac 3 (Arde´che, southeast France): occupation types. Journal of Ar- Rohling, Eelco J., Katherine M. Grant, Andrew P. Roberts, and Juan-Cruz chaeological Science 32:1283–1301. Larrasoan˜a. 2013. Paleoclimate variability in the Mediterranean and Red Moncel, Marie-He´le`ne, Anne-Marie Moigne, Youssef Sam, and Jean Combier. Sea regions during the last 500,000 years: implications for hominin migra- 2011. The emergence of Neanderthal technical behavior: new evidence from tions. Current Anthropology 54(suppl. 8):S183–S201. Orgnac 3 (Level 1, MIS 8), southeastern France. Current Anthropology 52(1): Rolland, Nicholas. 2004. Was the emergence of homebases and domestic fire 37–75. a punctuated event? a review of the Middle Pleistocene record in Eurasia. Monigal, Katheryn. 2002. The Levantine leptolithic: blade production from Asian Perspectives 43(2):248–280. the Lower Paleolithic to the dawn of the Upper Paleolithic. PhD dissertation, Rotts, Veerle. 2009. The functional analysis of the Mousterian and Micoquian Southern Methodist University, Dallas, TX. assemblages of Sesselfelsgrotte, Germany: aspects of tool use and hafting Morin, Euge`ne, and Ve´ronique Laroulandie. 2012. Presumed symbolic use of in the European Late Middle Palaeolithic. Quarta¨r 56:37–66. diurnal raptors by Neanderthals. PLoS ONE 7(3):e32856, doi:10.1371 Ruebens, Karen. 2006. A typological dilemma: Micoquian elements in con- /journal.pone.0032856. tinental northwestern Europe during the Last Glacial Cycle (MIS 5d–3). Olle´, Andreu, Marina Mosquera, Xose´Pedro Rodrı´guez, Arturo de Lombera- Lithics 27:58–73. Hermida, Marı´a Dolores Garcı´a-Anto´n, Paula Garcı´a-Medrano, Luna Pen˜a, Sandgathe, Dennis, Harold Dibble, Paul Goldberg, Shannon McPherron, Alain et al. 2013. The Early and Middle Pleistocene technological record from Turq, Laura Niven, and Jamie Hodgkins. 2011. On the role of fire in Nean- Sierra de Atapuerca (Burgos, Spain). Quaternary International 295:138–167. dertal adaptations in western Europe: evidence from Pech de l’Aze´IV and Otte, Marcel. 1995. The nature of Levallois. In The definition and interpretation Roc de Marsal. Paleoanthropology 2011:216–242. of Levallois technology. Harold Dibble and Ofer Bar-Yosef, eds. Pp. 117– Santoja, Manuel, and Arturo Pe´rez Gonza´le. 2010. Mid-Pleistocene Acheulean 243. Madison, WI: Prehistory Press. industrial complex in the Iberian Peninsula. Quaternary International 223/ Otte, Marcel, Is¸ın Yalc¸inkaya, Janusz Kozlowski, Ofer Bar-Yosef, Ignacio Lopez 224:154–161. Bayon, and Harun Tas¸kıran. 1998. Long-term technical evolution and hu- Santoja, Manuel, and Paola Villa. 2006. The Acheulean of western Europe. man remains in the Anatolian Palaeolithic. Journal of Human Evolution 34: In Axe age: Acheulian tool-making from quarry to discard. N. Goren-Inbar 413–431. and G. Sharon, eds. Pp. 429–478. London: Equinox. Otte, Marcel, Is¸ın Yalc¸inkaya, Janusz Kozlowski, Ofer Bar-Yosef, Harun Tas¸- Schick, Kathy. 1994. The Movius line reconsidered. In Integrative paths to the S268 Current Anthropology Volume 54, Supplement 8, December 2013

past. R. Corruccini and R. Chiochon, eds. Pp. 569–596. Engelwood Cliffs, and the hare: small game use, the broad spectrum revolution, and Paleolithic NJ: Prentice Hall. demography. Current Anthropology 41(1):39–73. Schiegel, Solveg, Paul Goldberg, Ofer Bar-Yosef, and Stephen Weiner. 1996. Stiner, Mary C., Natalie D. Munro, Todd A. Surovell, Eitan Tchernov, and Ash deposits in Hayonim and Kebara Caves, Israel: macroscopic, micro- Ofer Bar-Yosef. 1999. Paleolithic population growth pulses evidenced by scopic and mineralogical observations, and their archaeological implication. small animal exploitation. Science 283:190–194. Journal of Archaeological Science 23:763–781. Surovell, Todd A. 2008. Toward a behavioral ecology of lithic technology: cases Sharon, Gonen. 2007. Acheulian large flake industries: technology, chronology from Paleoindian archaeology. Tucson: University of Arizona Press and significance. BAR International Series 1701. Oxford: Archaeopress. Surovell, Todd A., and P. Jeffery Brantingham. 2007. A note on the use of Sharon, Gonen, and Peter Beaumont. 2006. Victoria West: a highly standard- temporal frequency distributions in studies of prehistoric demography. Jour- ized prepared core technology. In Axe age: Acheulian tool-making from nal of Archaeological Science 34:1868–1877. quarry to discard. N. Goren-Inbar and G. Sharon, eds. Pp. 181–199. London: Surovell, Todd A., J. Finely, G. M. Smith, P. Jeffery Brantingham, and Robert Equinox. L. Kelly. 2009. Correcting temporal frequency distributions for taphonomic Shea, John J. 1988. Spear points from the Middle Paleolithic of the Levant. bias. Journal of Archaeological Science 36:1715–1724. Journal of Field Archaeology 15:441–456. Tennie, Claudio, Josep Call, and Michael Tomasello. 2009. Ratcheting up the ———. 2006. The origins of lithic projectile point technology: evidence from ratchet: on the evolution of cumulative culture. Philosophical Transitions of Africa, the Levant and Europe. Journal of Archaeological Science 33:823–846. the Royal Society B 364(1528):2405–2415. ———. 2011. Homo sapiens is as Homo sapiens was. Current Anthropology Thieme, Hartmut, and Reinhard Maier, eds. 1995. Archa¨ologische ausgrabungen 52(1):1–35. im braunkohlentagebau Scho¨ningen. Hannover: Landkreis Helmstedt. Shimelmitz, Ron, Ran Barkai, and Avi Gopher. 2011. Systematic blade pro- Tomasello, Michael. 1999. The cultural origins of human cognition. Cambridge, duction at late Lower Paleolithic (400–200 ka) Qesem Cave, Israel. Journal MA: Harvard University Press. of Human Evolution 61(4):458–479. Tostevin, Gilbert. 2007. Social intimacy, artefact visibility, and acculturation Slimak, Ludovic. 1999. Mise en e´vidence d’une composante laminaire et la- models of Neanderthal-modern human interaction. In Rethinking the hu- mellaire dans un complex Mouste´rien du sud de la France. Pale´o 11:89– man revolution: new behavioural and biological perspectives on the origins and 110. dispersal of modern humans. Paul Mellars, Katie Boyle, Ofer Bar-Yosef, and Slimak, Ludovic, and Y. Giraud. 2007. Circulations sur plusieurs centaines de Chris Stringer, eds. Pp. 341–357. Cambridge: MacDonald Institute for Ar- kilome`tres durant le pale´olithique moyen: contribution a` la connaissance chaeological Research. des socie´te´s Ne´andertaliennes. Comptes Rendus Palevol 6:359–368. Tuffreau, Alain. 1995. The variability of Levallois technology in northern Slimak, Ludovic, Steven Kuhn, He´le`ne Roche, Damase Mouralis, Hilke Bui- France and neighboring areas. In The definition and interpretation of Levallois tenhuis, Nur Balkan-Atlı, Didier Binder, Catherine Kuzucuog˘lu, and Herve technology. Harold Dibble and Ofer Bar-Yosef, eds. Pp. 413–427. Madison, Guillou. 2008. Kaletepe Deresi 3 (Turkey): archaeological evidence for early WI: Prehistory Press. human settlement in Central Anatolia. Journal of Human Evolution 54(1): Turq, Alain. 2000. Le pale´olithique infe´rieur et moyen entre les Valee´s de la 99–111. Dordogne et do Lot. Pale´o (suppl. 2). Les Eyzies: SAMRA. Snodgrass, J. Josh, and William Leonard. 2009. Neandertal energetics revisited: Vallverdu´, Josep, Susana Alonso, Ame`lia Bargallo´, Rau´l Bartrolı´, Gerard Cam- insights into population dynamics and life history evolution. Paleoanthro- peny, A´ ngel Carrancho, Isabel Expo´sito, et al. 2012. Combustion structures pology 209:220–237, doi:10.4207/PA.2009.ART31. of archaeological level O and Mousterian activity areas with use of fire at Sorenson, Mark V., and William R. Leonard. 2001. Neandertal energetics and the Abric Romanı´ rockshelter (NE Iberian Peninsula). Quaternary Inter- foraging efficiency. Journal of Human Evolution 40:483–495. national 247:313–324. Soressi, Marie. 2002. Le Mouste´rien de tradition Acheule´enne du sud-ouest Villa, Paola. 2001. Early Italy and the colonization of western Europe. Qua- de la France. PhD dissertation, University of Bordeaux 1. ternary International 75:113–130. Soressi, Marie, and Francesco D’Errico. 2007. Pigments, gravures, parures: les ———. 2009. The Lower to Middle Paleolithic transition. In Sourcebook of comportements symboliques controverse´s des Ne´andertaliens. In Ne´ander- Paleolithic transitions. Marta Camps and Parth Chuahan, eds. Pp. 265–270. taliens: biologie et cultures. Bernard Vandermeersch, Bruno Maureille, and New York: Springer. Yves Coppens, eds. Pp. 297–309. Documents Pre´historiques 23. Paris, Villa, Paola, Paolo Boscato, Filomena Ranaldo, and Annamaria Ronchitelli. CTHS. 2009. Stone tools for the hunt: points with impact scars from a Middle Soriano, Sylvain. 2000. Outillage bifacial et outillage sur e´clat au pale´olithique Paleolithic site in southern Italy. Journal of Archaeological Science 36(3):850– ancien et moyen: coexistence et interaction. PhD dissertation, Universite´ 859. Paris X Nanterre. Villa, Paola, and Michel Lenoir. 2009. Hunting and hunting weapons of the Speth, John D., Liliane Meignen, Ofer Bar-Yosef, and Paul Goldberg. 2012. Lower and Middle Paleolithic of Europe. In The evolution of hominin diets: Spatial organization of Middle Paleolithic occupation X in Kebara Cave integrating approaches to the study of Paleolithic subsistence. Jean-Jacques (Israel): concentrations of animal bones. Quaternary International 247:85– Hublin and Michael Richards, eds. Pp. 59–87. Dordrecht: Springer. 102. White, Mark. 2000. The Clactonian question: on the interpretation of core- Stiner, Mary C. 2013. An unshakable Middle Paleolithic? trends versus con- and-flake assemblages in the British Lower Paleolithic. Journal of World servatism in the predatory niche and their social ramifications. Current Prehistory 14(1):1–62. Anthropology 54(suppl. 8):S288–S304. White, Mark, and Nicholas Ashton. 2003. Lower Paleolithic core technology Stiner, Mary C., Ran Barkai, and Avi Gopher. 2011. Hearth-side socioeco- and the origins of the Levallois method in northwestern Europe. Current nomics, hunting and paleoecology during the late Lower Paleolithic at Anthropology 44:598–608. Qesem Cave, Israel. Journal of Human Evolution 60(2):213–233. White, Mark, Nick Ashton, and Beccy Scott. 2011. The emergence, diversity Stiner, Mary C., Avi Gopher, and Ran Barkai. 2009. Cooperative hunting and and significance of the Mode 3 (prepared core) technologies. In The ancient meat sharing 400–200 kya at Qesem Cave, Israel. Proceedings of the National human occupation of Britain. Nick Ashton, Simon G. Lewis, and Chris B. Academy of Sciences of the USA 106:13207–13212. Stringer, eds. Pp. 53–66. Amsterdam: Elsevier. Stiner, Mary C., and Steven Kuhn. 2009. Paleolithic diet and the division of Wiessner, Polly. 1983. Style and social information in Kalahari San projectile labor in Mediterranean Eurasia. In The evolution of hominid diets: integrating points. American Antiquity 48:253–276. approaches to the study of Paleolithic subsistence. M. Richards and J. J. Hublin, Wurz, Sarah. 2013. Technological trends in the Middle Stone Age of South eds. Pp. 155–167. New York: Springer. Africa between MIS 7 and MIS 3. Current Anthropology 54(suppl. 8):S305– Stiner, Mary C., Steven Kuhn, Todd A. Surovell, Paul Goldberg, Amy V. S319. Margaris, Liliane Meignen, Stephen Weiner, and Ofer Bar-Yosef.2005. Bone, Wynn, Thomas, and Frederick Coolidge. 2004. The expert Neandertal mind. ash, and shell preservation in Hayonim Cave. In The faunas of Hayonim Journal of Human Evolution 46:467–487. cave, Israel. Mary C. Stiner, ed. Pp. 59–79. American School of Prehistoric Zilha˜o, Joa˜o, Diego E. Angelucci, Ernestina Badal-Garcı´a, Francesco d’Errico, Research Bulletin 48. Cambridge, MA: Peabody Museum of Archaeology Flore´al Daniel, Laure Dayet, Katerina Douka, et al. 2010. Symbolic use of and Ethnology, Harvard University. marine shells and mineral pigments by Iberian Neandertals. Proceedings of Stiner, Mary C., Natalie D. Munro, and Todd A. Surovell. 2000. The tortoise the National Academy of Sciences of the USA 107(3):1023–1028. Current Anthropology Volume 54, Supplement 8, December 2013 S269

The Evolutionary Implications of Variation in Human Hunting Strategies and Diet Breadth during the Middle Stone Age of Southern Africa

by Jamie L. Clark and Andrew W. Kandel

CAϩ Online-Only Material: Supplement A

In this paper, we analyze faunal data from southern Africa in order to explore the nature and extent of variability in human hunting strategies and diet breadth during the Middle Stone Age (MSA). Our analysis incorporates data from eight sites that span marine isotope stages (MIS) 6–3 (∼170–40 ka). The sample includes both coastal and inland sites; while it primarily derives from cave and rock shelter contexts (Blombos, Die Kelders, Diepkloof, Klasies River, Pinnacle Point, Sibudu, and Ysterfontein), we also include one open-air site (Florisbad). The data indicate marked changes in subsistence over time—of particular note is a spike in the exploitation of small ungulates and other small mammals during MIS 4. Trends in shellfish utilization also suggest an increasing dietary breadth over time, although shellfish consistently represent only a small portion of the overall diet. We close with a discussion of several causal mechanisms (environment, demography, technology, and cognition) that could account for the patterning evidenced in the data. While environmental change appears to play an important role in shaping the character of the data, we argue that no single causal factor can account for the full range of variability.

Introduction 2011). On the Eurasian side, this is due in large part to the long-held view that the MP was a period of relative stasis in The period known as the Middle Paleolithic (MP) in Eurasia human behavior (see Kuhn and Hovers 2006 for a detailed and the Middle Stone Age (MSA) in Africa (roughly spanning discussion). While somewhat greater attention has been paid 300–30 ka) was a critical one in the development of our to variation over both space and time within the MSA (e.g., species. This period witnesses the rise and spread of anatom- Clark 1993; McBrearty and Brooks 2000), particularly as ex- ically modern humans (AMH) and also incorporates both the pressed in the lithic record, the view of MSA behavior as florescence—and demise—of the Neanderthals. Yet our largely static has also been fairly well entrenched (see Klein knowledge about the nature and extent of variability in human 2009 for a more detailed discussion). This is particularly the behavior during this period remains limited despite a growing case when considering subsistence behavior; Klein (2009) ar- recognition that a deeper understanding of this variability may gued that “in most instances where controlled comparisons be critical to answering larger questions about the evolution are possible, differences in species abundance among Mous- of human culture and the ultimate success of AMH over terian/MSA sites or between Mousterian/MSA sites . . . appear archaic populations such as the Neanderthals (e.g., Clark to reflect differences in site environment rather than in oc- 2011; Kuhn and Hovers 2006 and references therein; Shea cupant behavior” (553). Our goals for this paper are twofold. Our primary goal is to present the results of a large-scale attempt to explore var- Jamie L. Clark is Assistant Professor in the Department of iation in human hunting behavior and diet breadth in the Anthropology at the University of Alaska Fairbanks (P.O.Box 757720, MSA of southern Africa. To this end, we have compiled faunal Fairbanks, Alaska 99775-7720, U.S.A. [[email protected]]). data from eight sites, including both coastal and inland lo- Andrew W. Kandel is Project Archaeologist in the Role of Culture in Early Expansions of Humans (ROCEEH) at the Heidelberg calities; while we hoped to incorporate material from the ∼ Academy of Sciences and Humanities (Ru¨melinstraße 23, 72070 entire span of the MSA ( 300–30 ka; MIS 8–3), the nature Tu¨bingen, Germany). This paper was submitted 3 VII 13, accepted of the available data forced us to restrict our study to materials 7 VIII 13, and electronically published 8 XI 13. spanning ∼170–40 ka. Our second goal is to evaluate the

᭧ 2013 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved. 0011-3204/2013/54S8-0009$10.00. DOI: 10.1086/673386 S270 Current Anthropology Volume 54, Supplement 8, December 2013 relative importance of environmental, demographic, tech- Mediterranean are applicable across a wider geographic re- nological, and cognitive changes on the observed variability gion. However, a recent study by Delagnes and Rendu (2011) in subsistence behavior. based on lithic and faunal materials recovered from 68 MP assemblages in southwestern France (spanning MIS 6–3) doc- umented clear (and seemingly interrelated) shifts in tech- Previous Research nology and subsistence among late Neanderthals in the region, The Eurasian record. The majority of research on variation particularly after MIS 4. The authors were unable to identify in hunting behavior within the Late Pleistocene (and its po- a single factor that could account for the documented vari- tential evolutionary implications) has focused on comparisons ability, and ultimately they stressed the importance of mul- between the MP and Upper Paleolithic (UP) in Eurasia. The ticausal explanatory models for understanding behavioral di- most influential work in this regard has been that of Mary versity in the archaeological record. Stiner and colleagues (e.g., Stiner 2002, 2006; Stiner and Kuhn 2006; Stiner and Munro 2002; Stiner, Munro, and Surovell The African record. Just as research on variation in the 2000), who utilized data from across the Mediterranean to Eurasian record has primarily focused on comparisons of the demonstrate a marked shift in subsistence around the time MP and UP, so too has research on the African record largely of the MP-UP transition, most notably as related to the ex- centered on comparisons of the MSA and Later Stone Age ploitation of small game. While MP populations primarily (LSA). Best known in this respect is the work of Klein and exploited slow-moving, slow-growing small game such as tor- colleagues (e.g., Klein 1979, 2001, 2009; Klein and Cruz-Uribe toises and marine mollusks, UP hunters focused on agile, fast- 1996, 2000; Steele and Klein 2009), who have argued that maturing animals such as hares and birds. This change MSA populations were less effective hunters than their LSA marked what Stiner considered to be a categorical shift in counterparts. Utilizing data from several MSA and LSA as- human predator-prey dynamics (Stiner 2006). However, this semblages, Klein demonstrated that LSA sites generally pre- pattern is not necessarily a universal one. Work by Adler and serve higher frequencies of dangerous prey (particularly buf- colleagues in the Caucasus (Adler and Bar-Oz 2009; Adler et falo; e.g., Klein and Cruz-Uribe 2000). He interprets these al. 2006) demonstrated relatively little change in human sub- data as indicating that MSA foragers were limited in their sistence behavior across the MP-UP transition. Similarly, ability to capture certain types of ungulate prey. However, when Gaudzinski-Windheuser and Niven (2009) compiled this argument has been contested; most notably, Faith (2008) faunal data from several Paleolithic sites in NW Europe, they utilized a wider range of samples and a variety of statistical found that subsistence behavior was relatively consistent be- methods to argue that dangerous game (including buffaloes tween the MP and earliest UP, with the most marked changes and wild pigs) are equally abundant in both the MSA and in subsistence occurring later in the UP. LSA. Although Weaver, Steele, and Klein (2011) critiqued Large-scale, specifically directed analyses of changes in hu- Faith’s methodology, their reanalysis nonetheless corroborates man subsistence behavior within the MP are rare (exceptions Faith’s primary result that MSA sites do not appear to be include Delagnes and Rendu 2011; Gaudzinski 2006; Stiner dominated by eland or be deficient in dangerous game com- 2006, 2013; Stiner and Kuhn 1992). Outside the long-held pared with LSA sites (see Faith 2011b). notion of behavioral stasis during this period, there are other, Steele and Klein (2009) presented the results of a much more practical reasons why such studies are uncommon. more comprehensive comparison of MSA and LSA subsis- Gaudzinski (2006) highlighted some of these reasons, includ- tence. In this study, they applied the framework used for ing a dearth of faunal assemblages subjected to detailed taph- exploring variation in subsistence strategies in the Eurasian onomic analyses, which raises concerns about the compara- MP and UP to the MSA/LSA record from coastal South Africa, bility of data sets. Coarse chronological resolution has also utilizing data from nine archaeological sites. Although fo- prevented the development of “far-reaching conclusions” cusing on the coast, Steele and Klein deliberately included concerning evolutionary trends in subsistence behavior dur- sites from different ecological zones, arguing that combining ing the MP (Gaudzinski 2006). these zones made it more likely that any patterns identified Stiner (2006, 2013) looked at variation in MP subsistence in the data reflected changes in human behavior rather than ecology in the Mediterranean region. She found that very few climatic/environmental variation. trends were apparent within the MP with the possible excep- Steele and Klein (2009) looked for variation in several di- tion of mild harvesting pressure on slow-turnover prey pop- mensions: (1) large-game exploitation (using both prey mor- ulations after 50 ka. Although limiting their study to change tality profiles and measures of processing intensity), (2) the over time at a single site (Kebara Cave, Israel), work by Speth abundance and diversity of small game (particularly small, and Clark (2006) was consistent with these results; they argued slow game such as tortoises and shellfish vs. small, fast game that during the last ∼12,000 years of the MP, the inhabitants such as hares and birds), and (3) shellfish and tortoise size, of Kebara overhunted their large-game resources while also which can serve as an indicator of collection intensity. They intensifying their use of lower-ranked gazelle and fallow deer. concluded that LSA populations consumed a wider variety of It is not yet clear whether the results obtained for the resources and exploited the available resources more inten- Clark and Kandel Variation in Middle Stone Age Hunting Strategies S271 sively than their MSA counterparts, and they suggested that Materials these differences related both to larger and denser human Criteria for Inclusion populations and perhaps to the development of LSA tech- nologies. Given that our goal is to explore variation in human hunting As was the case for the Eurasian MP, larger-scale attempts strategies and dietary breadth over time, we limited our anal- to explore variation in human hunting behavior within the ysis to well-dated sites that preserved both faunal and lithic MSA have been relatively limited (exceptions include Dus- remains. A further requirement was that the assemblages were seldorp 2012; Lombard and Clark 2008; Thompson 2010b). primarily anthropogenic in origin (see fig. 1 for a map of sites There are a number of reasons for this. First, as was the case discussed in the text and table A1 in CAϩ online supplement for the MP, this is due in part to notions of relative stasis in A for a list of excluded sites). As such, we excluded remains human behavior during the MSA (e.g., Klein 2009). Second, from well-known MSA sites such as Boomplaas (Faith 2011a), the number of relatively well-published MSA faunal assem- Equus Cave (Klein, Cruz-Uribe, and Beaumont 1991), the blages was quite limited until recently. Just as the scale of “old collections” at Florisbad (Brink 1988; Kuman and Clarke research into the MSA has markedly increased in the past two 1986), Herolds Bay Cave (Brink and Deacon 1982), Kalkbank decades, so too has the number of sites with published faunal (Hutson and Cain 2008), and Plovers Lake (de Ruiter et al. data. Since 2000, details on the fauna from six key MSA sites 2008), which appeared to be either natural death sites or (Blombos, Die Kelders, Diepkloof, Pinnacle Point 13B, Si- accumulated primarily by agents other than humans. budu, and Ysterfontein) have been published, with data from We also chose to limit our analysis to sites for which mam- several other sites expected in the near future (e.g., Boom- malian faunal data were reported as number of identified plaas, Kudu Koppie, Melikane, and Sehonghong). Finally, specimens (NISP) rather than minimum number of individ- broader-scale comparisons of MSA faunal assemblages have uals (MNI). While a detailed discussion of the relative merits been hindered by the use of different analytical and quantifi- (and faults) of these quantification methods is beyond the cation methods such that all data sets are not necessarily scope of this paper, our choice was based more on practical directly comparable (see Thompson 2010b for a detailed dis- considerations. We wanted to limit our analysis to a single cussion of some of the theoretical, empirical, and method- quantification method, and a larger proportion of sites pre- ological obstacles facing those attempting larger comparative sented mammalian faunal data in NISP form. As such, we studies of MSA subsistence). The view of MSA subsistence behavior as static was so entrenched that as recently as 2008, Lombard and Clark fo- cused their attention on documenting whether variation in human hunting behavior even existed over the course of the MSA. Combining lithic and faunal data sets, they summarized current interpretations about hunting behavior spanning from the lowermost pre–Still Bay at Blombos (∼100 ka) through the post–Howieson’s Poort at Sibudu (∼58 ka); however, they did not attempt to identify any broader directional trends in the data. Thompson (2010b) explored variation in faunal exploita- tion at two MSA sites that preserve deposits spanning ∼170– 73 ka. She personally analyzed the faunal material from Blom- bos and Pinnacle Point 13B, promoting comparability of the two data sets. Looking at variation in dietary composition, processing, and transport behavior, she concluded that sub- sistence decisions were not uniform over time or space. She did not identify any clear temporal trends but rather proposed that the identified variation may be closely related to differ- Figure 1. Map of MSA sites discussed in the text. Dots indicate ences in site context. sites that were included in the study; Xs indicate sites that were In light of the recent publication of faunal data from a excluded. A11 p Apollo 11, BBC p Blombos Cave, BC p number of well-excavated MSA sites preserving extensive, Border Cave, BFA p Bundu Farm, BPA p Boomplaas Cave, p p p well-stratified deposits, the time is ripe for a larger-scale con- BRS Bushman Rock Shelter, DK1 Die Kelders 1, DRS p p sideration of variation in MSA subsistence behavior over time. Diepkloof Rock Shelter, EQC Equus Cave, FLO Florisbad, HB p Herold’s Bay, KB p Kalkbank, KK p Kudu Koppie, While recognizing that this type of study is not entirely KRM p Klasies River Main Site, MLK p Melikane, PLL p straightforward (see below for a more detailed discussion of Plover’s Lake, PP13B p Pinnacle Point Cave 13B, SCB p Strath- the caveats), we believe that the available data are sufficiently alan Cave B, SEH p Sehonghong, SIB p Sibudu, YFT1 p comparable to produce meaningful results. Ysterfontein 1. Table 1. Mammalian assemblages included in the analysis and grouped by marine isotope stage (MIS)

Age range NISP Assemblage (ka) (mammals) References MIS 6: PP13B LB Silt 349–152 6 Marean et al. 2010; Rector and Reed 2010 PP13B LC-MSA Lower 174–153 2,826 Marean et al. 2010; Thompson 2010a PP13B Lower DB Sand 164–159 147 Marean et al. 2010; Rector and Reed 2010; Thompson 2010a MIS 5: YFT1 Lower 140–124 1,222 Avery et al. 2008 YFT1 Middle 128–114 266 Avery et al. 2008 YFT1 Upper 135–123 1,836 Avery et al. 2008 PP13B LBG Sand 134–94 1,391 Marean et al. 2010; Rector and Reed 2010; Thompson 2010a PP13B LC-MSA Middle/Upper 130–120 314 Marean et al. 2010; Thompson 2010a FLO Unit F 127–115 268 Brink 1987; Kuman, Inbar, and Clarke 1999 DRS Lower MSA 118–90 300 Steele and Klein 2013; Tribolo et al. 2013 DRS MSA-Mike n.d. 63 Steele and Klein 2013; Tribolo et al. 2013 DRS Pre-SB Lynn 110–90 37 Steele and Klein 2013; Tribolo et al. 2013 DRS Still Bay 119–99 166 Steele and Klein 2013; Tribolo et al. 2013 DRS Early HP 119–95 290 Steele and Klein 2013; Tribolo et al. 2013 KRM LBS 115–110 419 Feathers 2002; Wurz 2002; van Pletzen 2000 KRM Cave 1 (38/LBS) 115–110 853 Feathers 2002; R. G. Klein, unpublished data PP13B Lower Roof Spall 114–106 1,510 Marean et al. 2010; Rector and Reed 2010; Thompson 2010a KRM Cave 1 (37/RBS) 110–95 1,173 Feathers 2002; Wurz 2002; R. G. Klein, unpublished data KRM Cave 1 (17/SASL) 110–95 1,493 Feathers 2002; Wurz 2002; R. G. Klein, unpublished data KRM Cave 1 (16/SASU) 110–95 1,794 Feathers 2002; Wurz 2002; R. G. Klein, unpublished data KRM Cave 1 (15/SAS) 110–95 944 Feathers 2002; Wurz 2002; R. G. Klein, unpublished data KRM SASb 110–95 1,938 Feathers 2002; van Pletzen 2000; Wurz 2002 BBC M3 105–91 287 Henshilwood et al. 2011; Thompson and Henshilwood 2011 BBC M3 105–91 1,212 Henshilwood et al. 2001, 2011 PP13B Upper DB Sand 102–91 2,950 Marean et al. 2010; Rector and Reed 2010; Thompson 2010a PP13B LB Sand 2 102–91 4 Marean et al. 2010; Rector and Reed 2010 PP13B SB Sand/ Upper Roof Spall 98–91 5,330 Marean et al. 2010; Rector and Reed 2010; Thompson 2010a DRS MSA-Jack 98–80 114 Steele and Klein 2013; Tribolo et al. 2013 KRM Cave 1 (14/SASR) 95–85 3,357 Feathers 2002; Wurz 2002; R. G. Klein, unpublished data KRM SASm 95–72 884 Jacobs et al. 2008a; van Pletzen 2000 KRM SASt 95–72 205 Jacobs et al. 2008a; van Pletzen 2000 PP13B LB Sand 1 94–91 763 Marean et al. 2010; Rector and Reed 2010; Thompson 2010a DRS Intermediate HPa 94–57 955 Steele and Klein 2013; Tribolo et al. 2013 BBC M2 lower 91–77 121 Henshilwood et al. 2011; Thompson and Henshilwood 2011 BBC M2 upper 80–74 468 Henshilwood et al. 2011; Thompson and Henshilwood 2011 BBC M2 91–74 1,167 Henshilwood et al. 2001, 2011 BBC M1 74–68 934 Henshilwood et al. 2011; Thompson and Henshilwood 2011 BBC M1 74–68 1,727 Henshilwood et al. 2001, 2011 MIS 4: KRM Cave 1 (13/WS) 78–63 186 d’Errico, Garcia Moreno, and Rifkin 2012; R. G. Klein, un- published data KRM Upper 72–58 1,017 Jacobs et al. 2008a; van Pletzen 2000 DK1 Layer 15 70–60 336 Feathers and Bush 2000; Klein and Cruz-Uribe 2000 DK1 Layer 14 70–60 2,126 Feathers and Bush 2000; Klein and Cruz-Uribe 2000 DK1 Layer 13 70–60 374 Feathers and Bush 2000; Klein and Cruz-Uribe 2000 DK1 Layer 12 70–60 960 Feathers and Bush 2000; Klein and Cruz-Uribe 2000 DK1 Layer 11 70–60 458 Feathers and Bush 2000; Klein and Cruz-Uribe 2000 DK1 Layer 11 70–60 541 Feathers and Bush 2000; Marean et al. 2000 DK1 Layer 10 70–60 1,996 Feathers and Bush 2000; Klein and Cruz-Uribe 2000 DK1 Layer 10 70–60 2,424 Feathers and Bush 2000; Marean et al. 2000 DK1 Layer 9 70–60 4,414 Feathers and Bush 2000; Klein and Cruz-Uribe 2000 DK1 Layer 8 70–60 107,939 Feathers and Bush 2000; Klein and Cruz-Uribe 2000 DK1 Layer 7 70–60 9,200 Feathers and Bush 2000; Klein and Cruz-Uribe 2000 DK1 Layer 6 70–60 56,476 Feathers and Bush 2000; Klein and Cruz-Uribe 2000 DK1 Layer 4/5 70–60 15,414 Feathers and Bush 2000; Klein and Cruz-Uribe 2000 SIBHP 67–60 5,463 Jacobs et al. 2008a; J. L. Clark, unpublished data MIS 3: SIB Post HP MSA 2 62–56 1,090 Jacobs et al. 2008b; J. L. Clark, unpublished data SIB Post HP MSA 1 62–56 843 Jacobs et al. 2008b; J. L. Clark, unpublished data Clark and Kandel Variation in Middle Stone Age Hunting Strategies S273

Table 1 (Continued) Age range NISP Assemblage (ka) (mammals) References DRS Late HP 57–47 1,023 Steele and Klein 2013; Tribolo et al. 2013 DRS Post HP n.d. 481 Steele and Klein 2013; Tribolo et al. 2013 SIB Late MSA 52–44 1,985 Jacobs et al. 2008b; Wadley, Plug, and Clark 2008 SIB Final MSA 42–35 303 Jacobs et al. 2008b; Wadley, Plug, and Clark 2008 Note. Age ranges are best estimates based on the data found in the literature. Assemblages are ordered chronologically except in cases that would reverse their stratigraphic sequence. NISP p number of identified specimens; n.d. p no date. a The Intermediate HP at DRS contains layers dating to both MIS 5 and MIS 4. Because the faunal data were provided only for the unit as a whole and because the majority of the dates fall within MIS 5, we included it in the MIS 5 sample. See CAϩ online supplement A. excluded sites with only MNI data—for example, the Wendt cluded here comes from excavations that employed modern 1968–1972 excavations at Apollo 11 (Thackeray 1979), Border recovery strategies, including the use of fine mesh sieves (!5 Cave (Klein 1977), Kudu Koppie (Pollarolo et al. 2010), and mm), and thus recovery biases should be limited. By restrict- the Opperman 1988–1991 excavations at Strathalan Cave B ing our analysis to sites providing NISP counts for mam- (Brink’s data in Opperman 1996). Although Klein (1976) only malian fauna, we avoid potential issues arising from the use presented MNI data from Cave 1 at Klasies River, he kindly of multiple quantification methods and also avoid any po- provided us with NISP counts for the same sample, allowing tential complications related to the nonstandardized calcu- us to include these results in our analysis. However, because lation of MNI values. Although more thorough taphonomic researchers primarily presented MNI counts for the molluscan analyses are clearly warranted, we are confident that humans fauna, we report shellfish data using this quantification were the major contributors to the faunas under consider- method. ation. Finally, similar to Steele and Klein (2009), we feel that We excluded other sites because faunal remains were as- the inclusion of remains from a variety of ecological zones signed to taxa in a way that was inconsistent with the other (including inland and coastal sites as well as sites from sum- data sets (e.g., while NISP data are available for the new mer and winter rainfall areas) makes it more likely that any excavations at Apollo 11, Vogelsang et al. [2010] utilized cat- trends identified in the data signify real differences in human egories such as “small or medium antelope,” making it dif- behavior rather than simply reflecting environmental change. ficult to compare data). Finally, the nature of the available dates meant that some sites could not be assigned to a specific Methods marine isotope stage (MIS); this was the case for Bundu Farm (Brink’s data in Kiberd 2006; Hutson 2012) and Bushman For all analyses reported here, we grouped the assemblages Rock Shelter (Badenhorst and Plug 2012; Louw 1969; Plug by MIS; this made the resulting data easier to manage and 1981). had the added benefit of making evaluating whether the po- tential variance in the data was driven primarily by climate change more straightforward. The number of mammalian Sites/Assemblages Included in the Study assemblages dated to MIS 5 and 4 (n p 35 and 16, respec- Based on these criteria, the total vertebrate sample comprises tively) are larger than those for MIS 6 and 3 (n p 3 and 6, 60 assemblages from eight sites (table 1): Blombos Cave respectively); in fact, the sample from MIS 6 is limited to one (BBC), Die Kelders Cave 1 (DK1), Diepkloof Rock Shelter site (PP13B), while that from MIS 3 is limited to two (DRS (DRS), Florisbad (FLO), Klasies River Main Site (KRM), Pin- and SIB). For shellfish, MIS 5 also contained the largest num- nacle Point Cave 13B (PP13B), Sibudu Cave (SIB), and Yst- ber of assemblages (n p 22 ), with MIS 6, 4, and 3 represented erfontein 1 (YFT1). Shellfish data were available for 28 as- by two assemblages each. Given the nature of the sample and semblages from five of these sites (table 2): BBC, DRS, KRM, the variation in sample size across MIS, our analyses are by PP13B, and YFT1. More detailed information about site con- necessity more qualitative than quantitative in nature. text and the faunal remains can be found in supplement A. We considered grouping the assemblages by archaeological While we recognize that these sites may have been excavated cultures in order to explore other potential dimensions of using different recovery strategies and that the zooarchaeol- variation in hunting behavior, as some proportion of the stone ogists responsible for producing and publishing the faunal tools likely served as weapon tips. However, this consideration data may have employed differing analytical strategies, we proved to be untenable in large part because there are too contend that the assemblages included here are sufficiently few conventions for assigning material to specific MSA cul- comparable so as to provide a reliable basis for a consideration tures. Especially before and after the better-known Still Bay of variation in human hunting behavior during the course of (SB) and Howieson’s Poort (HP) phases of the MSA, lithic the MSA. With a single exception—the Singer and Wymer assemblages are highly variable and trends are poorly under- (1982) excavations at Klasies River—all of the material in- stood (see Wurz 2013). S274 Current Anthropology Volume 54, Supplement 8, December 2013

Table 2. Shellfish assemblages included in the analysis grouped by marine isotope stage (MIS)

Assemblage MNI Weight (g) kcal People days References MIS 6: PP13B Lower LC-MSA 28 156 64 .0 Jerardino and Marean 2010 PP13B West Lower Sand 1 2 1 .0 Jerardino and Marean 2010 MIS 5: YFT1 Lower 7,087 109,744 49,149 24.6 Avery et al. 2008 YFT1 Middle 1,231 14,109 7,404 3.7 Avery et al. 2008 YFT1 Upper 1,540 25,870 13,146 6.6 Avery et al. 2008 PP13B West Middle Sand 11 30 18 .01 Jerardino and Marean 2010 PP13B Middle LC-MSA 53 398 175 .1 Jerardino and Marean 2010 PP13B Upper LC-MSA 35 156 64 .03 Jerardino and Marean 2010 DRS Lower MSA . . . 22 17 .01 Steele and Klein 2013 DRS MSA-Mike . . . 1 1 .001 Steele and Klein 2013 DRS Pre-SB Lynn . . . 17 14 .01 Steele and Klein 2013 DRS Still Bay . . . 16 10 .005 Steele and Klein 2013 DRS Early HP . . . 69 35 .02 Steele and Klein 2013 KRM LBS (MSA I) . . . 6,577 ...... Langejans et al. 2012 PP13B Lower Roof Spall 238 2,066 846 .4 Jerardino and Marean 2010 KRM SAS (MSA II) . . . 4,656 ...... Langejans et al. 2012 BBC M3 2,692 98,218 75,456 37.7 Langejans et al. 2012 PP13B West Upper Sand 49 369 206 .1 Jerardino and Marean 2010 PP13B Upper Roof Spall 234 2,292 1,012 .5 Jerardino and Marean 2010 PP13B SB Sand 68 823 362 .2 Jerardino and Marean 2010 DRS MSA-Jack . . . 111 63 .03 Steele and Klein 2013 DRS Intermediate HP . . . 92 45 .02 Steele and Klein 2013 BBC M2 3,118 24,253 15,867 7.9 Langejans et al. 2012 BBC M1 3,203 24,363 14,801 7.4 Langejans et al. 2012 MIS 4: KRM Upper (HP) 605 ...... Langejans et al. 2012 KRM Upper (MSA III) 213 ...... Langejans et al. 2012 MIS 3: DRS Late HP . . . 996 630 .3 Steele and Klein 2013 DRS Post-HP . . . 1065 783 .4 Steele and Klein 2013 Notes. Nutritional values for shellfish (kcal, people-days) calculated according to Buchanan (1988). See text for details on calculations. Assemblages are ordered chronologically except in cases that would reverse their stratigraphic sequence.

Exploring Variation in Hunting Strategies and Diet Breadth time with the assumption that the smallest ungulates would be of the lowest rank. Bovids are the most common ungulates Prey choice models predict that top ranked prey (those with in the region under consideration; given the number of dif- a higher energetic yield relative to search, pursuit, and han- ferent bovid species present in southern Africa and the re- dling costs) will always be taken on encounter; if encounter sulting difficulty in identifying fragmentary remains to spe- rates with preferred prey decline, hunters are expected to cies, the majority of the bovid remains in MSA assemblages broaden their diets by including more lower-yield resources are assigned to size class (see table 3 for details); these classes (e.g., Bird and O’Connell 2006; Kaplan and Hill 1992; Lupo have also been utilized more generally (cf. Thompson 2010a; 2007). Within this framework, there are a number of ways Thompson and Henshilwood 2011). For this analysis, we in- we can utilize the available data. Ethnographic and empirical cluded all ungulate remains identifiable to species as well as studies have shown that when acquired individually, prey rank those identifiable to size class. scales closely with body size such that large prey are generally higher ranked than smaller animals (Broughton and Grayson 1993; Hawkes, Hill, and O’Connell 1982; Kelly 1995; Lupo 2007; Lupo and Schmitt 2005). As such, an increase in the Large versus Small (!4.5-kg) Mammals exploitation of small (and presumably lower-ranked) animals may indicate an increase in dietary breadth. An increase in the exploitation of small game may also reflect an expansion of dietary breadth. Although work by Stiner and colleagues (e.g., Stiner, Munro, and Surovell 2000) fo- cused not just on the relative frequency of small game com- Ungulate Size Classes pared with larger game but more specifically on changes in The first way we approached the data was to explore variation the types of small game present in any given assemblage (i.e., in the exploitation of the various ungulate size classes over slow-moving/slow-growth prey vs. agile/fast-maturing prey), Clark and Kandel Variation in Middle Stone Age Hunting Strategies S275

Table 3. Size classes utilized for southern African faunal assemblages

Size classes Weight Brain (1974) Klein (1976) (kg) Species (list not all inclusive) Size 1 Small 4.5–23 Blue duiker, common duiker, klipspringer, steenbok Size 2 Small-medium 23–84 Mountain reedbuck, bushbuck, springbok, impala Size 3 Large-medium 85–295 Red hartebeest, wildebeest, roan antelope, kudu, zebra Size 4 Large 296–900 African buffalo, eland Size 5 Very large 1900 Giant buffalo (extinct) the available data limited us to a more general comparison cause the wild pigs are known to be aggressive and would of small versus larger game. Furthermore, because data on have been more dangerous to hunt than the bovids. The two nonmammalian game (such as tortoises, birds, and fish) were suids occupy different habitat types, with warthogs occupying not consistently available, we restricted our analysis to small more open environments and bushpigs occupying closed en- mammals. vironments. Thus variation in the relative frequency of suids Given that 4.5 kg is a common cutoff for small game among should not simply be a reflection of environmental change. zooarchaeologists working in southern Africa (cf. Marean et al. 2000; Thompson 2010a; Thompson and Henshilwood 2011), we included species with an average weight of less than Use of Nonvertebrate Resources: Shellfish Exploitation 4.5 kg in our “small mammal” category but excluded micro- mammal remains, defined as less than 0.75 or 0.3 kg, de- An increase in the exploitation of nonvertebrate resources pending on the analyst. We also included generic “small mam- may also indicate an expansion of dietary breadth. Here, we mal” NISP counts if these were clearly defined as representing are particularly interested in looking at variation in shellfish animals of the appropriate size. For this particular analysis, exploitation during the MSA. Shellfish represent a predictable we excluded those assemblages for which data on small mam- resource that is easy to collect, although they have a high mals were not provided. processing cost relative to the number of calories provided. Shellfish appear as early as ∼164 ka in the southern African MSA (Marean et al. 2007). While people collected shellfish in what often appears to be large amounts, the calories that Dangerous versus Docile Prey shellfish provide are surprisingly low compared with verte- A third approach to exploring changes in diet breadth—and brate resources (Buchanan 1988). This suggests that shellfish hunting strategies more broadly—involves a consideration of represented just one part of a varied MSA subsistence strategy. variation in the exploitation of dangerous or aggressive game. We explored variation in the diversity of shellfish over time As previously discussed, Klein has proposed that the exploi- by reviewing the published shellfish data and combining the tation of dangerous game serves as an indicator of hunting shells into groups (see table A2). Because we are interested ability (and thus perhaps cognitive capacity; Klein 2001; Klein in examining human diet, we excluded incidental species that and Cruz-Uribe 1996). On a broader level, though, one could were not likely exploited as food. Of the five sites for which argue that dangerous/aggressive game would be comparatively shellfish data are available, DRS and YFT1 represent South low-ranked resources, and thus an increase in the exploitation Africa’s cooler west coast, while BBC, KRM, and PP13B come of such taxa could serve as an indicator of increased dietary from the warmer south coast. Because of differences in ocean breadth (Clark 2009, 2011). This is because capturing this temperature along each coast, some shell species are not pres- game could potentially pose a greater risk (and thus would ent in both regions. Therefore, we examined the data from be higher cost) than the pursuit of more docile game. each coast separately. In exploring this issue, we began by comparing the relative We also explored variation in the relative contribution of frequencies of eland (Taurotragus oryx) and buffalo (Syncerus shellfish to the diet. To accomplish this, we converted shell caffer and Pelorovis antiquus) not only because this compar- weights into nutritional values to evaluate the dietary im- ison has been a popular one among those looking at variation portance of shellfish, although weights were not available for in human subsistence behavior (e.g., Dusseldorp 2012; Faith KRM. The nutritional values stem from Buchanan’s (1988) 2008; Klein 1979, 2001) but also because the species in ques- investigation of the dietary contribution of marine and ter- tion have broadly similar habitat requirements, and as such, restrial resources. He reports that two limpet species (Cymbula variation in relative frequency of these taxa should not simply granatina and Scutellastra granularis) yield an average of 350 reflect changes in environmental conditions over time. kJ of energy for each 100 g of shell present, whereas black We also compared the relative frequency of suids—warthog mussels (Choromytilus meridionalis) generate 150 kJ from the (Phacochoerus africanus) and bushpig (Potamochoerus larva- same shell mass. Because nutritional data were not available tus)—with the frequency of similar sized bovids (Size 2) be- for other shellfish, we assume that gastropods (including other S276 Current Anthropology Volume 54, Supplement 8, December 2013 limpet species) are similar to limpets and that bivalves are Results similar to the black mussels analyzed by Buchanan. We also assume a modern nutritional requirement of 2,000 kcal (8,370 Variation in Ungulate Size Classes over Time kJ) per person per day for calculating person-days of food. Figure 2a presents data on variation in the frequency of the The third way we examined variation in subsistence was various ungulate size classes by MIS in both graphical and to look at the relative frequency of ungulates to shellfish. For tabular format. While these data appear to show a very consistency in our comparison, we only used data from as- marked increase in the smallest ungulates in MIS 4, this pat- semblages that contained both ungulates and shellfish. We tern is potentially driven by a single assemblage, DK1 Layer recognize the potential problems inherent in comparing un- 6, which had an NISP of more than 18,000 Size 1 ungulates. gulate NISP with shellfish MNI values. However, because we Given the fact that raptors have been implicated as a major are interested in analyzing change over time, this index pro- contributor to the small ungulate fauna in other layers at the vides a relative measure of the intensity with which people site (particularly DK1 Layer 10), we removed this assemblage utilized shellfish. from the analysis to see how the pattern would change. How- ever, we should note that new work by Armstrong (2013) indicates that humans contributed to the collection of both Other Methods for Exploring Variation in Subsistence small ungulates and other small game at the site. Figure 2b Two lines of evidence that have been commonly utilized when presents the results of this analysis. The same general trend exploring variation in hunting strategies and/or diet breadth is apparent; we still see a marked increase in the smallest could not be considered here. The first relates to changes in ungulates from MIS 5 to MIS 4 with a subsequent (and even prey mortality profiles; Klein and colleagues (Klein 1979, more marked) reduction in MIS 3. If we consider the smallest 2001; Klein and Cruz-Uribe 1996; Steele and Klein 2009) have ungulates to be the lowest-ranked prey, these data are con- largely utilized this data in discussions about the hunting sistent with an expansion of dietary breadth from MIS 6 to abilities of MSA versus LSA populations; for example, they MIS 4 followed by a decline in diet breadth in MIS 3. have argued that when MSA populations do capture large, Another pattern in these data can be seen in the relative dangerous game, they tend to capture very young or old in- proportion of large and very large ungulates (Size 3 and dividuals as opposed to prime-aged adults. Unfortunately, age larger), which, according to prey choice models, should be data were not consistently available, so we could not explore the highest-ranked prey. Comparatively speaking, large and variability along this dimension. very large ungulates are more frequent during the interglacial/ 1 More intensive carcass use and processing has also been interstadial periods (MIS 5 and 3; 50% in both cases) than cited as evidence for subsistence intensification (and thus in MIS 6 and 4. Given that the larger ungulates in the sample wider dietary breadth), for example, the exploitation of mar- also tend to be those that occupy more open environments, row from skeletal parts with low marrow utility (or from small this could imply that variation in the presence of the largest game) and the production of bone grease have relatively high ungulates is linked to environmentally driven changes in the processing costs relative to energetic yield (e.g., Binford 1978; availability of this game. Munro and Bar-Oz 2005). A consideration of variation in carcass use and processing requires not only skeletal element Variation in Large versus Small Mammals over Time frequency data but also detailed taphonomic data. Unfortu- Figure 3a presents data on the occurrence of large versus small nately, these data are not yet available for most of the assem- mammals. As was the case for the previous analysis, one sam- blages under consideration. ple from DK1 was large enough to potentially skew the re- sults—DK1 Layer 8 contained more than 100,000 Cape dune mole rat (Bathyergus suillus) bones. Again, in the absence of Controlling for Environmental Change: Browsers detailed taphonomic analysis, it is unclear whether these were versus Grazers accumulated predominantly by humans. While Klein and Finally, given that the data under consideration incorporate Cruz-Uribe (2000) point out that those layers at DK1 with materials from four MISs, including two glacial/stadial periods the most Cape dune mole rat bones are also those with evi- (MIS 6 and 4) and two interglacial/interstadial periods (MIS dence for less intensive human occupation, new work by 5 and 3), we were also interested in exploring what degree of Armstrong (2013) implicates humans in the collection of at variance in the data might be driven primarily by environ- least some of these remains. Figure 3b shows the results of mental change. To this end, we compared the relative fre- the analysis with the assemblage from DK1 Layer 8 removed. quency of browsers and grazers by MIS. Classification was Again, the general pattern remained the same. In fact, the based on Gagnon and Chew (2000) and Skinner and Chim- data show the same trend as the ungulate size-class data—a imba (2005). Note that there are a number of extinct species relatively small increase in small mammals from MIS 6 to in the sample. Because we cannot be certain about their di- MIS 5, a very marked increase in MIS 4, and then an even etary preferences, we exclude extinct taxa from the analysis. more pronounced decline in MIS 3. If small mammals are Clark and Kandel Variation in Middle Stone Age Hunting Strategies S277

Figure 2. a, Ungulate size-class data by MIS (in NISP). b, Ungulate size-class data by MIS (in NISP) with DK1 Layer 6 removed from analysis. considered low-ranked prey, this indicates an increase in di- There is also a more general question about what propor- etary breadth from MIS 6 to MIS 4. tion of the small mammals present in MSA sites was intro- In considering the implications of these data, two points duced by human activity. The small mammals from many of deserve mention. First, as previously discussed, our sample the assemblages under consideration have not been subjected from MIS 6 is limited to data from PP13B (Thompson 2008, to detailed taphonomic analyses; for example, Milo’s (1994, 2010a). However, Thompson conducted detailed taphonomic 1998) taphonomic study of the KRM fauna was limited to analyses of the PP13B fauna and argued that the relatively the larger mammal (14.5 kg) remains, as was Marean et al.’s low frequency of small game was a real phenomenon and not (2000) taphonomic study of the material from DK1 layers 10 simply a reflection of taphonomic factors unique to this site. and 11. Even when taphonomic analyses have been com- In fact, she drew particular attention to the relative dearth of pleted, the primary accumulating agent for the small mam- small game at PP13B especially compared with other MSA mals is not always clear. For example, although Thompson’s sites along the southwestern coast of South Africa (Thompson (2010a) work at PP13B indicated some human involvement 2010a). with the small fauna, direct evidence for human interaction S278 Current Anthropology Volume 54, Supplement 8, December 2013

Figure 3. a, Relative frequency of large versus small (!4.5-kg) mammals by MIS (in NISP). b, Relative frequency of large versus small (!4.5-kg) mammals by MIS (in NISP) with DK1 Layer 8 removed from study. with the small game component of the HP fauna from Sibudu 5–3 because the sample size for these taxa in MIS 6 was 1. remains elusive (J. L. Clark, unpublished data). All this being The relative frequency of buffalo declines from MIS 5 to MIS said, the fact that the small ungulates, which do seem to be 4 and then subsequently increases in MIS 3. This could be primarily accumulated by human activity, and small mammal interpreted as an indication of a reduction in diet breadth fauna show the same trends could indicate that humans also during MIS 4. However, the fact that MIS 5 and 3 have similar accumulated the latter. proportions of buffalo could also be a reflection of more open environments during the interglacial/interstadial periods. Figure 4b presents the data on the relative proportion of Prey Characteristics: Dangerous versus Docile Game suids versus similar sized bovids (Size 2). While the sample Figure 4 presents data on the relative frequencies of dangerous size from MIS 6 is quite small (n p 67 ), there is a steady versus docile prey. Figure 4a shows the proportion of eland increase in suids over time, although their relative frequency versus buffalo; note that the analysis was restricted to MIS is always low. This pattern could suggest a gradual increase Clark and Kandel Variation in Middle Stone Age Hunting Strategies S279

Figure 4. Dangerous versus docile prey. a, NISP counts for buffalo (including the extinct Pelorovis antiquus) versus eland by MIS. b, NISP counts for suids versus similar sized bovids (Size 2) by MIS. in intensification over time; however, these data are difficult in MIS 6 but predominate in MIS 5 and then drop in MIS to interpret for a variety of reasons. For example, because 4. As a means of controlling for the unequal sample size in suids are typically less common in coastal settings, we expect the data set, we plotted the data from a single site, KRM, them to occur at different frequencies around the various where we found a similar trend (fig. 6b). Given that MIS 5 sites included in the analysis. is an interglacial period when sea level was high and com- parable with the modern coastline, the prevalence of shellfish in MIS 5 is to be expected. Shellfish Data The data presented in table 2 regarding the dietary con- Figure 5 shows increasing diversification in shellfish exploi- tribution of shellfish deserve particular note. These data tation from MIS 6 to MIS 4. This is consistent with an increase indicate that the amount of shellfish in BBC M3, the largest in dietary breadth over time. Figure 6a presents data on the assemblage for which shell weights were available, would relative frequency of ungulates to shellfish. Shellfish are rare have provided subsistence for a maximum of 38 person- S280 Current Anthropology Volume 54, Supplement 8, December 2013

Figure 5. Distribution of shell groups by MIS and region (MNI). days (ca. 75,000 kcal), which is the caloric equivalent of ers dominate the glacial/stadial periods (MIS 6 and 4), with a single bontebok (Damaliscus dorcas). The next largest grazers dominating during the interglacial/interstadial periods sample, from YFT1 Lower, would have provided 25 person- (MIS 5 and 3). This suggests more trees and brush during days (ca. 49,000 kcal), or the energy equivalent of one the glacial periods and more open environments during the bushbuck (Tragelaphus scriptus), one steenbok (Raphicerus interglacials. These data fit within our current understanding campestris), and one hare (Lepus spp.). In contrast, the of paleoenvironmental conditions during the period under contribution from all 10 layers at PP13B is 1.5 person- consideration; for example, Chase (2010) utilized multiple days (ca. 3,000 kcal), the equivalent of four Cape dune lines of evidence to demonstrate the presence of cool and mole rats. Because the reported samples represent only a humid conditions during MIS 4 followed by dramatic arid- portion of the total excavated assemblage, the overall di- ification in MIS 3 (see also Ziegler et al. 2013). etary contribution would have been greater. Nonetheless, these data average hundreds, if not thousands, of years of shell accumulation, belying the impression that shellfish Discussion played a major dietary role during this period. When we consider the data presented above, some broader trends emerge (see fig. 8 for a visual summary of our results). No matter which way we approach the data, MIS 6 preserves Browser/Grazer Data the least evidence of subsistence intensification. Moreover, Figure 7a presents data on the relative proportion of browsers several lines of evidence are consistent with an expansion of and grazers over time. Because a majority of the browsers dietary breadth from MIS 6 to MIS 4. Most notable is the from MIS 4 came from a single species from the HP deposits aforementioned similarity in the data relating to the exploi- at Sibudu (Philantomba monticola, or blue duiker; n p tation of the smallest (Size 1) ungulates and the small mam- 1,962), we removed the Sibudu HP assemblage from the anal- mals, both of which would presumably have been low-ranked ysis; the resulting data are presented in figure 7b. While the game. The data indicate a slight increase in the exploitation total sample for MIS 6 is small (n p 13 ), in both cases, brows- of these species from MIS 6 to MIS 5 with a marked inten- Clark and Kandel Variation in Middle Stone Age Hunting Strategies S281

Figure 6. a, Frequency of shellfish versus ungulates (shellfish data expressed as MNI, ungulate as NISP) by MIS. b, Frequency of shellfish versus ungulates (shellfish data expressed as MNI, ungulate as NISP) at KRM. sification in their collection in MIS 4 and significant decline similar lines, if the exploitation of shellfish were primarily a in the focus on these prey in MIS 3. The increasing diversity reflection of subsistence intensification, we might expect to in shellfish exploitation also fits within a model of expanding see an increase in the relative frequency of shellfish from MIS dietary breadth from MIS 6 to MIS 4. 5 to MIS 4, and yet, this is not the case. However, in both However, not all of the data are consistent with these re- cases, the results may ultimately have more to do with en- sults. For example, if buffalo are considered to be lower ranked vironmental change than with subsistence intensification in than eland (given the higher pursuit costs for this aggressive that shifts in climate could have affected availability or ac- species), one might expect to see an increase in the exploi- cessibility of these prey. tation of buffalo from MIS 6 to MIS 4 and a decline in MIS Taken as a whole, the data indicate marked variance in 3. While data from MIS 6 are unavailable, we actually found subsistence over time. We now consider the potential role of the opposite trend, with a decline in the relative frequency of a number of different causal mechanisms: environment, de- buffalo from MIS 5 to MIS 4 and an increase in MIS 3. Along mography, technology, and cognition. S282 Current Anthropology Volume 54, Supplement 8, December 2013

Figure 7. a, Browser/grazer data by MIS (in NISP). b, Browser/grazer data by MIS (in NISP) with Sibudu HP removed from analysis.

Explaining Variance in the Data: Linkages to Linkages to Demographic Change? Environmental Change? It is notoriously difficult to identify changes in demography As already discussed, some of the variance in the data seems in the MSA. It is possible that the observed expansion of to be best explained as a reflection of changes in the envi- dietary breadth from MIS 6 to MIS 4 is related to an increase ronment. In fact, even the large increase in both the smallest in population pressure caused by either expanding population ungulates and small mammals in MIS 4 could signify sub- sizes or the concentration of existing populations. However, sistence intensification resulting from changes in the land- as previously discussed, this pattern could also be related to scape during this glacial period. As such, the changes in the environmental change, and distinguishing between these two data may largely reflect shifts in prey availability driven by alternatives is not currently possible. One potential way of climate change. However, we do not believe that environ- addressing this would be to explore the evidence for variation mental change can explain the full range of variance; for in shellfish and tortoise size across the MSA, because a decline instance, if this were the case, one might expect a higher in the size of these species has been linked to population frequency of small ungulates/mammals in MIS 6, and this is growth/pressure (e.g., Steele and Klein 2005/2006; Stiner et not the pattern that we see. al. 1999). Clark and Kandel Variation in Middle Stone Age Hunting Strategies S283

a large lithic assemblage, no diagnostic HP tools have been recovered (Thackeray 2000). This implies that we need to think more carefully about the functional significance of the technology of the HP, at least as far as it can be related to subsistence behavior. This is further underscored by new data from Diepkloof, where the HP spans MIS 5–3, periods in which we found a lesser focus on small game (Porraz et al. 2013; Tribolo et al. 2013).

Linkages to Variation in Cognition? We should begin here by admitting our own personal biases on this point; we assume that all of the populations repre- sented in this analysis likely shared the same cognitive capacity and behavioral capabilities. That said, evaluating cognitive abilities via hunting strategies is far from straightforward. That Figure 8. Schematic representation of overall trends in the data. MSA peoples were capable hunters is no longer questioned (e.g., Clark and Plug 2008; Faith 2008, 2011b; Milo 1998; Thompson and Henshilwood 2011). Although we did see an Linkages to Variation in Hunting Technology? increase in the exploitation of one type of dangerous game— A lack of agreed-on conventions for assigning lithic material suids—over time, we are not convinced by the argument that to specific MSA cultures/techno-complexes has precluded us a greater focus on dangerous game primarily signifies in- from exploring the relative impact of technological change on creased cognitive abilities (e.g., Klein and Cruz-Uribe 1996). the variance present in our data set in any detail. While new This observation could indicate intensification caused by in- classification systems for MSA lithics have been proposed creased population size or reflect improvements in hunting (Conard, Porraz, and Wadley 2012; Lombard et al. 2012), it technology that would reduce the danger associated with the remains to be seen whether researchers will apply these guide- procurement of these species. Unfortunately, it is not cur- lines. However, it is worth noting that a number of scholars rently possible to distinguish between these alternatives. have utilized data from the SB and HP in an attempt to Wadley (2010) argues that the use of remote capture tech- explore the potential relationships between lithic and faunal nologies indicates an essentially modern cognition. She pro- change, although the results of these studies are often at odds posed that the increased focus on taxa from closed environ- with each other (e.g., Dusseldorp 2012; Lombard and Clark ments and the diverse small game assemblage evidenced in 2008; McCall 2007; Newlander and Clark 2010). the HP at Sibudu could reflect use of these technologies. If In terms of well-published faunal assemblages subjected to one agrees with this hypothesis, then our data suggest that detailed taphonomic analyses, data from the SB are currently all MIS 4 populations in southern Africa—even those without limited to those from Blombos. The HP is also limited to a the HP—were utilizing remote capture technology and thus single sample—that from Sibudu. While other sites included possessed complex cognitive abilities. in this analysis do contain SB and HP deposits (i.e., the SB is also present at Sibudu and Diepkloof, while the HP has Summary: The Implication of Variability in Subsistence been identified at Klasies and Diepkloof), full taxonomic and/ during the MSA or taphonomic data are thus far lacking. Given this, we feel that more specific comparisons of subsistence between these Although we cannot fully evaluate the influence of techno- two phases should await the publication of a larger number logical and demographic change on the observed patterns, it of assemblages. seems clear that no single causal mechanism will adequately This being said, we would like to make one important account for the range of variation evidenced in the data. Of point. The increase in small ungulates (and small game more course, we are not only interested in understanding the driv- generally) that we identified in MIS 4 has often been rec- ing forces behind this variability but also in its broader evo- ognized as a phenomenon specific to the HP. For example, lutionary implications. At present, it would appear that sub- Backwell, d’Errico, and Wadley (2008) argued that the bone sistence behavior in the MSA was more variable than it was points and backed tools associated with the HP “may signify in the MP (Stiner 2013; but see Delagnes and Rendu 2011). part of a hunting adaptation to small prey in a closed forest This could be a reflection of a greater degree of behavioral environment” (1577). However, our data make it apparent flexibility among MSA populations. And yet, it is interesting that this pattern cannot be uniquely linked to the technology to note that we could not identify clear directional trends of the HP. The DK1 fauna is similarly dominated by small associated with increasing intensification or behavioral com- ungulates and small mammals—and despite the recovery of plexity. In fact, we only identified one example of a continuous S284 Current Anthropology Volume 54, Supplement 8, December 2013 directional trend, the case of the increasing frequency of suids may be related to environmental change, or it could reflect over time. The patterns in the faunal data are thus consistent intensification driven by population growth, although at the with those from other classes of material culture. During the current time we cannot effectively reconstruct population MSA, certain innovative technologies and artifacts with sym- sizes during the different stages of the MSA. Interestingly, this bolic significance also seem to come and go rather than show- spike in smaller fauna has been linked to the particular tech- ing an accretional trend toward increasing complexity, and nologies of the HP. However, given that our sample includes this is something the MSA record actually does seem to share data from DK1, a site occupied at the same time as the HP with the MP (cf. Hovers and Belfer-Cohen 2006, 2013; Lom- but lacking its characteristic technology (Thackeray 2000), we bard 2012; Wurz 2013). argue that technology is not the force driving this pattern. While preparing this paper, we identified several factors Conclusions that will be critical to the success of future studies of this sort. First, more detailed taphonomic analyses are needed. This Large-scale, specifically directed studies aimed at character- particularly applies to the smaller fauna, as the accumulating izing and explaining the nature and extent of variability in agents of these taxa are often unclear. New work by Armstrong human behavior within the MSA have generally been lacking (2013) on the DK1 fauna is a step in the right direction. despite the fact that a deeper understanding of this variability Second, data from more sites will be necessary to ascertain is clearly relevant to larger questions about human behavioral how robust the identified patterns are. For example, it will evolution and the rise and spread of AMH. There are a num- be interesting to see whether MIS 6 will consistently show ber of reasons for this, both practical and theoretical. How- the least evidence for the exploitation of low-ranked prey. ever, in light of the recent publication of faunal data from a Third, finer grained analyses will require better dating of ex- number of key MSA sites, we deemed it valuable to conduct isting collections. Finally, we think that a consideration of the a large-scale exploration of variation in MSA subsistence be- effect of technological change on human subsistence behavior havior incorporating material from eight sites that span ∼174– could prove very insightful in understanding variation in the 40 ka (MIS 6–3). faunal data; however, this type of analysis will not truly be Because we were interested in exploring evidence for var- possible until the development of a more standardized system iation in diet breadth over time, we largely approached the for defining archaeological cultures in the MSA. Conard, Por- data from the perspective of prey choice models under which raz, and Wadley (2012), Lombard et al. (2012), and Porraz smaller game is considered lower-ranked prey. Thus, an in- et al. (2013) have made significant steps toward this goal. crease in the exploitation of these taxa should indicate a Although we may not yet fully understand the causes driv- broadening of the diet. Notably, multiple strands of evidence ing the variability identified in the faunal record, we feel that suggest an increase in dietary breadth that peaked in MIS 4; we have made an excellent start in documenting the nature not only was there a marked increase in the smallest ungulates and extent of variation in subsistence behavior during the (Size 1), but also there was also an equally large rise in the MSA. In fact, the degree of variance in the data exceeded our exploitation of small mammals (!4.5 kg). Both of these types expectations. Our results indicate that subsistence behavior of prey declined noticeably in MIS 3. The diversity of shellfish during the MSA was more highly variable than that in the also peaked in MIS 4, which also indicates increasing dietary MP, although the broader evolutionary implications of this breadth. Interestingly, although many scholars emphasize the are not immediately apparent. importance of shellfish exploitation in the MSA, the available data suggest that even the largest MSA shell accumulations reflect a small contribution to the overall diet. Explaining the variance in the data is not necessarily a Acknowledgments straightforward task. Some dimensions of variability do seem to be driven primarily by environmental change; for example, J. L. Clark would like to thank Erella Hovers and Steve Kuhn the increased frequency in shellfish in MIS 5 (relative to MIS for their invitation to participate in the symposium on which 6 and 4) makes sense given that sea levels would have been this issue of Current Anthropology is based. Thanks to Richard highest during this period. Furthermore, MIS 5 and 3 show Klein for providing unpublished data from Klasies River Cave a similar (and high) frequency of grazers, particularly com- 1, James Brink for providing PDFs on Florisbad, Leslie Aiello pared with MIS 6 and 4, which show a predominance of and Laurie Obbink at Wenner-Gren for their advice and as- browsers. This suggests greater shrub and tree cover during sistance, and Teresa Steele and an anonymous reviewer for the glacial periods and more open environments in the their helpful comments. We also wish to thank Aaron Arm- warmer phases, which is consistent with our broader under- strong, Shaw Badenhorst, Genevieve Dewar, Tyler Faith, and standing of paleoenvironmental conditions in the region (e.g., Jarod Hutson for providing supplementary information. Re- Chase 2010; Ziegler et al. 2013). search by J. L. Clark was supported by an Alexander von What remains unclear is the reason behind the spike in the Humboldt Foundation postdoctoral research fellowship, the smallest ungulates and small mammals during MIS 4. This Leakey Foundation, and the Palaeontological Scientific Trust. Clark and Kandel Variation in Middle Stone Age Hunting Strategies S285 References Cited nological, elemental and colorimetric analysis of an engraved ochre frag- ment from the Middle Stone Age levels of Klasies River Cave 1, South Adler, Daniel S., and Guy Bar-Oz. 2009. Seasonal patterns of prey acquisition Africa. Journal of Archaeological Science 39:942–952. and inter-group competition during the Middle and Upper Paleolithic of de Ruiter, Darryl J., Juliet K. Brophy, Patrick J. Lewis, Steven E. Churchill, the southern Caucasus. In The evolution of hominin diets: integrating ap- and Lee R. Berger. 2008. Faunal assemblage composition and paleoenvi- proaches to the study of Palaeolithic subsistence. Jean-Jacques Hublin and ronment of Plovers Lake, a Middle Stone Age locality in Gauteng Province, Michael P. Richards, eds. Pp. 127–140. Vertebrate Paleobiology and Pa- South Africa. Journal of Human Evolution 55:1102–1117. leoanthropology Series. Dordrecht: Springer. Dusseldorp, Gerrit L. 2012. Tracking the influence of technological change Adler, Daniel. S., Guy Bar-Oz, Anna Belfer-Cohen, and Ofer Bar-Yosef. 2006. on Middle Stone Age hunting strategies in South Africa. Quaternary In- Ahead of the game: Middle and Upper Paleolithic hunting behaviors in the ternational 270:70–79. southern Caucasus. Current Anthropology 47:89–118. Faith, J. Tyler. 2008. Eland, buffalo, and wild pigs: were Middle Stone Age Armstrong, Aaron. 2013. Small mammal utilization by Middle Stone Age humans ineffective hunters? Journal of Human Evolution 55:24–36. humans at Die Kelders Cave 1, Western Cape, South Africa: Paleoanthro- ———. 2011a. Late Quaternary megafaunal extinctions in southern Africa’s pology Society meeting abstracts, Honolulu, HI, 2–3 April 2012. Paleoan- Cape Floral Region. PhD dissertation, George Washington University, Wash- thropology 2013:A2. ington, DC. Avery, Graham, David Halkett, Jayson Orton, Teresa Steele, Madelon Tusenius, ———. 2011b. Ungulate biogeography, statistical methods, and the profi- and Richard G. Klein. 2008. The Ysterfontein 1 Middle Stone Age rock ciency of Middle Stone Age hunters. Journal of Human Evolution 60:315– shelter and the evolution of coastal foraging. South African Archaeological 317. Society Goodwin Series 10:66–89. Feathers, James K. 2002. Luminescence dating in less than ideal conditions: Backwell, Lucinda, Franceso d’Errico, and Lyn Wadley. 2008. Middle Stone case studies from Klasies River Main Site and Duinefontein, South Africa. Age bone tools from the Howiesons Poort layers, Sibudu Cave, South Africa. Journal of Archaeological Science 29:177–194. Journal of Archaeological Science 35:1566–1580. Feathers, James K., and Daniel A. Bush. 2000. Luminescence dating of Middle Badenhorst, Shaw, and Ina Plug. 2012. The faunal remains from the Middle Stone Age deposits at Die Kelders. Journal of Human Evolution 38:91–119. Stone Age levels of Bushman Rock Shelter in South Africa. South African Gagnon, Mario, and Amy E. Chew. 2000. Dietary preferences in extant African Archaeological Bulletin 67:16–31. Bovidae. Journal of Mammalogy 81:490–511. Binford, Lewis R. 1978. Nunamiut ethnoarchaeology. New York: Academic Gaudzinski, Sabine. 2006. Monospecific or species-dominated faunal assem- Press. blages during the Middle Paleolithic in Europe. In Transitions before the Bird, Douglas W., and James F. O’Connell. 2006. Behavioral ecology and transition: evolution and stability in the Middle Paleolithic and Middle Stone archaeology. Journal of Archaeological Research 14:143–188. Age. Erella Hovers and Steven L. Kuhn, eds. Pp. 137–147. New York: Brain, C. K. 1974. Some suggested procedures in the analysis of bone accu- Springer. mulations from southern African Quaternary sites. Annals of the Transvaal Gaudzinski-Windheuser, Sabine, and Laura Niven. 2009. Hominin subsistence Museum 29:1–7. patterns during the Middle and Late Paleolithic in northwestern Europe. Brink, James S. 1987. The archaeozoology of Florisbad, Orange Free State.Mem- In The evolution of hominin diets: integrating approaches to the study of oirs van die Nasionale Museum (Bloemfontein) no. 24. Bloemfontein, South Palaeolithic subsistence. Jean-Jacques Hublin and Michael P. Richards, eds. Africa: National Museum. Pp. 99–111. Vertebrate Paleobiology and Paleoanthropology Series. Dor- ———. 1988. The taphonomy and palaeoecology of the Florisbad spring drecht: Springer. fauna. Paleoecology of Africa 19:169–179. Hawkes, Kristen, Kim Hill, and James F. O’Connell. 1982. Why hunters gather: Brink, James S., and Hilary J. Deacon. 1982. A study of a last interglacial shell optimal foraging and the Ache of eastern Paraguay. American Ethnologist midden and bone accumulation at Herolds Bay, Cape Province, South 9:379–398. Africa. In Palaeoecology of Africa: Southern African Society for Quaternary Henshilwood, Curtis S., Francesco d’Errico, Karen L. van Niekerk, Yvan Co- Research, Proceedings of the VIth Biennial Conference held at the Transvaal quinot, Zenobia Jacobs, Stein-Erik Lauritzen, Michel Menu, and Renata Museum, Pretoria, 26–29 May 1981, vol. 15. J. C. Vogel, E. A. Voigt, T. C. Garcı´a-Moreno. 2011. A 100,000-year-old ochre-processing workshop at Partridge, E. M. v. Z. Bakker, and J. A. K. Coetzee, eds. Pp. 31–39. Rot- Blombos Cave, South Africa. Science 334:219–222. terdam: Balkema. Henshilwood, Curtis S., Judith C. Sealy, Royden Yates, Kathryn Cruz-Uribe, Broughton, Jack M., and Donald K. Grayson. 1993. Diet breadth, adaptive Paul Goldberg, Frederick E. Grine, Richard G. Klein, Cedric Poggenpoel, change and the White Mountain fauna. Journal of Archaeological Science Karen Van Niekerk, and Ian Watts. 2001. Blombos Cave, Southern Cape, 20:331–336. South Africa: Preliminary report on the 1992–1999 excavations of the Mid- Buchanan, W. F. 1988. Shellfish in prehistoric diet, Elands Bay, S.W. Cape Coast, dle Stone Age levels. Journal of Archaeological Science 28:421–448. South Africa. British Archaeological Reports International Series 455. Ox- Hovers, Erella, and Anna Belfer-Cohen. 2006. “Now you see it, now you ford: BAR. don’t”: modern human behavior in the Middle Paleolithic. In Transitions Chase, Brian M. 2010. South African paleoenvironments during marine ox- before the transition: evolution and stability in the Middle Paleolithic and ygen isotope stage 4: a context for the Howiesons Poort and Still Bay Middle Stone Age. Erella Hovers and Steven L. Kuhn, eds. Pp. 295–304. industries. Journal of Archaeological Science 37:1359–1366. New York: Springer. Clark, Jamie L. 2009. Testing models on the emergence and nature of modern ———. 2013. On variability and complexity: lessons from the Levantine human behavior: Middle Stone Age fauna from Sibudu Cave (South Africa). Middle Paleolithic record. Current Anthropology 54(suppl. 8):S337–S357. PhD dissertation, University of Michigan, Ann Arbor. Hutson, Jarod M. 2012. A comparative study of life and death at Middle Stone ———. 2011. The evolution of human culture during the later Pleistocene: Age open-air sites within the southern African interior. PhD dissertation, using fauna to test models on the emergence and nature of “modern” University of Nevada, Reno. human behavior. Journal of Anthropological Archaeology 30:273–291. Hutson, Jarod M., and Chester R. Cain. 2008. Reanalysis and reinterpretation Clark, Jamie L., and Ina Plug. 2008. Animal exploitation strategies during the of the Kalkbank faunal accumulation, Limpopo Province, South Africa. South African Middle Stone Age: Howiesons Poort and post–Howiesons Journal of Taphonomy 6:399–428. Poort fauna from Sibudu Cave. Journal of Human Evolution 54:886–898. Jacobs, Zenobia, Richard G. Roberts, Rex F. Galbraith, Hilary J. Deacon, Rainer Clark, J. D. 1993. African and Asian perspectives on the origins of modern Grun, Alex Mackay, Peter Mitchell, Ralf Vogelsang, and Lyn Wadley. 2008a. humans. In The origin of modern humans and the impact of chronometric Ages for the Middle Stone Age of southern Africa: implications for human dating. M. J. Aitken, Curtis B. Stringer, and Paul A. Mellars, eds. Pp. 148– behavior and dispersal. Science 322:733–735. 178. Princeton, NJ: Princeton University Press. Jacobs, Zenobia, Ann G. Wintle, Geoffrey A. T. Duller, Richard G. Roberts, Conard, Nicholas J., Guillaume Porraz, and Lyn Wadley. 2012 What is in a and Lyn Wadley. 2008b. New ages for the post–Howiesons Poort, late and name? characterizing the “post-HP” at Sibudu. South African Archaeological final Middle Stone Age at Sibudu Cave, South Africa. Journal of Archaeo- Bulletin 67:180–199. logical Science 35:1790–1807. Delagnes, Anne, and William Rendu. 2011. Shifts in Neandertal mobility, Jerardino, Antonieta, and Curtis W. Marean. 2010. Shellfish gathering, marine technology and subsistence strategies in western France. Journal of Archae- paleoecology and modern human behavior: perspectives from cave PP13B, ological Science 38:1771–1783. Pinnacle Point, South Africa. Journal of Human Evolution 59:412–424. d’Errico, Franceso, Renata Garcia Moreno, and Riaan F. Rifkin. 2012. Tech- Kaplan, Hillard, and Kim Hill. 1992. The evolutionary ecology of food ac- S286 Current Anthropology Volume 54, Supplement 8, December 2013

quisition. In Evolutionary ecology and human behavior. E. A. Smith and B. Herries, Panagiotis Karkanas, Peter J. Nilssen, and Erin Thompson. 2010. Winterhalder, eds. Pp. 167–201. Hawthorne, NY: Aldine. The stratigraphy of the Middle Stone Age sediments at Pinnacle Point Cave Kelly, Robert L. 1995. The foraging spectrum: diversity in hunter-gatherer life- 13B (Mossel Bay, Western Cape Province, South Africa). Journal of Human ways. Washington, DC: Smithsonian Institution. Evolution 59:234–255. Kiberd, Philip. 2006. Bundu Farm: a report on archaeological and palaeoen- McBrearty, Sally, and Alison S. Brooks. 2000. The revolution that wasn’t: a vironmental assemblages from a pan site in Bushmanland, Northern Cape, new interpretation of the origin of modern human behavior. Journal of South Africa. South African Archaeological Bulletin 61:189–201. Human Evolution 39:453–563. Klein, Richard G. 1976. The mammalian fauna of the Klasies River Mouth McCall, Grant S. 2007. Behavioral ecological models of lithic technological sites, southern Cape Province, South Africa. South African Archaeological change during the later Middle Stone Age of South Africa. Journal of Ar- Bulletin 31:75–98. chaeological Science 34:1738–1751. ———. 1977. The mammalian fauna from the Middle and Later Stone Age Milo, Richard G. 1994. Human-animal interactions in southern African pre- (Later Pleistocene) levels of Border Cave, Natal Province, South Africa. history: a microscopic study of bone damage signatures. PhD dissertation, South African Archaeological Bulletin 32:14–27. University of Chicago. ———. 1979. Environmental and ecological implications of large mammals ———. 1998. Evidence for hominid predation at Klasies River Mouth, South from Upper Pleistocene and Holocene sites in southern Africa. Annals of Africa, and its implication for the behavior of early modern humans. Journal the South African Museum 81:223–283. of Archaeological Science 25:99–133. ———. 2001. Southern Africa and modern human origins. Journal of An- Munro, Natalie D., and Guy Bar-Oz. 2005. Gazelle bone fat processing in the thropological Research 57:1–16. Levantine Epipaleolithic. Journal of Archaeological Science 32:223–239. ———. 2009. The human career: human biological and cultural origins.3rd Newlander, Khori, and Jamie L. Clark. 2010. One size doesn’t fit all? modeling edition. Chicago: University of Chicago Press. technological change in the Middle Stone Age of southern Africa: Paleo- Klein, Richard G., and Kathryn Cruz-Uribe. 1996. Exploitation of large bovids anthropology Society meeting abstracts, St. Louis, MO, 13–14 April 2010. and seals at Middle and Later Stone Age sites in South Africa. Journal of Paleoanthropology 2010:A23. Human Evolution 31:315–334. Opperman, H. 1996. Strathalan Cave B, Northeastern Cape Province, South ———. 2000. Middle and Later Stone Age large mammal and tortoise remains Africa: evidence for human behaviour 29,000–26,000 years ago. Quaternary from Die Kelders, Cave 1, Western Cape Province, South Africa. Journal of International 33:45–53. Human Evolution 38:169–195. Plug, Ina. 1981. Some research results on the Late Pleistocene and Early Klein, Richard G., Kathryn Cruz-Uribe, and Peter B. Beaumont. 1991. En- Holocene deposits of Bushman Rock Shelter, Eastern Transvaal. South Af- vironmental, ecological, and paleoanthropological implications of the late rican Archaeological Bulletin 36:14–21. Pleistocene mammalian fauna from Equus Cave, Northern Cape Province, ———. 2004. Resource exploitation: animal use during the Middle Stone Age South Africa. Quaternary Research 36:94–119. at Sibudu Cave, KwaZulu-Natal. South African Journal of Science 100:151– Kuhn, Steven L., and Erella Hovers. 2006. General introduction. In Transitions 158. before the transition: evolution and stability in the Middle Paleolithic and Pollarolo, Luca, J. Wilkins, K. Kuman, and L. Galletti. 2010. Site formation Middle Stone Age. Erella Hovers and Steven L. Kuhn, eds. Pp. 1–11. New at Kudu Koppie: a late Earlier and Middle Stone Age site in northern York: Springer. Limpopo Province, South Africa. Quaternary International 216:151–161. Kuman, Kathleen, and R. J. Clarke. 1986. Florisbad: new investigations at a Porraz, Guillaume, Pierre-Jean Texier, Will Archer, Michel Piboule, Jean-Phi- Middle Stone Age hominid site in South Africa. Geoarchaeology 1:103–125. lippe Rigaud, and Chantal Tribolo. 2013. Technological successions in the Kuman, Kathleen, Moshe Inbar, and R. J. Clarke. 1999. Palaeoenvironments Middle Stone Age sequence of Diepkloof Rock Shelter, Western Cape, South and cultural sequence of the Florisbad Middle Stone Age hominid site, Africa. Journal of Archaeological Science 40:3376–3400. South Africa. Journal of Archaeological Science 26:1409–1425. Langejans, Geeske H. J., Karen L. van Niekerk, Gerrit L. Dusseldorp, and J. Rector, Amy L., and Kaye E. Reed. 2010. Middle and late Pleistocene faunas Francis Thackeray. 2012. Middle Stone Age shellfish exploitation: potential of Pinnacle Point and their paleoecological implications. Journal of Human indications for mass collecting and resource intensification at Blombos Cave Evolution 59:340–357. and Klasies River, South Africa. Quaternary International 270:80–94. Shea, John J. 2011. Homo sapiens is as Homo sapiens was: behavioral variability Lombard, Marlize. 2012. Thinking through the Middle Stone Age of sub- vs. “behavioral modernity” in Paleolithic archaeology. Current Anthropology Saharan Africa. Quaternary International 270:140–155. 52:1–35. Lombard, Marlize, and Jamie L. Clark. 2008. Variability and change in Middle Singer, Ronald, and John Wymer. 1982. The Middle Stone Age at Klasies River Stone Age hunting behavior: aspects from the lithic and faunal records. In Mouth in South Africa. Chicago: University of Chicago Press. Animals and people: archaeozoological papers in honour of Ina Plug. Shaw Skinner, John D., and Christian T. Chimimba. 2005. The mammals of the Badenhorst, Peter Mitchell, and Jonathan C. Driver, eds. Pp. 46–56. British southern African sub-region. 3rd edition. Singapore: Cambridge University Archaeological Reports International Series 1849. Oxford: Archaeopress. Press. Lombard, Marlize, Lyn Wadley, Janette Deacon, Sarah Wurz, Isabelle Parsons, Speth, John D., and Jamie L. Clark. 2006. Hunting and overhunting in the Moleboheng Mohapi, Joane Swart, and Peter Mitchell. 2012. South African Levantine late Middle Paleolithic. Before Farming 3:1–42. and Lesotho Stone Age sequence updated (I). South African Archaeological Steele, Teresa E., and Richard G. Klein. 2005/2006. Mollusk and tortoise size Bulletin 67:123–144. as proxies for Stone Age population density in South Africa: implications Louw, A. W. 1969. Bushman Rock Shelter, Ohrigstad, Eastern Transvaal: a for the evolution of human cultural capacity. Munibe 57:221–237. preliminary investigation, 1965. South African Archaeological Bulletin 24: ———. 2009. Late Pleistocene subsistence strategies and resource intensifi- 39–51. cation in Africa. In The evolution of hominin diets: integrating approaches to Lupo, Karen D. 2007. Evolutionary foraging models in zooarchaeological anal- the study of Palaeolithic subsistence. Jean-Jacques Hublin and Michael P. ysis: recent applications and future challenges. Journal of Archaeological Richards, eds. Pp. 113–126. Vertebrate Paleobiology and Paleoanthropology. Research 15:143–189. Dordrecht: Springer. Lupo, Karen D., and Dave N. Schmitt. 2005. Small prey hunting technology ———. 2013. The Middle and Later Stone Age faunal remains from Diepkloof and zooarchaeological measures of taxonomic diversity and abundance: Rock Shelter, Western Cape, South Africa. Journal of Archaeological Science ethnoarchaeological evidence from central African forest foragers. Journal 40:3453–3462. of Anthropological Archaeology 24:335–353. Stiner, Mary C. 2002. Carnivory, coevolution, and the geographic spread of Marean, Curis W., Yoshiko Abe, Carol J. Frey, and Robert C. Randall. 2000. the genus Homo. Journal of Archaeological Research 10:1–63. Zooarchaeological and taphonomic analysis of the Die Kelders Cave 1 layers ———. 2006. Middle Paleolithic subsistence economy in the Mediterranean 10 and 11 Middle Stone Age larger mammal fauna. Journal of Human region. In Transitions before the transition: evolution and stability in the Evolution 38:197–233. Middle Paleolithic and Middle Stone Age. Erella Hovers and Steven L. Kuhn, Marean, Curtis W., Miryam Bar-Matthews, Jocelyn Bernatchez, Erich Fisher, eds. Pp. 213–231. New York: Springer. Paul Goldberg, Andy I. R. Herries, Zenobia Jacobs, et al. 2007. Early human ———. 2013. An unshakable Middle Paleolithic? trends versus conservatism use of marine resources and pigment in South Africa during the Middle in the predatory niche and their social ramifications. Current Anthropology Pleistocene. Nature 449:905–908. 54(suppl. 8):S288–S304. Marean, Curtis W., Miryam Bar-Matthews, Erich Fisher, Paul Goldberg, Andy Stiner, Mary C., and Steven. L. Kuhn. 1992. Subsistence, technology, and Clark and Kandel Variation in Middle Stone Age Hunting Strategies S287

adaptive variation in Middle Paleolithic Italy. American Anthropologist 94: Tribolo, Chantal, Norbert Mercier, Eric Douville, Jean-Louis Joron, Jean-Louis 306–339. Reyss, Daniel Rufer, Nadia Cantin, et al. 2013. OSL and TL dating of the ———. 2006. Changes in the “connectedness” and resilience of Paleolithic Middle Stone Age sequence at Diepkloof Rock Shelter (South Africa): a societies in Mediterranean ecosystems. Human Ecology 34:693–712. clarification. Journal of Archaeological Science 40:3401–3411. Stiner, Mary C., and Natalie D. Munro. 2002. Approaches to prehistoric diet van Pletzen, Liezl. 2000. The large mammal fauna from Klasies River. MA breadth, demography, and prey ranking systems in time and space. Journal dissertation, University of Stellenbosch. of Archaeological Method and Theory 9:181–214. Vogelsang, Ralf, Ju¨rgen Richter, Zenobia Jacobs, Barbara Eichhorn, Veerle Stiner, Mary C., Natalie D. Munro, and Todd A. Surovell. 2000. The tortoise Linseele, and Richard G. Roberts. 2010. New excavations of Middle Stone and the hare: small-game use, the broad spectrum revolution, and Paleo- Age deposits at Apollo 11 rockshelter, Namibia: stratigraphy, archaeology, lithic demography. Current Anthropology 41:39–73. chronology and past environments. Journal of African Archaeology 8:185– Stiner, Mary C., Natalie D. Munro, Todd A. Surovell, Eitan Tchernov, and 218. Ofer Bar-Yosef. 1999. Paleolithic population growth pulses evidenced by Wadley, Lyn. 2010. Were snares and traps used in the Middle Stone Age and small animal exploitation. Science 283:190–194. does it matter? a review and case study from Sibudu, South Africa. Journal Thackeray, Anne I. 2000. Middle Stone Age artefacts from the 1993 and 1995 of Human Evolution 58:179–192. excavations of Die Kelders Cave 1, South Africa. Journal of Human Evolution Wadley, Lyn, Ina Plug, and Jamie L. Clark. 2008. The contribution of Sibudu’s 43:15–26. fauna to an understanding of KwaZulu-Natal environments at 60, 50 and Thackeray, J. Francis. 1979. An analysis of faunal remains from archaeological 37 ka. In Animals and people: archaeozoological papers in honour of Ina Plug. sites in southern southwest Africa (Namibia). South African Archaeological Shaw Badenhorst, Peter Mitchell, and Jonathan C. Driver, eds. Pp. 34–45. Bulletin 34:18–33. British Archaeological Reports International Series 1849. Oxford: Archaeo- Thompson, Jessica C. 2008. Zooarchaeological tests for modern human be- havior at Blombos Cave and Pinnacle Point Cave 13B, southwestern Cape, press. South Africa. PhD dissertation, Arizona State University, Tempe. Weaver, Timothy D., Teresa E. Steele, and Richard G. Klein. 2011. The abun- ———. 2010a. Taphonomic analysis of the Middle Stone Age faunal assem- dance of eland, buffalo, and wild pigs in Middle and Later Stone Age sites. blage from Pinnacle Point Cave 13B, Western Cape, South Africa. Journal Journal of Human Evolution 60:309–314. of Human Evolution 59:321–339. Wurz, Sarah. 2002. Variability in the Middle Stone Age lithic sequence, ———. 2010b. Variability in Middle Stone Age faunal exploitation and use 115,000–60,000 years ago at Klasies River, South Africa. Journal of Archae- of the physical and social landscapes in the southwestern Cape, South Africa. ological Science 29:1001–1015. In Settlement dynamics of the Middle Paleolithic and Middle Stone Age, vol. ———. 2013. Technological trends in the Middle Stone Age of South Africa 3. Nicholas J. Conard and Anne Delagnes, eds. Pp. 11–38. Tu¨bingen: Kerns. between MIS 7 and MIS 3. Current Anthropology 54(suppl. 8):S305–S319. Thompson, Jessica C., and Christopher S. Henshilwood. 2011. Taphonomic Ziegler, Martin, Margit H. Simon, Ian R. Hall, Stephen Barker, Chris Stringer, analysis of the Middle Stone Age larger mammal faunal assemblage from and Rainer Zahn. 2013. Development of Middle Stone Age innovation Blombos Cave, southern Cape, South Africa. Journal of Human Evolution linked to rapid climate change. Nature Communications 4: doi:10.1038 60:746–767. /ncomms2897. S288 Current Anthropology Volume 54, Supplement 8, December 2013

An Unshakable Middle Paleolithic? Trends versus Conservatism in the Predatory Niche and Their Social Ramifications

by Mary C. Stiner

The great temporal and geographic span of the Middle Paleolithic (MP) raises many questions about behavioral variation within this period and its evolutionary significance. This paper focuses on MP predator economics and its social ramifications by examining the data for possible trends in the size of the hominin ecological footprint, hunting practices, trophic level, food sharing, and the intensity with which sites were occupied. Middle Paleolithic hominins were big game hunters, and they were rather specialized in their focus on ungulate prey. Low-cost gatherable small prey were a perennial if minor contribution to MP diets at lower latitudes, but the overall breadth of the meat diet remained narrow throughout the period. Discernible trends in the MP are few. Foraging innovations of the MP include marine shellfish exploitation by 120,000 years BP (possibly earlier), galvanization of the prime-age ungulate hunting niche, and hearth-centered domestic camps. The density of zooarchaeological material seems to increase during the last 30,000 years of MP existence, implying mild increases in human populations. Important aspects of carcass processing and meat sharing in the MP do not show much variation but do indicate close cooperation and habitual sharing among group members. Contrasts to late Lower Paleolithic butchery patterns may illuminate more formal patterns of meat sharing in the MP and after. The seeming rigidity of MP hunting economics could have been the secret to its widespread success for 200,000 years.

Introduction and when very late MP populations first came in contact with African Middle Stone Age (MSA) or UP populations. Variation in Middle Paleolithic (MP) behavior and ecology This review of the MP emphasizes the archaeological record can be explored at many temporal and geographic scales, from of animal exploitation. While predation is but one way of local to the quasi-global. Temporal variation is seen in MP examining past hominin behavior, predation is one of the site densities, for example, and much geographic variation most powerful interactive forces in the evolution of life. There can be found in the prey species hunted. One may argue, can be no question that predation habits define a number of nonetheless, that the MP is more clearly defined by devel- important themes in the human evolution story not simply opments with the outset and close of this long culture period with respect to food energetics and ecology but also to the than by strong trends within it. Of course all arguments about bases of social cooperation and demography. I have two goals the MP come down to how much we choose to make of the in writing this essay. The first is to document changes in variation we find, and usually this is accomplished by com- hunting strategies and dietary breadth within the Mediter- parisons with recent human behavior. Though useful to a ranean MP and between the MP and periods just before and point, modern comparators are something of an Achilles’ heel after it. One cannot hope to cover the full volume of work in evolutionary interpretation because hindsight has nothing that has been done on MP economics in Eurasia. I will instead to do with the evolutionary processes that gave rise to or were focus mainly on the archaeological record of the Mediterra- important during the MP. If compared with the Lower Pa- nean Basin while bringing information from other areas to leolithic, the MP was a time of innovation. If compared with the discussion as much as possible. The Mediterranean region the Upper Paleolithic (UP), the MP was arguably a time of is a key testing ground for studies of variation in hominin stability and inflexibility. The hallowed standard of early subsistence because of the great variety of edible foods it offers “modern human behavior” is only particularly relevant where (Blondel and Aronson 1999) and the high density of archae- ological research. I will consider whether the variation ob- Mary C. Stiner is Professor in the School of Anthropology at the served in the faunal record is better explained by environ- University of Arizona (P.O.Box 210030, Tucson, Arizona 85721-0030, mental fluctuations or by social, demographic, or cognitive U.S.A. [[email protected]]). This paper was submitted 3 changes. The second goal of this essay is to reconcile the scope VII 13, accepted 24 VII 13, and electronically published 29 X 13. of the observed faunal trends with the temporal and geo-

᭧ 2013 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved. 0011-3204/2013/54S8-0010$10.00. DOI: 10.1086/673285 Stiner Trends versus Conservatism in the Predatory Niche S289 graphic breadth of the MP culture period as a whole. Before ecological variation because of the latitudinal gradient, diverse the invasion of anatomically modern humans (AMH) into topographies, and complex maritime-continental interfaces. their range, MP populations experienced selective conditions Undaunted by variable environments, MP hominins thrived that were somewhat distinct from those that confronted them in Eurasia for about 200,000 years. They were expert hunters when new, competing populations were present. The value of of large game animals wherever they lived, and they moved predatory specialization, for example, and the potential for large quantities of food and tool stone around to suit their population growth would almost certainly have changed in needs. The large game species that they hunted followed re- the face of interspecific competition. gional variation in animal community composition (cf. I will also explore the position that the differences between Grayson and Delpech 1998; Griggo 2004; Mu¨nzel and Conard the MP and the UP or the MSA need not have been about 2004; Patou-Mathis 2000; Speth and Tchernov 2001; Stark- differences in native cognitive ability. Cognitive differences ovich 2011; Stiner 1994, 2005; Valensi and Psathi 2004). In might well have existed between MP and MSA or UP pop- this regard, MP hunters appear to have been very flexible. ulations, but the evidence we have in hand does not really They were much less flexible in their use of small animals, point to this. Alternative explanations for the differing evo- in strong contrast to recent hunter-gatherers, UP foragers, lutionary trajectories of the MP, MSA, and UP may lie in the and possibly some MSA foragers. From a modernist per- demographic potential and stability of these Late Pleistocene spective, it seems odd that MP hominins did not expand their populations and, intimately linked to these population char- meat diet much at all in response to variation in biotic di- acteristics, the longevity and stability of knowledge-sharing versity over the more than 200,000 years of their existence in institutions (sensu Henrich 2004; Poteete and Ostrom 2004). Eurasia. Physiology does not predict cultural differences or intelligence One need not be a MP apologist, on the other hand, to among recent humans, and the same may have been true find that opinion currently is stacked against recognizing among Pleistocene MP and MSA populations. Lest we forget, “modernity” of any sort in the MP. There is no basis for MP artifacts, or artifacts very much like them, were made claiming that MP populations were inherently less imaginative and used by both Neanderthals and earliest AMHs in the in solving problems. They imagined reasons to bury deceased Levant (Bar-Yosef et al. 1986; Vandermeersch 1989). Consid- group members (e.g., Bar-Yosef et al. 1988, 1992; Haydal et erable planning capability is indicated for the MP from the al. 1995); they turned to the sea for new foods by at least many pathways by which flake or blade blank forms were 120,000 years ago (Stiner 1994); they collected uniquely at- produced (cf. Boe¨da 1994; Bourguignon, Faivre, and Turq tractive objects such as rock crystals and pretty shells (e.g., 2004; Delagnes and Meignen 2006; Kuhn 1995). In fact, MP Zilha˜o et al. 2010); they used marine sponges (Stiner 1994) lithic reduction systems tend to be a good deal more com- and mineral pigments (Rendu 2010; Roebroeks et al. 2012); plex—and the outcomes more elegant—than most MSA sys- they hafted stone artifacts to organic handles (e.g., Boe¨da et tems for working stone. Zooarchaeological data meanwhile al. 1996; Koller, Baumer, and Mania 2001; Shea 1989); they indicate that MP hunting practices relied on deep environ- made scraping tools from marine shells (Darlas 2007:360– mental knowledge, forethought, and close cooperation among 361; Stiner 1994:187–188; Vitagliano 1984); they used feathers group members. from large birds (Peresani et al. 2011); and they invented or Any attempt to review variation in MP subsistence is faced refined many distinct techniques for regulating blank form with patchy evidence. More problematic, however, is the fact in stone working. Important findings on the MSA from the that possible cultural continuity or gradients between Eurasia African continent, such as very early shell ornaments and bone and North Africa are underexplored. We should in principle tools (e.g., Bouzouggar et al. 2007; d’Errico et al. 2005; Hen- expect evidence of behavioral variation to increase as the area shilwood et al. 2004), have primed us to expect precocious of a paleoculture’s distribution increases merely because of behavioral patterns in this part of the world more generally. the likelihood of cultural drift. What we bump up against In the interest of healthy debate, why do seemingly innovative instead is an academic legacy of imposed geographic and acts in the MP hardly register in the story of “revolutionary” paleoculture boundaries that undermines attempts to look at human developments? real variation. If we assume that the MP simply did not exist The elephant in the room is the apparently capacious brain in Africa, even in areas nearest to the Mediterranean Basin, size of the Neanderthals. Scaled to body mass, AMH and then much of the potential variability is eclipsed by an ar- recent humans have nothing on MP Neanderthals in this bitrary segregation of archaeological records at the scale of regard (Klein 1999; Rightmire 2003). Argue as we may for continents. If, on the other hand, paleocultures such as the different wiring within the brain, it could all be special plead- Aterian of North Africa can be included in the MP, then the ing given the equally great metabolic burden of Eurasian MP, variation embodied by the MP paleoculture will seem far African MSA, and recent human brains (Snodgrass, Leonard, greater. and Robertson 2009). Surprisingly less attention has been Let us assume for the sake of this discussion that the MP given to the conditions behind the final burst in hominin was confined strictly to Europe and the whole of western Asia. brain expansion during the Middle Pleistocene, which led into This is still a huge area and one that encompasses tremendous the MP. Behaviors loosely associated with the tail end of this S290 Current Anthropology Volume 54, Supplement 8, December 2013 neurological trend included more extensive use of caves where tentially exists within the spectra both of plant use and animal available, ubiquitous fire technology, and residential camps use. Unfortunately, plant macro- and microfossil evidence that supported diverse activities. Thereafter we see remarkable cannot tell us much about the proportional contributions of consistency in how MP people reaped the benefits of large high- versus low-cost plants to early diets. Some archaeolo- prey through extensive meat transport to base camps, fire- gists turn to the technological record for answers, focusing centered food processing, formalized butchering, and meat particularly on durable milling, scraping, and pounding sharing. equipment to learn about investment and processing costs. From my perspective, the lists of differences thought to Plant-heavy diets among recent foragers tend to correlate with distinguish the MP and late MSA (McBrearty and Brooks emphasis on staple plant seeds and/or nuts (Keeley 1995), 2000) or the MP from the Eurasian UP (Mellars 1989, 1996) and heavy, durable tools are needed to extract the carbohy- are reducible to three phenomena: (1) diet breadth and as- drates from them efficiently. Apart from fire, the MP generally sociated radiations weapons technology, (2) durable art (i.e., lacks such tools. A distinct line of evidence from light isotopes beads and inscribed ochre and ostrich eggshells), and (3) the tends to support the idea of high levels of carnivory in MP degree of cultural volatility or “changeability.” In these re- hominins in Europe (e.g., Bocherens 2009), but data on MP gards, some phases of the MSA and the UP do seem to have populations at lower latitudes are still too few. more in common with each other than either has with the As for variable investments in the meat diet, the zooar- MP. chaeological record is the richest source of information on the range of animals eaten and the relative importance of each Behavioral Variation Within and Bracketing type provided that the prey species possess skeletons of some the MP Period kind. Faunal records present their own set of challenges to interpretation, but the inorganic components of bone and The most obvious component of MP subsistence is large game carbonate shells are well preserved in many regions of the exploitation—prey selection and capture, carcass processing, world. Equally important, the preserved remains can be re- land use, and provisioning of sites with meat. Other important lated in a consistent if generalized way to consumable organic themes in MP subsistence are small game exploitation and parts within and across prey species. foraging technology. Also important are the nature of sites Large prey in the size range of gazelles/roe deer to aurochs/ and their functions as evidenced from material contents, oc- bison were the core meat sources during the MP (cf. Delagnes cupation intensity, and the diversity of activities represented. and Rendu 2011; Gaudzinski-Windheuser and Niven 2009; Residential hub sites are very much in evidence in the MP, Hoffecker and Cleghorn 2000; Rabinovich and Tchernov and their development is relevant to the nature of coopera- 1995; Speth and Tchernov 2007). In north-central Europe, tion, delayed food returns, and domestic arenas of social in- MP hominins exploited large prey irrespective of global teraction. warming through the Eemian Interglacial (Gaudzinski 2004). A concerted focus on large prey was also typical in the bi- otically diverse Mediterranean Basin, where 95%–99% of the Living High in the Food Web animal foods procured by weight were large herbivores (fig. It is safe to say that MP people were dedicated large game 1; Stiner 2005). Within the “large prey” size window, MP hunters. While they must also have eaten plant foods, espe- hunters responded easily to variation in the species available cially in lower latitude environments, MP hominins occupied to them, hunting mainly red deer, ibex, or wild goat in some many cold regions in which plant productivity was highly areas; aurochs, bison, horse, or reindeer in others; and camel seasonal. In the absence of storage and milling tools for mak- or fallow deer and gazelle in yet others (cf. Adler and Bar- ing calorie-rich seeds or nuts digestible, meat was the only Oz 2009; Burke 2000; Conard, Bolus, and Mu¨nzel 2012; David high-quality food available to northern MP populations for and Farizy 1999; Gaudzinski and Roebroeks 2000; Griggo many months of the year. Important rare finds of seeds or 2004; Hoffecker 2009; Rendu et al. 2012; Reynaud Savioz and charred nut hulls in combustion features (Barton et al. 1999; Morel 2005; Rosell et al. 2012; Stiner 2009). Madella et al. 2002) and phytolith and starch grains preserved Rhino and straight-tusked elephant or mammoth remains in Neanderthal dental calculus (Henry, Brooks, and Piperno also occur at some MP sites, particularly in open settings. 2011) simply confirm what we all expect (Jones 2009), namely, Whether these megafaunal species were hunted or scavenged that MP hominins ate plant foods and sometimes used fire remains an open question, but their meat clearly was exploited to cook them. in some cases (Bocherens 2009; Conard and Prindiville 2000; What too many discussions of diet breadth fail to grasp, David and Poulain 1990; Griggo 2004; Hoffecker 2009; Patou- however, is the core importance of search and processing costs Mathis 2000; see also Villa 1990). Megafauna remains in cave of different foods in relation to their nutritional yield. This sites are uncommon if present at all probably because of the is a problem of how differing proportions of high- and low- high cost of transporting their heavy bones. Megafauna gen- cost food sources in the diet reflect variable human invest- erally drop out of MP records in accordance with the ex- ments (sensu Stephens and Krebs 1986). Such a contrast po- tinction of these species in local ecosystems (e.g., Garrard Stiner Trends versus Conservatism in the Predatory Niche S291

Figure 1. Percentage of total prey biomass represented by ungulate prey in three Mediterranean faunal series (adapted from Stiner and Kuhn 2006). Major climate cycles represented by the marine oxygen isotope record are adapted from Martinson et al. (1987).

1983; Hoffecker 2009; Hortola` and Martı´nez Navarro 2013; Ungulate skeletal representation typically varies from whole Stuart 1991:488–502), but aurochs-sized and smaller ungu- bodies to a bias favoring meaty or marrow-rich limb parts lates never lost their importance to MP hunters. and heads (e.g., Burke 2000; Conard and Prindiville 2000; A long-standing signature of human hunting is the ten- Griggo 2004; Patou-Mathis 1993, 2000; Speth and Tchernov dency to focus on prime-age artiodactyl prey (Stiner 1990). 2001; Stiner 2005; Valensi 2000). Figure 4 exemplifies the In reality, human-generated ungulate mortality patterns vary range of variation typical of medium-sized species from con- much more than this, but most patterns fall between non- secutive layers in U¨ c¸ag˘ızlı Cave II (MP) and U¨ c¸ag˘ızlı Cave I selective and strongly biased to prime adults, averaging to a (early UP) in southern Turkey. Here and elsewhere, MP for- mild bias to prime adult prey (figs. 2, 3; Stiner 2005). This agers often left axial parts behind at kill sites or exploited bias in prey age selection is widespread in the MP and UP them in the field. Differential preservation of dense versus (Gaudzinski and Roebroeks 2000; Hoffecker and Cleghorn spongy bones (following Lyman 1984) may explain the an- 2000; Patou-Mathis 2000; Rendu 2010; Speth and Tchernov atomical biases in some cases (e.g., Hoffecker, Baryshnikov, 1998; Steele 2004; Stiner 1990, 2005; Yesherun, Bar-Oz, and and Potapova 1991; Lam and Pearson 2005). However, the Weinstein-Evron 2007). The bias has also been documented patterns are clearly anthropogenic in a great many other cases in a few late Lower Paleolithic (LP) sites such as at Waller- (e.g., Burke 2000; Conard and Prindiville 2000; Rendu 2010; theim in Germany (Gaudzinski 1995) and Qesem Cave in Rendu et al. 2012; Speth and Tchernov 2001, 2007; Starkovich Israel (Stiner, Barkai, and Gopher 2011). The ubiquity of 2011; Stiner 1994, 2002, 2005) because there is no correlation prime-focused hunting in the MP is striking, however, and to bone density, or similarly dense skeletal features are quan- may suggest a galvanization of this dimension of the large tified across the axial and appendicular anatomy to infer game hunting niche. body-part profiles (Stiner 2002). The ungulate body parts that MP hominins carried from The MP is rife with examples of delayed consumption of kill sites to camps varied with circumstance, but repetitive high-quality animal parts. Carrying meat and marrow-rich biases are found in caves and rock shelters. Ideally we would bones to a central place allowed for more thorough process- like to have equally reliable data on prey body-part profiles ing, but the large volumes of meat moved also testify to an across open and natural shelter contexts, but preservation ethic of premeditated sharing. Evidence for delayed meat con- often favors samples from shelters. Shelter faunas nonetheless sumption is also found at late LP sites such as at Qesem Cave provide us with some analytical benefits because food must (Stiner, Barkai, and Gopher 2011) and probably the open site be carried to them. Most group members were likely present of Gesher Benot Ya’akov (Rabinovich, Gaudzinski-Windheu- at these sites, making sharing difficult to avoid. Evidence for ser, and Goren-Inbar 2008), but the MP differs in the sheer meat sharing in shelters can be explored by (1) considering abundance of clear-cut cases. In comparison with the stratified the quality and quantities of meat carried into them, (2) late LP record of Qesem Cave (Gopher et al. 2005), where attempting to follow the path of meat redistribution on site, butchering activities involved mainly simple defleshing and and (3) comparing the scope of activities at camps that may marrow extraction, MP meat sharing also appears more for- relate to modes of economic cooperation. malized (Stiner, Barkai, and Gopher 2011). The types of tool S292 Current Anthropology Volume 54, Supplement 8, December 2013

but at times turned to very different foraging agendas while using coastal caves in Latium, scavenging the very same kinds of ungulates they hunted elsewhere in addition to collecting shellfish and tortoises” (Stiner 1994:371). With the pendular return to the view of MP folk as big game hunters, scavenging behavior seems to have sunk beneath our notice (but see Conard and Prindiville 2000). This is unfortunate, because nonconfrontational scavenging is a form of gathering about which we still know little for the MP. At the site of Grotta dei Moscerini in west-central Italy, probable evidence for scav- enging head parts, nearly always from old animals, coincides with an unambiguous record of collecting shellfish and tor- toises (Stiner 1994).

Small Game Exploitation and Its Implications Large game hunting practices generally fail to differentiate the economies of MP and UP societies in Eurasia (see Adler and Bar-Oz 2009; Adler et al. 2006; Gaudzinski 2000, 2004; Grayson and Delpech 2003; Hoffecker 2009; Mu¨nzel and Con- ard 2004; Stiner 1994, 2005; Stiner, Barkai, and Gopher 2011). Figure 2. Models a and b averaged values for observed ungulate Economic differences are more apparent in how these paleo- mortality patterns created by various human and nonhuman cultures filled gaps in large game availability with small an- predators. Shaded panels represent natural variations in the age imals. In warm-temperate and subtropical environments, MP structures of living ungulate populations and thus also nonse- hominins favored small animals that were easy to collect— lective mortality patterns and mortality patterns caused by at- tortoises, marine shellfish, ostrich eggs, and large lizards if tritional factors such as disease, accidents, and malnutrition. Each corner of the diagram represents a strong bias toward the des- available. This makes good sense from the viewpoint of a prey ignated prey age group. Observed means are for recent spotted choice (optimality) model, as the low handling costs of slow- hyena (1), wolf (2), Cape hunting dog (3), tiger (4), African lion moving or immobile small animals compensate for their small (5), Holocene and recent human hunters (6), Mediterranean body size. Small, quick animals—particularly birds, hares, and Epipaleolithic and Upper Paleolithic hunters (7), and Mediter- fish—generally have higher handling costs, and MP people ranean Middle Paleolithic hunters (8). An asterisk indicates the did not pursue these prey most of the time (Aura et al. 2002; average for the Acheulo-Yabrudian (late Lower Paleolithic) fallow Costamagno and Laroulandie 2004; Laroulandie 2004; Stiner, deer assemblages from Qesem Cave. Munro, and Surovell 2000). Whatever flexibility may have existed in MP foraging systems, it seldom extended to animals marks on the ungulate bones from Qesem Cave are redun- or plants with high capture or processing costs. dant, and the cut-mark orientations are relatively chaotic. The Although habitual use of small, quick animals is rare in latter observation indicates many procedural interruptions or the MP overall, fascinating exceptions exist in southwestern diverse positions while cutting flesh and perhaps a more in- Europe, usually involving rabbits (Aura et al. 2002), a colonial dividualized way of consuming shared meat. MP and early burrower, and some birds. La Canelettes in France contains UP butchering practices are more similar to one another, with significant numbers of cut-marked rabbit and bird bones much consistency in cut-mark orientations, suggesting a more alongside large game remains (Cochard et al. 2012). Another complex, socially canalized way of sharing meat. exception is found in the early MP at Bolomor Cave in Va- No discussion of the MP would be complete without some lencia, Spain. Here cut-marked rabbit remains occur in rel- mention of the scavenging controversy. In the 1980s, Lewis atively high percentages in multiple horizons along with some R. Binford challenged a long-standing assumption that early birds and many tortoise and ungulate remains (Blasco and hominins (particularly LP hominins) were big game hunters Ferna´ndez Peris 2009). The fact that these examples are lo- (Binford 1981). He even suggested that MSA and MP hom- calized occurrences that repeat through multiple occupations inins might have been obligate scavengers of other predators’ suggests that unique aspects of the locality made this unusual kills (Binford 1984:244–249, 1988). The second hypothesis manner of foraging feasible. Some instances of MP bird ex- was generally refuted by subsequent analyses (e.g., Chase 1986, ploitation, again very rare, associate with ochre use and may 1989; Grayson and Delpech 1994; Speth and Tchernov 2007; have been for obtaining large feathers (Peresani et al. 2011; Stiner 1990, 1994). I, a student of Binford, concluded from Rendu 2010). Use of quick, small prey is not the rule in the long study of MP faunas in Italy that “Mousterian people MP, however, and the above examples stand out because of were competent hunters of small, medium and large ungulates their rarity (Cochard et al. 2012). Stiner Trends versus Conservatism in the Predatory Niche S293

dence on large game and on slow-moving and slow-growing small animals such as Mediterranean tortoises implies that MP populations were consistently small and highly dispersed. Given that the first detectable human demographic pulse oc- curred more or less at the threshold of the MP-UP cultural transition, some very late Mousterian populations may have been uniquely affected by expansions of early UP populations in Eurasia. The African MSA is not as extensively documented as the Eurasian MP. Especially troubling is the data gap for the in- terval 60,000–20,000 years ago (within marine isotope stage [MIS] 3) in both South and North Africa (Steele and Klein 2009) and spotty reporting in East Africa. There are hints of a greater range of subsistence and technological behaviors in some areas and phases of the MSA (reviewed by McBrearty and Brooks 2000; Steele and Klein 2009). Scarce bird remains are reported in some North African sites, small mammal re- mains (hare and hyrax) in the Ethiopian site of Porc-Epic (Assefa 2006), catfish bones in Aduma Middle Awash Valley sites (Yellen et al. 2005), and early bone harpoons (Yellen et al. 1995) and grinding tools that may have been suitable for processing seeds or nuts (McBrearty and Brooks 2000). These cases do not seem to constitute a single trend but rather oscillate substantially with environmental changes (especially in MIS 4; Clark and Kandel 2013). The MSA data nonetheless suggest a contrast to the MP, even to the MP in warmer climes. Better documented jumps in diet breadth occur in African Late Stone Age sequences (Klein 1999; Steele and Klein 2009) Figure 3. Observed mortality pattern distribution in artiodactyl and post–Large Glacial Maximum sequences in Eurasia prey from Mediterranean Epipaleolithic (EPI), Upper Paleolithic (UP), and Middle Paleolithic (MP) assemblages from Israel (Hay- (Munro 2004; Stiner 2001; Stiner et al. 1999). onim Cave, Meged Rockshelter, Kebara Cave), Lebanon (Ksar Evidence of early shellfishing has attracted new attention. ’Akil), Turkey (U¨ c¸ag˘ızlı I Cave), and various cave sites in west- It is said by some to be evidence of precocious expansions central and northern Italy. Acheulo-Yabrudian (late Lower Pa- in dietary breadth and cognitive advances during the MSA in leolithic) cases from Qesem Cave appear as diamonds. South Africa (Marean et al. 2007). The MP has not figured much in this discussion, but it should (see Colonese et al. A major shift in the overall breadth of the meat diet in the 2011). Evidence for probable MP shellfishing in the Medi- Mediterranean region coincides with the MP-UP transition terranean region—mainly limpets, large clams, mussels, oys- (fig. 5; Stiner 2001; Stiner et al. 1999). Differences in small ters, and turbans—has been widely reported in archaeological game use between the two Paleolithic periods are made all works of the 1950s onward at Gibraltar (Baden-Powell 1964; the more fascinating by the fact that diet diversification and Garrod et al. 1928); on the Italian coasts of Liguria, Lazio, intensified use of resources can support a larger population and Puglia (Blanc 1958–1961; Palma di Cesnola 1965, 1969); base. In the Levant, this transition in small game use is linked and at Haua Fteah in Cyrenaica, North Africa (Klein and to harvesting pressure on sensitive tortoise populations. The Scott 1986). Detailed taphonomic studies are available for mean sizes of tortoises declined (Speth and Tchernov 2002; only some of these sites, but recent results confirm that hom- Stiner et al. 1999), and unnatural skewing is evidenced in the inins exploited shellfish at Cueva Perneras and Cueva de los age (size) structure of the harvested animals, which points to Aviones in Spain (Zilha˜o et al. 2010), Kalamakia Cave in heavy exploitation by humans in particular (Stiner 2005:139– Greece (Darlas 2007), Grotta dei Moscerini in Italy (Stiner 147). These findings imply that human populations in the 1994), U¨ c¸ag˘ızlı Cave II in southern Turkey (Stiner 2009), and region first exceeded the potential of high-ranked, high-return in Morocco (Steele and Alverez-Ferna´ndez 2011). Several of resources to support them after about 50,000 years ago (see the MP sites on Gibraltar contain shellfish remains, and al- Speth and Clark 2006 for effects within the late MP). De- though some are almost certainly archaeological, their attri- clining availability of high-yield resources, which raises search bution to hominin activities continues to be controversial (cf. costs, is considered the main reason why foragers turn to Barton 2000; Erlandson and Moss 2001; Fernandez-Jalvo and foods that give a lower return for the effort (Stephens and Andrews 2000; Finlayson et al. 2008; Freeman 1981; Klein Krebs 1986:17–24). Middle Paleolithic foragers’ great depen- and Steele 2008; Stringer et al. 2008). The earliest secure case S294 Current Anthropology Volume 54, Supplement 8, December 2013

Figure 4. Comparison of standardized skeletal element frequencies (observed MNE/expected MNE) by layer and anatomical region for medium-sized ungulate prey at the Epipaleolithic and early Upper Paleolithic site of UcI (layers EPI though I, youngest to oldest) and the Middle Paleolithic site of UcII (layers A through C-D in bottom row, arranged youngest to oldest) in southern Turkey. Unevenness in the anatomical profile indicates biases in body-part representation relative to the complete animal anatomy. Dental elements were not used for the calculations of head-part abundance. for MP shellfish exploitation comes from Grotta dei Mos- have bumped into Mediterranean Neanderthals already in the cerini, a deeply stratified cave site on the Gaetan coast of Italy habit of gathering mussels, limpets, and turbans from shore- that spans 115–65 ka (Stiner 1994:180–192). line rocks and digging for clams buried in the sand. The idea The earliest record of MSA shellfish exploitation at Pinnacle that the coast was a main corridor for colonization is doubtful Point (PP13B) in South Africa has been dated to about 160 in any case given that good shellfishing areas were virtually ka (Marean et al. 2007). Taken at face value, shellfishing on absent along the coast from Egypt to northern Galilee. Im- the South African coast began roughly 40,000–50,000 years portantly, Clark and Kandel (2013) show that the quantities before it appeared on Mediterranean shores. However, geo- of shells in coastal MSA are low as a rule and on par with logical studies of shoreline changes indicate that much of the those in the MP sites. Mediterranean archaeological record is now submerged or has been lost to the erosion with marine transgression following Labor Allocation and Foraging Equipment the Last Interglacial (Lambeck 1996; Van Andel and Tzedakis 1996). Given that all of the known shell-bearing MP sites This is not a paper on Paleolithic technology, but a few re- occur at or just a few meters above modern sea level, one marks about foraging implements are relevant to any consid- can assume that earlier coastal occupations have been scoured eration of hominin subsistence. First, it is clear that late LP clean by waves or inundated by rising seas (reviewed in Bailey and MP hominins routinely hunted large game animals long and Flemming 2008; Colonese et al. 2011). For the moment, before the development of stone-tipped and bone-tipped 115–110 ka is merely the minimum age for shellfish collecting weapons. In Eurasia, stone-tipped weapons appear in the later in the Mediterranean region. Suggesting significantly different MP as localized traditions. Hafted Levallois points are re- behavioral thresholds between the MP and MSA for marine ported in the Levant (Shea 1989, 1993), for example, and foraging therefore is premature. bifacial Blattzspitzen in Germany (Mu¨ller-Beck 1988) along Marean et al. (2007) argue that early shellfishing in South with rare pointed bone artifacts (Conard, Bolus, and Mu¨nzel Africa fueled a demographic expansion of MSA populations 2012). Modest weapons diversification therefore is suggested along coastlines northward into Eurasia. The energetic basis by the late MP. Bifacial stone and bone points are also reported of this proposition is highly questionable, but even if such a in the African MSA (McBrearty and Brooks 2000; Shea 2009). thing occurred, the expanding MSA populations would soon Because stone-tipped weapons are not universal to the MP Stiner Trends versus Conservatism in the Predatory Niche S295

eventually came to support. We have seen that MP people could obtain quick animals if they wanted to but seldom chose to do so. Early radiations in small game hunting equipment are reported in some MSA records of Africa (e.g., fishing; McBrearty and Brooks 2000), apparently in connection with expanding dietary breadth. Expanding dietary breadth has labor and social connota- tions. The benefits of niche (labor) separation within human groups should increase as the diet broadens over long periods of time because of less overlap or symmetry in personal sched- ules and the locations where various foods can be obtained. Such within-population diversification would be most ad- vantageous in environments where key resources occur at disparate locations or times or if distinct methods and tools Figure 5. Changes in evenness in the exploitation of three small are required to obtain them efficiently. These conditions may prey categories based on the inverse of Simpson’s index (1 p also result in greater autonomy with respect to what individ- p least even and the narrowest diet; 3 most even and therefore uals add to a pool of shareable foods. the broadest meat diet). Prey are low-cost slow game and higher- To appreciate the social ramifications of expanding diet cost quick-running terrestrial mammals and quick-flying birds; fish are very rare and therefore are excluded. Black-filled circles breadth across the MP-UP, one must consider diversification represent the U¨ c¸ag˘ızlı II-I faunal series; unfilled and gray-filled in technology and diet simultaneously. Upper Paleolithic pop- circles denote other Mediterranean series in Italy and Israel (from ulations readily diversified along either dimension or both in Stiner 2001). Sources: late MP from Kebara Cave ca. 65–50 ka response to environmental circumstance. They produced, for (Speth and Tchernov 2001); Paleolithic Italy, Israel, and Turkey example, weatherproof clothing and related items to support (Munro 2004; Stiner 1994, 2001, 2005, 2009; Stiner, Barkai and the food quest in cold areas and diverse food harvesting equip- Gopher 2011); Greece, Klissoura Cave 1 (Starkovich 2011); ment in warmer areas. Acknowledging that we know little Franchthi Cave (Stiner and Munro 2011). (Adapted from Stiner 2001, 2005.) about organic artifacts of any Paleolithic period, the MP none- theless shows less flexibility in both technology and the scope of the meat diet. There is considerable evidence for hide work- or the MSA, however, wooden spears were almost certainly ing in the MP (Meignen et al. 1989; Villa, Bon, and Castel the default weapon of choice. In Eurasia, this represents a 2001), suggesting that European Neanderthals wore skin gar- continuation of late LP hunting equipment (cf. Jacob-Friesen ments. However, evidence of working dry hide (cured leather) 1956; Thieme 1997). in the form of microwear traces on stone artifacts is com- The evidence forces us to decouple suppositions about Pa- paratively scarce in the MP (e.g., Anderson-Gerfaud 1990; leolithic hunting capabilities and complex hunting weapons. Beyries 1987; Lemorini 2000; Martı´nez-Molina 2005). It is Improvements in weapons design are typical of the UP (e.g., only with the UP that the types of artifacts commonly as- Knecht 1997), and increased weapons efficiency or reliability sociated ethnographically with tailored, weather-resistant may have reduced an individual’s procurement time, risk per clothing—bone needles and awls—also became a regular part foray, and the minimum hunting party size needed to capture of the Eurasian archaeological record. Upper Paleolithic so- large animals. Middle Paleolithic hunters nonetheless man- ciety seems to have differed from MP society because of a aged very well without complex weapons, relying instead on wider range of economic and social roles. extensive cooperation among group members. The impor- Kuhn and Stiner (2006) hypothesize that MP women, chil- tance of cooperation in MP big game hunting is underscored dren, and men all participated more consistently and directly by the apparently low population densities of the period. Close in large game hunting than is generally seen among recent teamwork would have been utterly essential to MP hunting foragers or UP foragers. This is not to say that socioeconomic success and maintaining minimum party size a perennial con- roles among MP individuals by age or gender were identical. cern. Rather, the point is that a comparatively narrow economic While there is no necessary link between the ability to bring focus on large game and very low population densities con- down large game animals and weapons complexity, there is strained labor organization and land use in historically unique a predictable relation between the exploitation of small, quick ways. Like other predators that hunt socially, humans can animals and technologies that reduce capture costs in Pa- gain significant advantages over large prey if some members leolithic Eurasia. This is because the quick prey are small and of the hunting party act as artificial surrounds or funnels that the returns are too low relative to human metabolic needs in move the quarry toward the killers. (Only substantive in- the absence of tools that make harvesting efficient (sensu creases in weapon efficiency can alter this dynamic.) Individ- Oswalt 1976). Making and maintaining the harvesting tools ual roles in MP hunting would have varied from direct phys- also incurs costs, which some later Paleolithic infrastructures ical contact with prey to diverse, safer tasks for helpers. In S296 Current Anthropology Volume 54, Supplement 8, December 2013 such small-scale societies that lacked sophisticated killing density of material in MP sites must relate to the season of tools, group members had to be available on short notice occupation (e.g., Boyle 2000; Hoffecker and Cleghorn 2000; much of the time. Beating the bushes, carrying meat and Patou-Mathis 2000; Pike-Tay et al. 1999), and it seems likely bones back to camps, processing carcasses, and tanning hides that cold-weather camps may have lasted longer (e.g., Conard, were all essential to gaining high payoffs from a hunt. More Bolus, and Mu¨nzel 2012). However, the generally opposing helpers meant that more meat and bone could be carried back relation between hominin and carnivore presence signals an to camps, cutting losses to scavengers and allowing more to important divide in the function of sites within MP territories. be gained from the meat, marrow, and hides of large prey This phenomenon continues through the early UP in some animals. Given the perennial risk of groups being stretched regions, suggesting that low population densities could be part too thin and the apparently limited development of regional of the explanation. social networks, within-group proximity would also have been The MP site record is not entirely trendless. There is an the main safety net for MP people. increase in the number of sites after about 70 ka (e.g., Mirazo´n Lahr and Foley 2003; Van Andel et al. 2003). The ravages of time complicate interpretation of this trend but cannot ex- Site Function and Occupation Intensity plain it away. Moreover, a higher proportion of the later MP Two modes have been noted in MP site data: (1) ephemeral sites were occupied more intensively. For example, human occupations with high mobility and (2) relatively intensive occupation intensity in the long consecutive sequences of occupations with high material inputs. The two conditions Hayonim (early MP) and Kebara Caves (late MP) has been do not seem to represent a single continuum. The relative compared through integrated analyses of faunal and lithic frequencies of ungulate and lithic materials are positively and assemblages, hearth features, and wood ash deposits (Meig- strongly correlated in the richer cave sites (Moncel and Dau- nen, Speth, and Stiner 2006; Meignen et al. 2010). The MP jeard 2012; Riel-Salvatore and Barton 2004; Stiner and Kuhn occupations in Hayonim Cave are repetitive in content but 1992), implying that tool manufacture and maintenance were small in scale. Thermoluminescence dates suggest a sediment intimately connected to large game exploitation. The relation accumulation rate of about 1 m per 10,000–15,000 years. By is weaker or nonexistent in sites with scant materials and high contrast, each meter accumulated in roughly 3,000 years in rates of tool retouch and exotic raw materials (e.g., Grotta Kebara Cave (Goldberg and Bar-Yosef 1998), and the densities Guattari, Grotta dei Moscerini [Kuhn 1995], and Yarımburgaz of lithic artifacts are also much higher. Cave in Thrace [Kuhn and Stiner 2010]). Delagnes and Rendu While the intensity of human presence at Hayonim was (2011) propose that the duration or length of lithic reduction generally low, all stages of stone tool production are in evi- sequences may also have varied with mobility and hunting dence (Meignen et al. 2010), and raw material came mainly strategies. The sparse hominin occupations in caves tend to from local sources (Delage, Meignen, and Bar-Yosef 2000). intercalate or be intermixed with carnivore occupations (Bru- Ungulates and tortoises were exploited repeatedly, and evi- gal and Jaubert 1991; Gamble 1999; Patou-Mathis 2000; Speth dence of fire-aided food preparation is widespread (Stiner and Tchernov 1998; Stiner 1994; Stiner, Arsebu¨k, and Howell 2005; Weiner, Goldberg, and Bar-Yosef 2002). Also common 1996; Straus 1982; Villa and Soressi 2000). Gamble (1986) is evidence of postdepositional burning of bone and lithic was among the first to examine the distribution of this phe- material, indicating considerable reuse of certain locations nomenon across the Mediterranean area. Where hominid within the cave (Stiner et al. 2001). The variety of economic components are thin, carnivore components often are thick activities represented tells us that Hayonim cave served mainly and easily recognized (e.g., Aura et al. 2002; Brugal and Jau- as a residential camp. Complete on-site core reduction and bert 1991). a diversified tool kit rule out the possibility of a special use Other MP sites instead are dominated by hominid-gen- site (Meignen et al. 2010). The pace of material buildup (den- erated debris. Early MP examples in the Levant include Hay- sity) was significantly higher overall during the late MP in onim Cave (Stiner 2005) and Misliya Cave (Yesherun, Bar- Kebara Cave (Albert et al. 2000, 2003; Goldberg and Bar- Oz, and Weinstein-Evron 2007). The quantities of faunal Yosef 1998; Meignen et al. 2010; Weiner, Goldberg, and Bar- material in these sites are not tremendous relative to sediment Yosef 2002). A concentrated bone midden occurs along the volume, but all of the high-quality ungulate body parts that north wall in Units XI-IX (Speth and Tchernov 1998), and we would expect to be valued by hunters are represented along carcass processing and consumption occurred at spatially dis- with hearth traces and evidence of butchering and marrow crete locations inside the cave (Speth and Clark 2006; Speth processing. Later MP examples are both more common and and Tchernov 2007). Hearth features are prevalent and deeply often richer in material, such as in Kebara Cave (Bar-Yosef stacked with little or any sediment formation between lenses et al. 1992; Speth and Tchernov 1998) and Qafzeh Cave (Ra- (Bar-Yosef et al. 1992; Meignen, Bar-Yosef, and Goldberg binovich and Tchernov 1995). Comparable situations are re- 1989). A full suite of lithic production is represented in Kebara ported throughout Europe (e.g., Chase 1986; David and Pou- (Meignen et al. 2010), generally on local flint, and the tools lain 1990; Farizy, David, and Jaubert 1994; Gaudzinski 1995; were used for diverse tasks, including butchery, woodworking, Jaubert et al. 1990; Stiner 1994). Some of the variation in the cutting, and scraping of hard and medium materials (Plisson Stiner Trends versus Conservatism in the Predatory Niche S297 and Beyries 1998; Shea 1991). The absence of exotic raw tial (Goldberg et al. 2012) is not fully resolved, but probably material and the complete reduction sequence of cores in the it is due to the latter. site indicate large, well-stocked encampments of prolonged duration (Meignen et al. 2010). Some MP layers in Kebara Evidence Summary Cave display considerable heterogeneity in site structure, ap- parently in response to rapid debris buildup. The relegation It is a good deal easier to find subsistence trends between the of bone trash along the peripheries of habitation areas is MP and the periods before and after it than within the MP expected to increase with the duration of an occupation (Bin- itself. The transition from the late LP to early MP seems ford 1978, 1991, 1998; Brooks 1984; Galanidou 2000; Gorecki gradual with respect to dependence on large game, basic hunt- 1991). This condition is not seen in the early MP of Hayonim ing equipment, aspects of blade technology, and prey age Cave even though this site and Kebara served mainly as res- selection. One possible contrast to the late LP is a more com- idential encampments. There evidently was more impetus to plex pattern of meat butchering and sharing based on cut- manage domestic space in the later, denser site. mark data, but more work is needed on this issue. Another Residential camps are places where all group members may contrast to the LP is the consistent and rather more complex congregate and engage in a wide range of activities. By this “residential” nature of many MP sites, including the ubiquity definition, residential camps are clearly in evidence through- of fire. The emergence of burial customs and pigment use out the MP and UP. If we try to count the number of distinct also set the MP apart from the late LP. These practices almost activities in dense sites, however, there seem to be fewer rep- certainly relate to important social developments in the MP resented in MP camps than in UP camps (and fewer still in even if we do not know just what they were about. The shifts the late LP camps). Trends in the complexity of site activity between the MP and early UP tend to be more abrupt, al- histories may be undermined by differential preservation but though not with respect to big game hunting or the dense- not where durable classes of technological material are in- ephemeral site contrast within territories. However, the UP volved. displays novel solutions for managing foraging risk through There is at least one other aspect of site use—the centrality clear expansions in dietary breadth, the use of artistic media, of fire—that may define the MP period as a whole. Fire tech- and rapid technological diversification. A number of archae- nology emerged well before the MP (Alperson-Afil and ologists argue that these qualities of the UP contributed to Goren-Inbar 2010; Gowlett et al. 2005; Karkanas et al. 2007; greater population stability and environmental carrying ca- Preece et al. 2006), but the record of fire becomes ubiquitous pacity. with the beginning of the MP (e.g., Albert, Berna, and Gold- These observations have led to an impression of MP pop- berg 2012; Ferna´ndez Peris et al. 2012; Goldberg et al. 2012; ulations as proverbially stuck in the mud, left behind by evo- Roebroeks and Villa 2011; Rolland 2004; Vallverdu´etal. lution. Sampling and preservation issues notwithstanding, the 2012). Hearths were magnets for carcass processing activities late MP may hint at a few subtle changes within the period. in MP residential sites (Rosell et al. 2012; Speth 2006; Stiner Hominin populations may have grown, based on site structure 2005; Vallverdu´ et al. 2012; Vaquero and Pasto´ 2001), and evidence and numbers of sites, although there are few if any they probably were intensely social spaces as well (Foley and indications of increased predator pressure on large or small Gamble 2009; Gamble 1999). Evidence of fire is not preserved animals resources (but see Speth and Tchernov 2007 on Lev- in every MP site, but it is as common in MP sites as it is in antine cases). Dated burials also cluster in the later part of UP sites if time-dependent preservation is taken into account the MP. And the late introduction of stone-tipped hunting (Roebroeks and Villa 2011). Sandgathe et al. (2011) take a weapons suggests mild increases in hunting efficiency in some different view, arguing that fire only became a regular part regions. MP meat diets nonetheless remained narrow nearly of life in the late MP, and that earlier use of fire was limited everywhere. to curation of live coals obtained from natural fires. It is Part of the explanation for greater diet breadth in the Af- difficult to reconcile Sandgathe et al.’s interpretation against rican MSA could be environmental, because so much of the the rich, long-term fire records in the late LP site of Qesem MP range falls in higher latitude environments, where the Cave based on the distribution of burned bones (Stiner, choice of animal and plant foods would have been less varied. Barkai, and Gopher 2011), the early MP of Hayonim Cave The same cannot be argued for the contrast between the (Weiner, Goldberg, and Bar-Yosef 2002) and Bolomor Cave Mediterranean MP (including the late MP) and the early UP, (Ferna´ndez Peris et al. 2012), or geologist A. Segre’s obser- as both existed sequentially in the same parts of the world vations from the excavation of Grotta dei Moscerini (Stiner and each weathered many shifts in climate, environment, and 1994:46–52). Repeated use of fire in these early cultural series food supplies. Dietary and technological evidence show us spans distinct climate cycles and environmental regimes. It is that both MP and UP societies depended on systems of di- interesting that MP fire records are much clearer in some vided, collaborative labor, but MP activities indicate a nar- regions than in others and not necessarily in the coldest en- rower spectrum of on-site activities and probably greater uni- vironments. Whether this anomaly reflects regional behavioral formity in ranging patterns of group members. differences or taphonomic differences in preservation poten- The most readily visible differences between the early and S298 Current Anthropology Volume 54, Supplement 8, December 2013 late MP seem to be about numbers of people—rates of site of specialization along with a narrower tolerance range in one visitation, population density at the regional scale, and pos- or more dimensions of niche (Pianka 1978:253–256). Spe- sibly social group sizes. The potential for population growth cialization brings greater efficiency in the exploitation of cer- in the MP would have been severely constrained by frequent tain core resources that in turn may reduce the group’s ability residential moves and an economy based on a high-quality to be efficient in exploiting others. Because of this, it is dif- but unpredictable nutrient supply (Boone 2002; Mu¨nzel and ficult for specialists to retreat from time-hardened success. Conard 2004; Stiner, Munro, and Surovell 2000). Of course, Middle Paleolithic populations went far down the path of MP females maintained viable levels of reproductive success large game specialization, a path made possible in part by through thick and thin. How did they pull it off, if the social high ungulate biomass during the ice ages (e.g., Discamps safety nets of the MP did not extend very far beyond the 2011). boundaries of small groups? In the absence of reliable regional The low population densities of the MP raise questions networks, group coherence, vigilant concern for group mem- about the importance of cultural drift, because isolation can bers, and close cooperation would have been utterly critical foster regional variation and distinct trajectories of change to MP survival. One would also expect males to remain in over the long term. Of course high mobility and the tele- natal territories through adulthood if large game hunting were graphic scent of fire meant that MP folk were not totally cut the core of the economy. Recent findings by Lalueza-Fox et off from one another. Considerable temporal and spatial var- al. (2011) on the late MP individuals from El Sidron Cave iation nonetheless exists in the details of MP stone working. suggest that may indeed have been the case. Rather than periodic, directional diffusion of novel ways of doing things, we see rather stochastic patterns or reappearance Why Is There Not More Variation of a set array of technological variants in different areas (e.g., within the MP? Delagnes and Meignen 2006; Kuhn 2013; Moncel and Dau- jeard 2012; Peresani 2012; Vaquero et al. 2012). This pattern One of the great challenges to studying the MP is explaining may be a consequence of demographic factors in that low- the appearance of stasis in spite of the considerable intelli- density populations are subject to comparatively high rates gence of these hominins. Middle Paleolithic humans existed of local extinction and recolonization. These conditions on narrow meat diets across a wide range of latitudes. Why would reduce the probability of cumulative cultural evolution was there not more variation in small game use in the bi- via complex learning traditions while promoting some degree otically diverse Mediterranean Basin? The prey animals were of nondirectional variation in material culture (Premo and there. The typically low population densities of the MP are Kuhn 2010; Stiner and Kuhn 2006). The MP has the ap- part of the answer. A heavy dependence on large game implies pearance of a stable adaptation with plenty of flexibility built a very high position in the food web, and it seems that con- into it. But it was headed nowhere in particular—no evidence sumer abundance seldom exceeded the supply of high-quality of strong directional selection—consistent with the status of foods at the regional scale. While large game animals can yield a successful and well-tuned system. The MP was not a dead high average return rates, they associate with high variances end in itself but rather the victim of the historically unique in supply (reviewed in Kelly 1995; Kuhn and Stiner 2006) colonization by a competing population. and thus limit the reproductive potential of any highly car- nivorous population. These populations compensate with A Zooarchaeological Agenda high mobility, which also facilitates recolonization where local extinctions occurred. The evidence synthesized in this essay exposes some clear-cut It is striking that even the late MP cultural record changed opportunities for further research. Specifically, how different only a little, because this is generally when AMH populations were MSA and MP population responses to food supplies if, expanded into Eurasia. The conservative nature of MP culture for example, we compare them during climate intervals of may have been the outcome of a very successful and long- more stability and less stability at a millennial scale (see also standing adaptation, one that became fixed via the process of Clark and Kandel 2013)? So-called Mediterranean-type en- specialization. Paleolithic archaeologists, including me, have vironments exist in southern Europe, the Levant, North Af- vacillated to and fro in attempts to classify hunter-gatherer rica, and South Africa. The temporal and spatial records are foraging as more specialized or generalized. Some of the con- not yet well connected in climate science, but these regions fusion stems from how the terms may be applied at several can serve as geographic laboratories. The null hypothesis distinct scales of behavior. All hominins were omnivores, for would be that climate is not the driver of human behavioral example, and omnivorousness connotes a “generalist” feeding evolution. Siddall and colleagues (Siddall, Chappell, and Pot- strategy. Specialization is only informative if considered for ter 2006; Siddall et al. 2003) identify the climate intervals of one well-controlled aspect of econiche at a time. Middle Pa- 40–80, 120–140, and 160–200 ka as particularly variable based leolithic hominins were unusually specialized in their focus on Red Sea–Ocean proxies for global sea level changes. They on large game, with limited supplementation from small game identify the climate intervals of 80–120, 140–160, and 200– animals. Streamlining and elegance are common signatures 220 ka as least variable. The most recent contrasting pair of Stiner Trends versus Conservatism in the Predatory Niche S299 intervals, 40–80 and 80–120 ka, seem particularly suitable for the game: Middle and Upper Paleolithic hunting behaviors in the southern Caucasus. Current Anthropology 47:89–118. comparisons of MP and MSA strategies because faunal and Albert, R. M., O. Bar-Yosef, L. Meignen, and S. Weiner. 2003. Quantitative other data are fairly abundant. phytolith study of hearths from the Natufian and Middle Palaeolithic levels In order to undertake such a comparison, two other things of Hayonim Cave (Galilee, Israel). Journal of Archaeological Science 30:461– 480. must be done. First, zooarchaeologists must get away from Albert, R. M., F. Berna, and P. Goldberg. 2012. Insights on Neanderthal fire making lists of anecdotal evidence and focus on systematic, use at Kebara Cave (Israel) through high-resolution study of prehistoric quantitative analysis of the full scope of the animal diet (big, combustion features: evidence from phytoliths and thin sections. Quater- nary International 247:278–293. small, vertebrate, and shelled invertebrate). Richard Klein and Albert, R. M., S. Weiner, O. Bar-Yosef, and L. Meignen. 2000. Phytoliths in colleagues have set a remarkable precedent for a holistic mode the Middle Palaeolithic deposits of Kebara Cave, Mt. Carmel, Israel: study of data presentation in South Africa, a practice that sadly is of the plant materials used for fuel and other purposes. Journal of Archae- ological Science 27:931–947. out of vogue. The same limitations once applied to zooar- Alperson-Afil, N., and N. Goren-Inbar. 2010. Ancient flames and controlled chaeological work in Eurasia, but this has changed in the past use of fire, vol. 2 of The Acheulian site of Gesher Benot Ya’aqov. Dordrecht: few decades, and now there are complete faunal accounts for Springer. Anderson-Gerfaud, P. 1990. Aspects of behavior in the Middle Paleolithic: much of southern Eurasia. Second, although we cannot hope functional analysis of stone tools from southwest France. In The emergence to calibrate quantitative faunal data to plant consumption for of modern humans: an archaeological perspective. P. Mellars, ed. Pp. 389– the Paleolithic in any region, what we can do is compare the 418. Edinburgh: Edinburgh University Press. Assefa, Z. 2006. Faunal remains from Porc-Epic: paleoecological and zooar- faunal record in quantity and character with the record of chaeological investigations from a Middle Stone Age sites in southeastern durable plant processing equipment in a more controlled Ethiopia. Journal of Human Evolution 51(1):50–75. manner. The idea would be to look for evidence of heightened Aura, J. E., V. B. Villaverde, M. Perez Ripoll, R. Martinez Valle, and P. Cal- atayud Guillem. 2002. Big game and small prey: Paleolithic and Epipaleo- processing investments in plant use. Scattered reports, mainly lithic economy from Valencia (Spain). Journal of Archaeological Method and anecdotal, suggest that ground-stone artifacts are fairly com- Theory 9(3):215–268. mon in some MSA records. Given that these artifacts are not Baden-Powell, D. F. W. 1964. Gorham’s Cave, Gibraltar: report on the climatic equivalent of the marine Mollusca. In The excavation of Gorham’s Cave: a notable part of MP records, here lies a potential contrast Gibraltar, 1951–1954. J. Waechter, ed. Pp. 216–218. Bulletin of the Institute waiting for systematic consideration. Of course small MSA of Archaeology 4. London: University of London. milling equipment suitable for preparing mineral and organic Bailey, G. N., and N. C. Flemming. 2008. Archaeology of the continental shelf: marine resources, submerged landscapes and underwater archaeology. Qua- pigments must be distinguished from large implements suit- ternary Science Review 27:2153–2165. able for seed processing, but experimental evidence provides Barton, R. N. E. 2000. Mousterian hearths and shellfish: late Neanderthal some expectations. Such a strategy might provide an alter- activities on Gibraltar. In Neanderthals on the edge: papers presented at the conference marking the 150th anniversary of the Forbes’ Quarry discovery, native way of looking at the problem of how and why the Gibraltar. C. B. Stringer, R. N. E. Barton, and J. C. Finlayson, eds. Pp. 211– MP and MSA seem different from one another and assessing 220. Oxford: Oxbow. the independence of their evolutionary trajectories. Barton, R. N. E., A. P. Currant, Y. Fernandez-Jalvo, J. C. Finlayson, P.Goldberg, R. Macphail, P. B. Pettitt, and C. B. Stringer. 1999. Gibraltar Neanderthals and results of recent excavations in Gorham’s, Vanguard and Ibex Caves. Antiquity 73:13–23. Bar-Yosef, O., H. Laville, L. Meignen, A. M. Tillier, B. Vandermeersch, B. Arensburg, A. Belfer-Cohen, P. Goldberg, Y. Rak, and E. Tchernov. 1988. Acknowledgments La sepulture neanderthalienne de Kebara (unite XII). In L’homme de Ne- anderthal, vol. 5. M. Otte, ed. Pp. 17–24. Lie`ge: ERAUL. This paper is an outcome of the 2012 Wenner-Gren Sym- Bar-Yosef, O., B. Vandermeersch, B. Arensburg, A. Belfer-Cohen, P. Goldberg, posium held in Sweden and titled “Alternative Paths to Com- H. Laville, L. Meignen, et al. 1992. The excavations in Kebara Cave, Mt. plexity: Evolutionary Trajectories in the Middle Paleolithic Carmel. Current Anthropology 33(5):497–550. Bar-Yosef, O., B. Vandermeersch, B. Arensburg, P. Goldberg, H. Laville, and Middle Stone Age.” I am deeply grateful to the organizers L. Meignen, Y. Rak, E. Tchernov, and A.-M. Tillier. 1986. New data of this conference, Erella Hovers and Steve Kuhn, and to on the origin of modern man in the Levant. Current Anthropology 27: Wenner-Gren colleagues Leslie Aiello and Laurie Obbink for 63–64. their many efforts in making this one of the most stimulating Beyries, S. 1987. Variabilite´de l’industrie lithique asu Mousterien: approche functionelle sur quelques gisments franc¸aises. BAR International Series 328. and enriching symposia in which I have had the honor to Oxford: British Archaeological Reports. participate. I also thank Jamie Clark and two anonymous Binford, L. R. 1978. Nunamiut ethnoarchaeology. New York: Academic Press. Current Anthropology reviewers for their thoughtful and crit- ———. 1981. Bones: ancient men and modern myths. New York: Academic Press. ical remarks on an earlier version of this paper that certainly ———. 1984. Faunal remains from Klasies River Mouth. New York: Academic helped improve it. Press. ———. 1988. E´ tude taphonomique des restes fauniques de la grotte Vaufrey. In La Grotte Vaufrey a`Ce´nac et Saint-Julien (Dordogne): pale´oenvironnements, References Cited chronologie et activite´s humaines. J.-P. Rigaud, ed. Pp. 535–563. Me´moires de la Socie´te´ Pre´historique Franc¸aise, vol. 19. Paris: CNRS. Adler, D. S., and G. Bar-Oz. 2009. Seasonal patterns of prey acquisition and ———. 1991. When the going gets tough, the tough get going: Nunamiut inter-group competition during the Middle and Upper Palaeolithic of the local groups, camping patterns, and economic organization. In Ethnoar- southern Caucasus. In Evolution of human diets: integrating approaches to chaeological approaches to mobile campsites: hunter-gatherer and pastoralist the study of Palaeolithic subsistence. J.-J. Hublin and M. Richards, eds. Pp. case studies. C. S. Gamble and W. A. Boismer, eds. Pp. 25–137. Ann Arbor, 127–140. Dordrecht: Springer. MI: International Monographs in Prehistory. Adler, D. S., G. Bar-Oz, A. Belfer-Cohen, and O. Bar-Yosef. 2006. Ahead of ———. 1998. Hearth and home: the spatial analysis of ethnographically doc- S300 Current Anthropology Volume 54, Supplement 8, December 2013

umented rock shelter occupations as a template for distinguishing between du comple´ment alimentaire aux ressources utilitaires. J.-P. Brugal and J. Desse, human and hominid use of sheltered space. In Thirteenth UISPP Congress eds. Pp. 369–382. Antibes: APCDA. Proceedings: Forli, 8–14 September 1996. N. J. Conard and F. Wendorf, eds. Darlas, A. 2007. Le Mouste´rien de Gre`ce a` la lumie`re des re´centes recherches. Pp. 229–239. Forli, Italy: ABACO. L’anthropologie 111:346–366. Blanc, A. C. 1958–1961. Industria musteriana su calcare e su valve di Meretrix David, F., and C. Farizy. 1999. Mauran (Haute-Garonne, France). In The role chione associata con fossili di elefante e rinocerante: nuovi giacimenti costieri of early humans in the accumulation of European Lower and Middle Palaeo- del Capo de Leuca. Quaternaria 5:308–313. lithic bone assemblages (Ergebisse eines Kolloquiums). Pp. 267–277. Mono- Blasco, R., and J. Ferna´ndez Peris. 2009. Middle Pleistocene bird consumption graphien des Ro¨misch-Germanischen Zentralmuseums 42. Mainz: Rom- at level XI of Bolomor Cave (Valencia, Spain). Journal of Archaeological isch-Germanischen Zentralmuseums. Science 36:2213–2223. David, F., and T. Poulain. 1990. La faune de grands mammiferes des niveaux ———. 2012. A uniquely broad spectrum diet during the Middle Pleis- XI et XC de la Grotte du Renne a` Arcy-sur-Cure (Yonne): e´tude pre´limi- tocene at Bolomor Cave (Valencia, Spain). Quaternary International naire. In Pale´olithique Moyen Recent et Pale´olithique Superieur Ancien en 252:16–31. Europe. C. Farizy, ed. Pp. 319–323. Me´moires du Muse´e de Pre´histoire d’I˚le Blondel, J., and J. Aronson. 1999. Biology and wildlife of the Mediterranean de France 3. I˚le de France: E´ ditions de l’Association pour la Promotion de region. Oxford: Oxford University Press. la Recherche Arche´ologique. Bocherens, H. 2009. Neanderthal dietary habits: review of the isotopic evi- Delage, C., L. Meignen, and O. Bar-Yosef. 2000. Chert procurement and the dence. In Evolution of human diets: integrating approaches to the study of organization of lithic production in the Mousterian of Hayonim Cave (Is- Palaeolithic subsistence. J.-J. Hublin and M. Richards, eds. Pp. 214–250. rael). Journal of Human Evolution 38:A10–A11. Dordrecht: Springer. Delagnes, A., and L. Meignen. 2006. Diversity of lithic production systems Boe¨da, E. 1994. Le concept Levallois: variabilite´des me´thodes. Paris: CNRS. during the Middle Paleolithic in France: are there any chronological trends? Boe¨da, E., J. Connan, D. Dessort, S. Muhesen, N. Mercier, H. Valladas, and In Transitions before the transition: evolution and stability in the Middle N. Tisne´rat. 1996. Bitumen as a hafting material on Middle Palaeolithic Paleolithic and Middle Stone Age. E. Hovers and S. L. Kuhn, eds. Pp. 85– artefacts. Nature 380:336–338. 107. Santa Barbara, CA: Springer. Boone, J. 2002. Subsistence strategies and early human population history: Delagnes, A., and W. Rendu. 2011. Shifts in Neandertal mobility, technology an evolutionary ecological perspective. World Archaeology 34:6–25. and subsistence strategies in western France. Journal of Archaeological Science Bourguignon, L., J. P. Faivre, and A. Turq. 2004. Ramification des chaıˆnes 38:1771–1783. ope´ratoires: une spe´cificite´ du Mouste´rien? Pale´o 16:37–48. d’Errico, F., C. Henshilwood, M. Vanhaeren, and K. Van Niekerke. 2005. Bouzouggar A., R. N. E. Barton, M. Vanhaeren, F. D’errico, S. Collcutt, Nassarius kraussianus shell beads from Blombos Cave: evidence for T. Higham, E. Hodge, et al. 2007. 82,000-year-old shell beads from symbolic behaviour in the Middle Stone Age. Journal of Human Evo- North Africa and implications for the origins of modern human be- lution 48:3–24. havior. Proceedings of the National Academy of Sciences of the USA Discamps, E. 2011. Hommes et hye`nes face aux recompositions des com- 104(24):9964–9969. munaute´s d’Ongule´s (MIS 5–3): e´le´ments pour un cadre pale´oe´cologique Boyle, K. V. 2000. Reconstructing Middle Palaeolithic subsistence strategies des socie´te´s du Pale´olithique moyen et supe´rieur ancien d’Europe de in the south of France. International Journal of Osteoarchaeology 10:336– l’Ouest. PhD dissertation, Universite´ Bordeaux 1. 356. Erlandson, J. M., and M. L. Moss. 2001. Shellfish feeders, carrion eaters, and Brooks, A. 1984. San land-use patterns, past and present: implications for the archaeology of aquatic adaptations. American Antiquity 66(3):413–432. southern African prehistory. In Frontiers: southern African archaeology today. Farizy, C., F. David, and J. Jaubert, eds. 1994. Hommes et bisons du Pale´olithique M. Hall, G. Avery, D. M. Avery, M. L. Wilson, and A. J. B. Humphreys, Moyen a` Mauran (Haute Garonne). Gallia pre´histoire, suppl. 30. Paris: eds. Pp. 40–52. BAR International Series 207. Oxford: British Archaeological Reports. CNRS. Brugal, J.-P., and J. Jaubert. 1991. Les gisements pale´ontologiques Ple´istoce`nes Ferna´ndez-Jalvo, Y., and P. Andrews. 2000. The taphonomy of Pleistocene a` indices de fre´quentation humaine: un nouveau type de comportement caves, with particular reference to Gibraltar. In Neanderthals on the edge: de pre´dation? Pale´o 3:15–41. papers presented at the conference marking the 150th anniversary of the Forbes’ Burke, A. 2000. The view from Starosele: faunal exploitation at a Middle Quarry discovery, Gibraltar. C. B. Stringer, R. N. E. Barton, and J. C. Fin- Palaeolithic site in western Crimea. International Journal of Osteoarchaeology layson, eds. Pp. 171–182. Oxford: Oxbow. 10:325–335. Ferna´ndez Peris, J., V. Barciela Gonza´lez, R. Blasco, F. Cuartero, H. Fluck, P. Chase, P. G. 1986. The hunters of Combe Grenal: approaches to Middle Paleolithic San˜udo Die, and C. Verdasco. 2012. The earliest evidence of hearths in subsistence in Europe. BAR International Series 286. Oxford: British Ar- southern Europe: the case of Bolomor Cave (Valencia, Spain). Quaternary chaeological Reports. International 247:267–277. ———. 1989. How different was Middle Palaeolithic subsistence? a zooar- Finlayson, C., A. F. Darren, F. J. Jime´nez-Espejo, J. S. Carrio´n, G. Finlayson, chaeological perspective on the Middle to Upper Palaeolithic transition. In F. Giles Pacheco, J. Rodrı´guez Vidal, C. B. Stringer, and F. Martı´nez. 2008. The human revolution: behavioural and biological perspectives on the origins Gorham’s cave, Gibraltar: the persistence of a Neanderthal population. of modern humans. P. Mellars and C. Stringer, eds. Pp. 321–337. Princeton, Quaternary International 181:64–71. NJ: Princeton University Press. Foley, R., and C. Gamble. 2009. The ecology of social transitions in human Clark, J. L., and A. W. Kandel. 2013. The evolutionary implications of variation evolution. Philosophical Transactions of the Royal Society B 364:3267– in human hunting strategies and diet breadth during the Middle Stone Age 3279. of southern Africa. Current Anthropology 54(suppl. 8):S269–S287. Freeman, L. G. 1981. The fat of the land: notes on Paleolithic diet in Iberia. Cochard, D., J.-P. Brugal, E. Morin, and L. Meignen. 2012. Evidence of small In Omnivorous primates: gathering and hunting in human evolution.R.S. fast game exploitation in the Middle Paleolithic of Les Canalettes Aveyron, O. Harding and G. Teleki, eds. Pp. 104–165. New York: Columbia University France. Quaternary International 264:32–51. Press. Colonese, A., M. A. Mannino, D. E. Bar-Yosef Mayer, D. A. Fa, C. J. Galanidou, N. 2000. Patterns in caves: foragers, horticulturalists, and the use Finlayson, D. Lubell, and M. C. Stiner. 2011. Marine mollusc exploi- of space. Journal of Anthropological Archaeology 19:243–275. tation in Mediterranean prehistory: an overview. Quaternary Interna- Gamble, C. 1986. The Palaeolithic settlement of Europe. Cambridge: Cambridge tional 239:86–103. University Press. Conard, N. J., M. Bolus, and S. C. Mu¨nzel. 2012. Middle Paleolithic land use, ———. 1999. The Hohlenstein-Stadel revisited: the role of early humans in spatial organization and settlement intensity in the Swabian Jura, south- the accumulation of European Lower and Middle Palaeolithic bone assem- western Germany. Quaternary International 247:236–245. blages. In The role of early humans in the accumulation of European Lower Conard, N. J., and T. J. Prindiville. 2000. Middle Palaeolithic hunting econ- and Middle Palaeolithic bone assemblages (Ergebisse eines Kolloquiums). Pp. omies in the Rhineland. International Journal of Osteoarchaeology 10:286– 305–324. Monographien des Ro¨misch-Germanischen Zentralmuseums 42. 309. Mainz: Romisch-Germanischen Zentralmuseums. Costamagno, S., and V. Laroulandie. 2004. L’exploitation des petits verte´bre´s Garrard, A. N. 1983. The Palaeolithic faunal remains from Adlun and dans les Pyre´ne´es franc¸aises du Pale´olithique au Me´solithique: un inventaire their ecological context. In Adlun in the Stone Age: the excavations of taphonomique et arche´ozoologique. In Petits animaux et socie´te´s humaines: D. A. E. Garrod in the Lebanon, 1958–1963. D. A. Roe, ed. Pp. 397– Stiner Trends versus Conservatism in the Predatory Niche S301

413. BAR International Series 159(ii). Cambridge: British Archaeolog- northwestern Caucasus and the ecology of the Neanderthals. International ical Reports. Journal of Osteoarchaeology 10:368–378. Garrod, D. A. E., L. H. D. Buxton, G. E. Smith, D. M. A. Bate, R. C. Spiller, Hoffecker, J. R. 2009. Neanderthal and modern human diet in eastern Europe. M. A. C. Hinton, and P. Fischer. 1928. Excavation of a Mousterian rock- In Evolution of human diets: integrating approaches to the study of Palaeolithic shelter at Devil’s Tower, Gibraltar. Journal of the Royal Anthropological In- subsistence. J.-J. Hublin and M. Richards, eds. Pp. 87–98. Dordrecht: stitute of Great Britain and Ireland 58:33–113. Springer. Gaudzinski, S. 1995. Wallertheim revisited: a reanalysis of the fauna from the Hortola`, P., and B. Martı´nez-Navarro. 2013. The Quaternary megafaunal ex- Middle Palaeolithic site of Wallertheim (Rheinhessen/Germany). Journal of tinction and the fate of Neanderthals: an integrative working hypothesis. Archaeological Science 22:51–66. Quaternary International 295:69–72. ———. 2000. On the variability of Middle Palaeolithic procurement tactics: Jacob-Friesen, K. H. 1956. Eiszeitliche Elephantenja¨ger in der Lu¨neburger the case of Salzgitter Lebenstedt, Northern Germany. International Journal Heide. Jarbu¨ch des Ro¨misch-Germanischen Zentralmuseums, Mainz 3:1– of Osteoarchaeology 10:396–406. 22. ———. 2004. A matter of high resolution? the Eemian Interglacial (OIS 5e) Jaubert, J., M. Lorblanchet, H. Laville, R. Slott-Moller, A. Turq, and J.-P. in north-central Europe and Middle Palaeolithic subsistence. International Brugal. 1990. Les chasseurs d’aurochs de la Borde. Documents d’Archeologie Journal of Osteoarchaeology 14(3/4):201–211. Francaise, no. 27. Paris: Maison des Sciences de l’Homme. Gaudzinski, S., and W. Roebroeks. 2000. Adults only: reindeer hunting at the Jones, M. 2009. Moving north: archaeobotanical evidence for plant diet in Middle Palaeolithic site Salzgitter Lebenstedt, northern Germany. Journal Middle and Upper Paleolithic Europe. In Evolution of human diets: inte- of Human Evolution 38:497–521. grating approaches to the study of Palaeolithic subsistence. J.-J. Hublin and Gaudzinski-Windheuser, S., and L. Niven. 2009. Hominin subsistence M. Richards, eds. Pp. 171–180. Dordrecht: Springer. patterns during the Middle and Late Paleolithic in northwestern Eu- Karkanas, P., R. Shahack-Gross, A. Ayalon, M. Bar-Matthews, R. Barkai, A. rope. In Evolution of human diets: integrating approaches to the study Frumkin, A. Gopher, and M. C. Stiner. 2007. Evidence for habitual use of of Palaeolithic subsistence. J.-J. Hublin and M. Richards, eds. Pp. 99– fire at the end of the Lower Paleolithic: site formation processes at Qesem 111. Dordrecht: Springer. Cave, Israel. Journal of Human Evolution 53(2):197–212. Goldberg, P., and O. Bar-Yosef. 1998. Site formation processes in Kebara and Keeley, L. H. 1995. Protoagricultural practices among hunter-gatherers: a Hayonim caves and their significance in Levantine prehistoric caves. In cross-cultural survey. In Last hunters–first farmers: new perspectives on the Neanderthals and modern humans in western Asia. T. Akazawa, K. Aoki, and prehistoric transition to agriculture. T. D. Price and A. B. Gebauer, eds. Pp. O. Bar-Yosef, eds. Pp. 107–125. New York: Plenum. 243–272. Santa Fe, NM: School of American Research. Goldberg, P., H. Dibble, F. Berna, D. Sandgathe, S. J. P. McPherron, and Kelly, R. 1995. The foraging spectrum: diversity in hunter-gatherer lifeways. A. Turq. 2012. New evidence on Neandertal use of fire: examples from Washington, DC: Smithsonian Institution. Roc de Marsal and Pech de l’Aze´IV.Quaternary International 247:325– Klein, R. G. 1999. The human career: human biological and cultural origins. 340. 2nd edition. Chicago: University of Chicago Press. Klein, R. G., and K. Scott. 1986. Reanalysis of faunal assemblages from the Gopher, A., R. Barkai, R. Shimelmitz, M. Khalaily, C. Lemorini, I. Heshkovitz, Haua Fteah and other Late Quaternary archaeological sites in Cyrenaican and M. C. Stiner. 2005. Qesem Cave: an Amudian site in central Israel. Libya. Journal of Archaeological Science 13:515–542. Journal of the Israel Prehistory Society 35:69–92. Klein, R. G., and T. Steele. 2008. Gibraltar data are too sparse to inform on Gorecki, P. P. 1991. Horticulturalists as hunter-gatherers: rock shelter Neanderthal exploitation of coastal resources. Proceedings of the National usage in Papua New Guinea. In Ethnoarchaeological approaches to mo- Academy of Sciences of the USA 105(51):E115. bile campsites: hunter-gatherer and pastoralist case studies. C. S. Gamble Knecht, H. 1997. Projectile technology. New York: Plenum. and W. A. Boismer, eds. Pp. 237–262. Ann Arbor, MI: International Koller, J., U. Baumer, and D. Mania. 2001. High-tech in the Middle Palaeo- Monographs in Prehistory. lithic: Neandertal-manufactured pitch identified. European Journal of Ar- Gowlett, J. A. J., J. Hallos, S. Hounsell, V. Brant, and N. C. Debenham. 2005. chaeology 4:385–397. Beeches Pit: archaeology, assemblage dynamics and early fire history of a Kuhn, S. L. 1995. Mousterian lithic technology: an ecological perspective. Prince- Middle Pleistocene site in East Anglia, UK. Journal of Eurasian Prehistory ton, NJ: Princeton University Press. 3:3–38. ———. 2013. Roots of the Middle Paleolithic in Eurasia. Current Anthropology Grayson, D. K. and F. Delpech. 1994. The evidence for Middle Palaeolithic 54(suppl. 8):S255–S268. scavenging from Couche VIII, Grotte Vaufrey (Dordogne, France). Journal Kuhn, S. L., and M. C. Stiner. 2006. What’s a mother to do? a hypothesis of Archaeological Science 21:359–375. about the division of labor among Neandertals and modern humans in ———. 1998. Changing diet breadth in the early Upper Palaeolithic of south- Eurasia. Current Anthropology 47:953–980. western France. Journal of Archaeological Science 25:1119–1129. ———. 2010. Summary of findings and conclusion. In Culture and biology ———. 2003. Ungulates and the Middle-to-Upper Paleolithic transition at at a crossroads: the Middle Pleistocene record of Yarimburgaz Cave (Thrace, Grotte XVI (Dordogne, France). Journal of Archaeological Science 30:1633– Turkey). F. C. Howell, G. Arsebu¨k, S. L. Kuhn, M. O¨ zbas¸aran, and M. C. 1648. Stiner, eds. Pp. 301–319. Istanbul: Zero Books/Ege. Griggo, C. 2004. Mousterian fauna from Dederiyeh Cave and comparisons Lalueza-Fox, C., A. Rosas, A. Estalrrich, E. Gigli, P. F. Campos, A. Garcı´a- with fauna from Umm El Tlel and Douara Cave. Pale´orient 30(1):149– Tabernero, S. Garcı´a-Vargas, et al. 2011. Genetic evidence for partrilocal 162. mating behavior among Neandertal groups. Proceedings of the National Haydal, J., T. Oguchi, T. Akazawa, S. Muhesen, Y. Dodo, Y. Mizoguchi, Y. Academy of Sciences of the USA 108(1):250–253. Abe, Y. Nishiaki, and S. Ohta. 1995. Neanderthal infant burial from the Lam, Y. M., and O. M. Pearson. 2005. Bone density studies and the inter- Dederiyeh cave in Syria. Pale´orient 21:77–86. pretation of the faunal record. Evolutionary Anthropology 14(3):99–108. Henrich, J. 2004. Demography and cultural evolution: how adaptive cultural Lambeck, K. 1996. Sea-level change and shore-line evolution in Aegean Greece processes can produce maladaptive losses: the Tasmanian case. American since Upper Paleolithic time. Antiquity 70:588–611. Antiquity 69(2):197–214. Laroulandie, V. 2004. Exploitation des ressources aviaires durant le Pale´o- Henry, A. G., A. S. Brooks, and D. R. Piperno. 2011. Microfossils in calculus lithique en France: bilan critique et perspectives. In Petits animaux et socie´te´s demonstrate consumption of plants and cooked foods in Neanderthal diets humaines: du comple´ment alimentaire aux ressources utilitaires. J.-P. Brugal (Shanidar III, Iraq; Spy I and II, Belgium). Proceedings of the National and J. Desse, eds. Pp. 163–172. Antibes: APCDA. Academy of Sciences of the USA 108(2):486–491. Lemorini, C. 2000. Reconnaıˆtre de tactiques d’exploitation du milieu au pale´o- Henshilwood, C., F. d’Errico, M. Vanhaeren, K. Van Niekerk, and Z. lithique moyen: la contribution de l’analyse fonctionelle: e´tude fonctionelle des Jacobs. 2004. Middle Stone Age shell beads from South Africa. Science industries lithiques au Grotta Breuil (Latium, Italie) et de La Colombette 304:404. (Bonnieux, Valcluse, France). BAR International Series 858. Oxford: British Hoffecker, J. F., G. Baryshnikov, and O. Potapova. 1991. Vertebrate re- Archaeological Reports. mains from the Mousterian site of Il’skaya I (northern Caucasus, Lyman, R. L. 1984. Bone density and differential survivorship of fossil classes. USSR): new analysis and interpretation. Journal of Archaeological Sci- Journal of Anthropological Archaeology 3:259–299. ence 18:113–147. Madella, M., E. Hovers, M. K. Jones, P. Goldberg, and Y. Goren. 2002. The Hoffecker, J. F., and N. Cleghorn. 2000. Mousterian hunting patterns in the exploitation of plant resources by Neanderthals in Amud Cave (Israel): the S302 Current Anthropology Volume 54, Supplement 8, December 2013

evidence from phytolith studies. Journal of Archaeological Science 29:703– Peresani, M., I. Fiore, M. Gala, M. Romandini, and A. Tagliacozzo. 2011. Late 719. Neandertals and the intentional removal of feathers as evidenced from bird Marean, C. W., M. Bar-Matthews, J. Bernatchez, E. Fisher, P. Goldberg, A. I. bone taphonomy at Fumane Cave 44 ky B.P., Italy. Proceedings of the Na- R. Herries, Z. Jacobs, et al. 2007. Early human use of marine resources and tional Academy of Sciences of the USA 108:3888–3893. pigment in South Africa during the Middle Pleistocene. Nature 449:905– Pianka, E. R. 1978. Evolutionary ecology. 2nd edition. New York: Harper & 909. Row. Martı´nez-Molina, K. 2005. Ana´lisis functional de industrias lı´ticas del Plies- Pike-Tay, A., V. Cabrera Valde´s, and F. Bernaldo de Quiro´s. 1999. Seasonal toceno superior: el Paleolı´tico Medio del Abric Romanı´ (Capellades, Bar- variations of the Middle–Upper Paleolithic transition at El Castillo, Cueva celona) y del Paleolı´tico Superior de U¨ c¸ag˘ızlı (Hatay, Turquı´a) y del Molı´ Morin and El Pendo (Cantabria, Spain). Journal of Human Evolution 36: del Salt (Vimbodı´, Tarragona): cambios en los patrones funcionales entre 283–317. el Paleolı´tico Medio y el Superior. PhD dissertation, Universitat Rovira I Plisson, H., and S. Beyries. 1998. Pointes ou outils triangulaires? donne´es Virgili, Tarragona. fonctionnelles dans le Mouste´rien Levantin. Pale´orient 24(1):5–24. Martinson, D. G., N. G. Pisias, J. D. Hays, J. Imbrie, T. C. Moore, and N. J. Poteete, A. R., and E. Ostrom. 2004. Heterogeneity, group size and collective Shackleton. 1987. Age dating and the orbital theory of the ice ages: devel- action: the role of institutions in forest management. Development and opment of a high-resolution 0 to 300,000-year chronostratigraphy. Qua- Change 35(3):435–461. ternary Research 27:1–29. Preece, R. C., J. A. J. Gowlett, S. A. Parfitt, D. R. Bridgland, and S. G. Lewis. McBrearty, S., and A. Brooks. 2000. The revolution that wasn’t: a new inter- 2006. Humans in the Hoxnian: habitat, context and fire use at Beeches Pit, pretation of the origin of modern human behavior. Journal of Human West Stow, Suffolk, UK. Journal of Quaternary Science 21:485–496. Evolution 39:456–463. Premo, L. S., and S. L. Kuhn. 2010. Modeling effects of local extinctions on Meignen, L., O. Bar-Yosef, and P. Goldberg. 1989. Les structures de com- culture change and diversity in the Paleolithic. PLoS ONE 5:15582. bustion Mouste´riennes de la grotte de Ke´bara (Mont Carmel, Israe¨l). In Rabinovich, R., S. Gaudzinski-Windheuser, and N. Goren-Inbar. 2008. Sys- Nature et fonctions des foyers prehistoriques. M. Olive and Y. Taborin, eds. tematic butchering of fallow deer (Dama) at the early middle Pleistocene Pp. 141–146. Memoires du Muse´e de Pre´histoire d’I˚le de France, vol. 2. Acheulian site of Gesher Benot Ya‘aqov (Israel). Journal of Human Evolution Nemours: APRAIF. 54(1):134–149. Meignen, L., O. Bar-Yosef, M. Stiner, S. Kuhn, P. Goldberg, and S. Weiner. Rabinovich, R., and E. Tchernov. 1995. Chronological, paleoecological and 2010. Apport des analyses mine´ralogiques (en spectrome´trie infrarouge a` taphonomical aspects of the Middle Paleolithic site of Qafzeh, Israel. In transforme´e de fourier) a` l’interpre´tation des structures anthropiques: les Archaeozoology of the Near East, vol. 2. H. Buitenhuis and H.-P. Uerpmann, concentrations osseuses dans les niveaux Mouste´riens des Grottes de Ke´bara eds. Pp. 45–71. Leiden: Backhuys. et Hayonim (Israe¨l). PALEO 3(suppl.):93–107. Rendu, W. 2010. Hunting behavior and Neanderthal adaptability in the Late Meignen, L., J. D. Speth, and M. C. Stiner. 2006. Middle Paleolithic settlement Pleistocene site of Pech-de-l’Aze´I.Journal of Archaeological Science 37:1798– patterns in the Levant. In Transitions before the transition: evolution and 1810. stability in the Middle Paleolithic and Middle Stone Age. E. Hovers and S. Rendu, W., S. Costamagno, L. Meignen, and M. C. Soulier. 2012. Monospecific L. Kuhn, eds. Pp. 149–169. New York: Kluwer. faunal spectra in Mousterian contexts: implications for social behavior. Mellars, P. 1989. Major issues in the emergence of modern humans. Current Quaternary International 247:50–58. Anthropology 30:349–385. Reynaud Savioz, N., and P. Morel. 2005. La faune de Nadaouiyeh Aı¨n Askar ———. 1996. The Neanderthal legacy: an archaeological perspective from west- (Syrie centrale, ple´istoce`ne moyen): aperc¸u et perspectives. Revue Pale´o- ern Europe. Princeton, NJ: Princeton University Press. biologie Gene`ve 10:31–35. Mirazo´n Lahr, M., and R. Foley. 2003. Demography, dispersal and human Riel-Salvatore, J., and C. M. Barton. 2004. Late Pleistocene technology, eco- evolution in the Last Glacial period. In Neandertals and modern humans in nomic behavior, and land-use dynamics in southern Italy. American Antiq- the European landscape during the Last Glaciation. T. H. Van Andel and W. uity 69(2):257–274. Davies, eds. Pp. 241–256. Cambridge: McDonald Institute for Archaeolog- Rightmire, G. P. 2003. Brain size and encephalization in early to mid- ical Research, University of Cambridge. Pleistocene Homo. American Journal of Physical Anthropology 124(2): Moncel, M.-H., and C. Daujeard. 2012. The variability of the Middle Palaeo- 109–123. lithic on the right bank of the Middle Rhoˆne Valley (southeast France): Roebroeks, W., M. J. Sier, T. Kellberg Nielsen, D. de Loecker, J. M. Pare´s, C. technical traditions or functional choices? Quaternary International 247: E. S. Arps, and H. J. Mu¨cher. 2012. Use of red ochre by early Neandertals. 103–124. Proceedings of the National Academy of Sciences of the USA 109(6):1889– Mu¨ller-Beck, H. 1988. The ecosystem of the “Middle Paleolithic” (late Lower 1894. Paleolithic) in the Upper Danube region: a stepping stone to the Upper Roebroeks, W., and P. Villa. 2011. On the earliest evidence for habitual use Paleolithic. In Upper Pleistocene prehistory in western Eurasia. H. Dibble of fire in Europe. Proceedings of the National Academy of Sciences of the and A. Montet-White, eds. Pp. 233–254. Monograph 54. Philadelphia: Uni- USA 108:5209–5214. versity Museum, University of Pennsylvania. Rolland, N. 2004. Was the emergence of home bases and domestic fire a Munro, N. 2004. Zooarchaeological measures of hunting pressure and oc- punctuated event? a review of the Middle Pleistocene record in Eurasia. cupation intensity in the Natufian. Current Anthropology 45(suppl.):S5–S33. Asian Perspectives 43:248–280. Mu¨nzel, S., and N. J. Conard. 2004. Change and continuity in subsistence Rosell, J., I. Ca´ceres, R. Blasco, M. Benna`sar, P. Bravo, G. Campeny, M. during the Middle and Upper Palaeolithic in the Ach Valley of Swabia Esteban-Nadal, et al. 2012. A zooarchaeological contribution to establish (south-west Germany). International Journal of Osteoarchaeology 14:225– occupational patterns at Level J of Abric Romanı´(Barcelona, Spain). Qua- 243. ternary International 247:69–84. Oswalt, W. H. 1976. An anthropological analysis of food-getting technology. New Sandgathe, D. M., H. L. Dibble, P. Goldberg, S. P. Mcpherron, A. Turq, L. York: Wiley. Niven, and J. Hodgkins. 2011. On the role of fire in Neandertal adaptations Palma di Cesnola, A. 1965. Notizie preliminari sulla terza campagna di scavi in western Europe: evidence from Pech de l’Aze´ IV and Roc de Marsal, nella Grotta del Cavallo (Lecce). Rivista di Scienze Preistoriche 25:3–87. France. Paleoanthropology 2011:216Ϫ242. ———. 1969. Il musteriano della Grotta del Poggio a Marina di Camerota Shea, J. 1989. A functional study of the lithic industries associated with hom- (Salerno). In Estratto dagli scritti sul Quaternario in onore di Angelo Pasa. inid fossils in the Kebara and Qafzeh caves, Israel. In The human revolution. Pp. 95–135. Verona: Museo Civico di Storia Naturale. P. Mellars and C. Stringer, eds. Pp. 611–625. Princeton, NJ: Princeton Patou-Mathis, M. 1993. E´ tude taphonomique et pale´oethnologique de la faune University Press. de l’Abri des Canalettes. In L’Abri des Canalettes: un habitat Mouste´rien sur ———. 1991. The behavioral significance of Levantine Mousterian industrial les Grands Causses (Nant, Aveyron). L. Meignen, ed. Pp. 199–237. Paris: variability. PhD dissertation, Harvard University, Cambridge, MA. CNRS. ———. 1993. Lithic use-wear evidence for hunting by Neandertals and early ———. 2000. Neanderthal subsistence behaviours in Europe. International modern humans from the Levantine Mousterian. In Hunting and animal Journal of Osteoarchaeology 10:379–395. exploitation in the later Palaeolithic and Mesolithic of Eurasia. G. L. Peterkin, Peresani, M. 2012. Fifty thousand years of flint knapping and tool shaping H. Bricker, and P. Mellars, eds. Pp. 189–197. Archaeological Papers of the across the Mousterian and Uluzzian sequence of Fumane Cave. Quaternary American Anthropological Association, vol. 4. Washington, DC: American International 247:125–150. Anthropological Association. Stiner Trends versus Conservatism in the Predatory Niche S303

———. 2009. The impact of projectile weaponry on Late Pleistocene hominin Stiner, M. C., R. Barkai, and A. Gopher. 2011. Hearth-side socioeconomics, evolution. In Evolution of human diets: integrating approaches to the study hunting and paleoecology during the late Lower Paleolithic at Qesem Cave, of Palaeolithic subsistence. J.-J. Hublin and M. Richards, eds. Pp. 189–199. Israel. Journal of Human Evolution 60(2):213–233. Dordrecht: Springer. Stiner, M. C., and S. L. Kuhn. 1992. Subsistence, technology, and adaptive Siddall, M., J. Chappell, and E.-K. Potter. 2006. Eustatic sea level during past variation in Middle Paleolithic Italy. American Anthropologist 94:12– interglacials. In The climate of past interglacials. F. Sirocko, T. Litt, M. Claus- 46. sen, and M.-F. Sanchez-Goni, eds. Pp. 75–92. Developments in Quaternary ———. 2006. Changes in the “connectedness” and resilience of Paleo- Science 7. Amsterdam: Elsevier. lithic societies in Mediterranean ecosystems. Human Ecology 34(5):693– Siddall, M., E. Rohling, A. Almogi-Labin, C. Hemleben, D. Meischner, I. 712. Schmelzer, and D. A. Smeed. 2003. Sea-level fluctuations during the last Stiner, M. C., S. L. Kuhn, T. A. Surovell, P. Goldberg, L. Meignen, S. Weiner, glacial cycle. Nature 423(6942):853–858. and O. Bar-Yosef. 2001. Bone preservation in Hayonim Cave (Israel): a Snodgrass, J. J., W. R. Leonard, and M. L. Robertson. 2009. The energetics macroscopic and mineralogical study. Journal of Archaeological Science 28: of encephalization in early hominids. In Evolution of human diets: integrating 643–659. approaches to the study of Palaeolithic subsistence. J.-J. Hublin and M. Rich- Stiner, M. C., and N. D. Munro. 2011. On the evolution of Paleolithic diet ards, eds. Pp. 15–29. Dordrecht: Springer. and landscape at Franchthi Cave (Peloponnese, Greece). Journal of Human Speth, J. D. 2006. Housekeeping, Neandertal-style: hearth placement and mid- Evolution 60:618–636. den formation in Kebara Cave (Israel). In Transitions before the transition: Stiner, M. C., N. D. Munro, and T. A. Surovell. 2000. The tortoise and the evolution and stability in the Middle Paleolithic and Middle Stone Age.E. hare: small game use, the broad spectrum revolution, and Palaeolithic de- Hovers and S. L. Kuhn, eds. Pp. 171–188. New York: Springer. mography. Current Anthropology 41:39–73. Speth J. D., and J. L. Clark. 2006. Hunting and overhunting in the Levantine Stiner, M. C., N. D. Munro, T. A. Surovell, E. Tchernov, and O. Bar-Yosef. late Middle Palaeolithic. Before Farming 3:1–42. 1999. Paleolithic population growth pulses evidenced by small animal ex- Speth, J. D., and E. Tchernov. 1998. The role of hunting and scavenging in ploitation. Science 283:190–194. Neanderthal procurement strategies: new evidence from Kebara Cave (Is- Straus, L. G. 1982. Carnivores and cave sites in Cantabrian Spain. Journal of rael). In Neanderthals and modern humans in west Asia. T. Akazawa, K. Anthropological Research 38:75–96. Aoki, and O. Bar-Yosef, eds. Pp. 223–239. New York: Plenum. Stringer, C. B., J. C. Finlayson, R. N. E. Barton, Y. Ferna´ndez-Jalvo, I. Ca´ceres, ———. 2001. Neandertal hunting and meat-processing in the Near East: R. C. Sabin, E. J. Rhodes, et al. 2008. Neanderthal exploitation of marine evidence from Kebara Cave (Israel). In The early human diet: the role of mammals in Gibraltar. Proceedings of the National Academy of Sciences of meat. C. Stanford and H. Bunn, eds. Pp. 52–72. Oxford: Oxford University the USA 105(38):14319–14324. Press. Stuart, A. J. 1991. Mammalian extinctions in the Late Pleistocene of northern ———. 2002. Middle Paleolithic tortoise use at Kebara Cave (Israel). Journal Eurasia and North America. Biological Reviews 66:453–562. of Archaeological Science 29(5):471–483. Thieme, H. 1997. Lower Palaeolithic hunting spears from Germany. Nature ———. 2007. The Middle Paleolithic occupations at Kebara Cave: a faunal 385:807–810. perspective. In Kebara Cave Mt. Carmel, Israel: the Middle Paleolithic ar- Valensi, P. 2000. The archaeozoology of Lazaret Cave (Nice, France). Inter- chaeology, part 1. O. Bar-Yosef and L. Meignen, eds. Pp. 165–260. American national Journal of Osteoarchaeology 10:357–367. School of Prehistoric Research Bulletin 49. Cambridge, MA: Peabody Mu- Valensi, P., and E. Psathi. 2004. Faunal exploitation during the Middle Pa- seum. laeolithic in south-eastern France and north-western Italy. International Starkovich, B. 2011. Subsistence change during the Middle to Upper Paleo- Journal of Osteoarchaeology 14:256–272. lithic at Klissoura Cave 1 (Peloponnese, Greece). PhD dissertation, Uni- Vallverdu´, J., S. Alonso, A. Bargallo´, R. Bartrolı´, G. Campeny, A` .Car- versity of Arizona, Tucson. rancho, I. Expo´sito, et al. 2012. Combustion structures of archaeolog- Steele, T. 2004. Variation in mortality profiles of red deer (Cervus elaphus)in ical level O and Mousterian activity areas with use of fire at the Abric Middle Palaeolithic assemblages from western Europe. International Journal Romanı´ rockshelter (NE Iberian Peninsula). Quaternary International of Osteoarchaeology 14:307–320. 247:313–324. Steele, T., and E. Alverez-Fernandez. 2011. Initial investigations into the ex- Van Andel, T. H., W. Davies, B. Weninger, and O. Jo¨ris. 2003. Archaeological ploitation of coastal resources in North Africa during the Late Pleistocene dates as proxies for the spatial and temporal human presence in Europe: at Grottes des Contrebandiers, Morocco. In Trekking the shore: changing a discourse on the method. In Neandertals and modern humans in the coastlines and the antiquity of coastal settlement. N. F. Bicho, J. A. Haws, European landscape during the Last Glaciation. T. H. Van Andel and W. and L. G. Davis, eds. Pp. 383–403. New York: Springer. Davies, eds. Pp. 21–29. Cambridge: McDonald Institute for Archaeological Steele, T. E., and R. G. Klein. 2009. Late Pleistocene subsistence strategies and Research, University of Cambridge. resource intensification in Africa. In Evolution of human diets: integrating Van Andel T. H., and P. C. Tzedakis. 1996. Palaeolithic landscape of Europe approaches to the study of Palaeolithic subsistence. J.-J. Hublin and M. Rich- and environs, 150,000–25,000 years ago: an overview. Quaternary Research ards, eds. Pp. 113–126. Dordrecht: Springer. Review 15:481–500. Stephens, D. W., and J. R. Krebs. 1986. Foraging theory. Princeton, NJ: Prince- Vandermeersch, B. 1989. The evolution of modern humans: recent evidence ton University Press. from southwest Asia. In The human revolution: behavioural and biological Stiner, M. C. 1990. The use of mortality patterns in archaeological studies of perspectives on the origins of modern humans. P. Mellars and C. B. Stringer, hominid predatory adaptations. Journal of Anthropological Archaeology 9: eds. Pp. 155–164. Edinburgh: Edinburgh University Press. 305–351. Vaquero, M., M. G. Chacon, M. D. Garcia-Anton, B. Gomez de Soler, K. ———. 1994. Honor among thieves: a zooarchaeological study of Neandertal Martinez, and F. Cuartero. 2012. Time and space in the formation of lithic ecology. Princeton, NJ: Princeton University Press. assemblages: the example of Abric Romanı´Level J. Quaternary International ———. 2001. Thirty years on the “broad spectrum revolution” and Palaeo- 247:162–181. lithic demography. Proceedings of the National Academy of Sciences of the Vaquero, M., and I. Pasto´. 2001. The definition of spatial units in Middle USA 98(13):6993–6996. Palaeolithic sites: the hearth-related assemblages. Journal of Archaeological ———. 2002. On in situ attrition and vertebrate body part profiles. Journal Science 28:1209–1220. of Archaeological Science 29:979–991. Villa, P. 1990. Torralba and Aridos: elephant exploitation in Middle Pleistocene ———. 2005. The faunas of Hayonim Cave (Israel): a 200,000-year record of Spain. Journal of Human Evolution 19:299–309. Paleolithic diet, demography and society. American School of Prehistoric Villa, P., F. Bon, and J.-C. Castel. 2001. Fuel, fire and fireplaces in the Paleolithic Research Bulletin 48. Cambridge, MA: Peabody Museum. of western Europe: review of Economie des combustibles au Pale´olithique,I, ———. 2009. Prey choice, site occupation intensity and economic diversity by The´ry-Parisot, ed. Review of Archaeology 23(1):33–42. across the Middle to early Upper Palaeolithic at U¨ c¸ag˘ızlı Caves I and II Villa, P., and M. Soressi. 2000. Stone tools in carnivore sites: the case of Bois (Hatay, Turkey). Before Farming. http://www.waspress.co.uk/journals Roche. Journal of Anthropological Research 56:187–215. /beforefarming/journal_20093/abstracts/index.php#200933. Vitagliano, S. 1984. Nota sul Pontiniano della Grotta dei Moscerini, Gaeta Stiner, M. C., G. Arsebu¨k, and F. C. Howell. 1996. Cave bears and Paleolithic (Latina). Il paleolitico e il mesolitico nel Lazio: atti della XXIV Riunione artifacts in Yarımburgaz Cave, Turkey: dissecting a palimpsest. Geoar- Scientifica. Pp. 155–164. Florence: Istituto Italiano di Preistoria e Proto- chaeology 11(4):279–327. storia. S304 Current Anthropology Volume 54, Supplement 8, December 2013

Weiner, S., P. Goldberg, and O. Bar-Yosef. 2002. Three-dimensional distri- fille, J. Feathers, et al. 2005. The archaeology of Aduma Middle Stone Age bution of minerals in the sediments of Hayonim Cave, Israel: diagenetic sites in the Awash Valley, Ethiopia. Paleoanthropology 10:25–100. processes and archaeological implications. Journal of Archaeological Science Yeshurun, R., G. Bar-Oz, and M. Weinstein-Evron. 2007. Modern hunting 29(11):1289–1308. behavior in the early Middle Paleolithic: faunal remains from Misliya Cave, Yellen, J. E., A. S. Brooks, E. Cornelissen, R. G. Klein, M. Mehlman, and K. Mount Carmel, Israel. Journal of Human Evolution 53(6):656–677. Stewart. 1995. A Middle Stone Age worked bone industry from Katanda, Zilha˜o, J., D. E. Angelucci, E. Badal-Garcı´a, F. D’errico, F. Daniel, L. Dayet, upper Semliki River valley (Kivu), Zaire. Science 268:553–556. K. Douka, et al. 2010. Symbolic use of marine shells and mineral pigments Yellen, J., A. S. Brooks, D. M. Helgren, M. Tappen, S. H. Ambrose, R. Bonne- by Iberian Neandertals. Proceedings of the National Academy of Sciences of the USA 107(3):1023–1028. Current Anthropology Volume 54, Supplement 8, December 2013 S305

Technological Trends in the Middle Stone Age of South Africa between MIS 7 and MIS 3

by Sarah Wurz

The range of technological elements that marks the Middle Stone Age originated more than 300,000 years ago and formed the basic tool kit for an extended period of time. No spatial and chronological patterns can be identified from the Early Middle Stone Age until marine isotope stage (MIS) 5, and there is no cumulative trend of increasing complexity and diversity; instead, periods of complexity come and go. The Howiesons Poort and Still Bay techno- complexes, broadly associated with MIS 4, are recognized across various ecological zones of South Africa. These techno-complexes contain relatively more retouched tools and exhibit heightened levels of what is described as innovative practices. The Howiesons Poort is the best understood industry in the Middle Stone Age of South Africa. Its unifying technological characteristic is the almost exclusive use of a blade and bladelet production system, but subtle changes in types of backed artefacts, other retouched tools, and raw material exploitation patterns occur through time. Technological preferences change in the ensuing MIS 3 period, and other strategies and implement types become popular, particularly unifacial points, but trends once again are less clear. This historical review of the technological diversity in the Middle Stone Age of South Africa emphasizes that the roots of some innovations may lie in the earlier Middle Stone Age and that innovation is best understood within the context of local historical trajectories of technological change in South Africa.

The Middle Stone Age period of South Africa is a key milieu Sibudu Cave, Border Cave, and Kathu Pan—also contribute within which Homo sapiens developed, diverged, and migrated significantly to the latest insights. (e.g., Behar et al. 2008; Dusseldorp, Lombard, and Wurz 2013; The dominant narrative for the MSA of South Africa Lombard, Schlebusch, and Soodyall, forthcoming; Pickrell et emerging from a number of recent publications (e.g., Comp- al. 2012; Schlebusch et al. 2012). Numerous Middle Stone ton 2011; McCall 2007; McCall and Thomas 2012; Mellars Age cave and open-air sites occur all over South Africa and 2006; Mourre, Villa, and Henshilwood 2010; Soriano, Villa, bear testimony to the successful adaptation of hunter-gath- and Wadley 2007; Villa, Delagnes, and Wadley 2005; Villa et erers over the past 300,000 years. Stratified sequences from al. 2010; Ziegler et al. 2013) is that “Middle Paleolithic–like” caves acted as culture-stratigraphic anchors since the “Middle variability occurred from the earliest MSA until around 77,000 Stone Age” (MSA) concept was established (Goodwin 1928). years ago. The first time that “Upper Paleolithic–like” vari- A recent exercise in ordering Stone Age sites with relatively ability developed was during marine isotope stage (MIS) 4, recent chronometric dates shows that some progress has been with a spurt of innovation and complexity associated with made in expanding the MSA spatiotemporal framework of the Still Bay and Howiesons Poort techno-complexes. The South Africa and Lesotho (Lombard and Haidle 2012). The innovations include formal bone tools, beads, processing of aim here is to describe MSA technological developments from ochre, engraved ochre and ostrich eggshell, and compound South Africa, a large area covering more than 1.2 million adhesive manufacture. The technological sophistication of the square kilometers. The considerable size of the area precludes Howiesons Poort and the Still Bay is evident in the manu- an exhaustive discussion, and only fairly general trends are facture of geometric-backed tools and bifacial points, specific addressed. The noticeable growth in investigations into the techniques of blade and bladelet reduction, pressure flaking, MSA in the past decade involve some cave sites from the Cape and heat treatment of stone. These levels of innovation and coastal area, which necessarily biases this discussion toward complexity are thought to signify symbolically mediated be- these locations, but sites from other regions—for example, havior (Henshilwood and Marean 2003) or cognitive com- plexity (Wadley, Hodgkiss, and Grant 2009), perhaps even Sarah Wurz is Senior Researcher in the Institute for Human conferring evolutionary advantage to the groups of Homo ∼ Evolution of the University of the Witwatersrand (Private Bag 3, PO sapiens who migrated out of Africa after 60,000 years ago WITS, 2050, Johannesburg, South Africa [[email protected]]). (Mellars 2006; Mourre, Villa, and Henshilwood 2010). After This paper was submitted 3 VII 13, accepted 18 VII 13, and ∼58,000 years there is a return to Middle Paleolithic–like electronically published 29 X 13. technological variability and a dearth of complex stone tools.

᭧ 2013 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved. 0011-3204/2013/54S8-0011$10.00. DOI: 10.1086/673283 S306 Current Anthropology Volume 54, Supplement 8, December 2013

Figure 1. Map of sites mentioned in the text.

The lack of perceived complexity and sophistication of the period, from 500,000 to 125,000 years ago, mainly in open- post–Howiesons Poort technologies indicates a return to air sites all over South Africa. The Fauresmith industry, con- “nonmodern” levels of behavior and cognition (but see Lom- sidered transitional between the Early Stone Age and the MSA, bard and Parsons 2010; Marean 2010; Wurz 2008). This nar- contains small handaxes, blades, and points. Its temporal des- rative exaggerates, but it does capture the dominant percep- ignation is unclear, and although relatively young dates tion of the development of technological complexity in the (around 150,000 years ago) at, for example, Bundu Farm and MSA of South Africa. A historical perspective on the spa- Rooidam (Herries 2011) are noted, dates of between 280,000 tiotemporal variability of stone tool assemblages dating to and 500,000 years ago may be more realistic (Beaumont and between ca. 300,000 and 40,000 years ago provides the op- Vogel 2006; Herries 2011; Porat et al. 2010). The Sangoan portunity to evaluate whether this narrative is the best-fit from the northern parts of South Africa, with its core axes model for the MSA of South Africa. The MISs are used as an and MSA debitage, is as yet undated but is in all probability organizational framework only as it is clear that terrestrial also a regionally distinct transitional industry (Kuman 2007). temperature and precipitation changes do not coincide with At Duinefontein in the Western Cape (fig. 1), large cutting them (Blome et al. 2012). tools occur as late as 125,000 years ago (Feathers 2002; Klein et al. 1999). The chronological ambiguities often associated Early Middle Stone Age with Middle Pleistocene sites (cf. Barham 2012) impede a clear understanding of when handaxes and other core tools The transition from the Early Stone Age to the MSA is am- ceased being part of the technological repertoire of Pleistocene biguous, and handaxes and other large cutting tools co-occur hunter-gatherers. The earliest MSA assemblage recorded with- with typical MSA flakes, blades, and points for an extended out Acheulean elements is an a ca. 280,000-year-old assem- Wurz Technological Trends in the Middle Stone Age of South Africa S307 blage from Florisbad (Gru¨n et al. 1996; Kuman, Inbar, and prehensively that the museum collections analyzed originate Clark 1999). It is not particularly diagnostic, and platform from the recently dated “stratum 4a” at Kathu Pan (Wadley cores and a small number of triangular flakes and blades are 2013). described. There has been very little opportunity to integrate paleoen- The Early MSA dates to between 300,000 and 130,000 years vironmental and technological data of the Early MSA because ago (Lombard et al. 2012; Volman 1984) and is characterized regional environmental proxies and technological trajectories by prepared-core technologies, including Levallois, discoid, still need to be developed. An exception is the “Cape Floral and blade methods. These methods had been invented more Region—South Coast Model” for the origins of modern hu- than half a million years ago. A recurring issue is whether mans (Marean 2010:432, 2011). In this model MIS 6, one of the origin of the Levallois in South Africa is in the Victoria the coldest and perhaps most arid stages of the Quaternary West industry (Van Riet Lowe 1945). In the Victoria West (Marean 2010; Petit et al. 1999), is related to a genetic bot- industry, a Levallois-like technique is employed to prepare a tleneck (e.g., Behar et al. 2008; Blum and Jakobsson 2011; domed surface for the removal of a large preferential flake, Fagundes et al. 2007; Gonder et al. 2007) when populations resulting in different shapes of discarded cores, including the crashed and survived only in favorable refugia. The 162,000- famous hoenderbek (hen beak) cores (Lycett 2011; McNabb year-old (Jacobs 2010) assemblage from Cave 13B at Pinnacle and Beaumont 2012). Subsequent investigations of these cores Point on the southern Cape coast is considered such a re- describe them as “proto-Levallois” (White, Ashton, and Scott fugium (Compton 2011; Marean 2010). The southern Cape 2011) or “para-Levallois” to avoid assumptions of phyloge- coast, being somewhat buffered from extreme aridity effects, netic relationships (Lycett 2009). At Canteen Kopje, Victoria would have provided predictable faunal, geophyte, and shell- West technology dating to more than 500,000 years ago occurs fish food resources. The continued occupation of sites such alongside prepared-core or Levallois technology (McNabb as Klasies River and Blombos Cave in MIS 5 on the southern and Beaumont 2011, 2012). Thus, a unilinear evolution from Cape coast is seen as a fluorescence of populations that sur- the Acheulean to Levallois technology is highly unlikely. The vived in the southern Cape refugium during MIS 6. The Cave early presence of Levallois at this site confirms that it has 13B assemblage is associated with the earliest use of shellfish deep roots in South Africa and co-occurs with typical Early (Marean et al. 2007) and with Levallois technology, points, Stone Age methods. In the South African MSA, “Levallois” blades, and bladelets (!10 mm in width; Marean et al. 2007: encompasses several types of prepared cores, including radial 906). The bladelets have been reported as the first of their cores (Volman 1984; Wilkins, Pollarolo, and Kuman 2010). kind in the South African MSA (Thompson, Williams, and Whether these cores conform strictly to the “Levallois” con- Minichillo 2010; but see Wilkins and Chazan 2012). cept or not is less important than appreciating that prepared- Certain genetic, paleoenvironmental, and archaeological core technology provided a new level of technological flexi- data imply that the southern Cape MIS 6 refugium scenario bility (White, Ashton, and Scott 2011). may need reconsideration. African resequencing data (Sjo¨din In South Africa, as in East Africa (Tryon and Faith 2013) et al. 2012) indicate, for example, that a demographic model and the Levant (Hovers and Belfer-Cohen 2013), laminar without an ancestral bottleneck in MIS 6 is a possibility (see technology occurs in the earlier part of the Middle Pleistocene. also Lombard, Schlebusch, and Soodyall, forthcoming). It is Large blades together with small Fauresmith handaxes, Le- further questionable whether the MIS 6 glacial resulted in vallois points and flakes are associated with sites from the extreme aridity in all areas of southern Africa. At Florisbad, Northern Cape (Beaumont and Morris 1990; Sampson 1974). an inland site, occupation events are related to wetter phases For one of these assemblages—Kathu Pan 1 stratum 4a, re- (Scott and Brink 1992), and one of these occurs in MIS 6, at .(years ago (Kuman, Inbar, and Clark 1999 21,000 ע cently redated to ∼500,000 years ago (Porat et al. 2010)—a 157,000 systematic blade production system is described (Wilkins and At Florisbad and the surrounding areas, highly productive Chazan 2012), confirming Sampson’s (1968) impression that ecosystems occurred periodically between 400,000 and large blade production has an ancient origin in South Africa. 100,000 years ago. The nine extant and six extinct species of The cores are centripetally prepared before the removal of grazers identified in various units reflect an open grassland blades with hard-hammer percussion. The blade cores are ecosystem similar to that of the Serengeti (Brink 2005; Brink “Levallois-like” (Wilkins and Chazan 2012:15), but they are and Lee-Thorpe 1992). Geological indicators, wetland species, not Levallois in concept because the flaking surfaces are much bird remains, and palynological and phytolith data signal the more convex than the lower platform surfaces, and the in- presence of pans or lakes in the central interior at these times tersection of the surfaces is not a plane. A number of bladelets (Loock and Grobler 1988; Manengold and Brink 2011; Scott (!12 mm in width) also occur in this assemblage. Some lam- and Rossouw 2005; Van Zinderen Bakker 1989; Visser and inar products are retouched into unifacial points, and these Joubert 1991). It is not surprising that this area was occupied and nonretouched triangular flakes show evidence for hafting by humans during productive times, as evidenced by the and use as spear tips (Wilkins et al. 2012). It is significant ∼160,000-year-old “Florisbad industry” (Sampson 1974), a that this level of complexity occurs at such an early date, and highly retouched industry with many side-scrapers. Elongated it would be more so if it could be demonstrated more com- flakes and a variety of core types, including prepared cores, S308 Current Anthropology Volume 54, Supplement 8, December 2013 occur (Kuman 1989). Florisbad and other inland sites such also occur (Wurz 2012). Assemblages from Florisbad, Yster- as Border Cave, Bundu Farm, Lincoln Cave, and Wonderwerk fontein 1, and Hoedjies Punt are broadly contemporaneous Cave show that hunter-gatherer groups lived beyond the with the Klasies River techno-complex. It is uncertain how southern Cape during MIS 6 (Lombard et al. 2012). the ∼120,000-year-old hornfels-dominated assemblage from Florisbad with blades, triangular flakes, bladelets, and few MIS 5 Middle Stone Age formal tools (Kuman, Inbar, and Clark 1999) compares. The Ysterfontein (YFT 1) techno-complex dating to either 1120 Only a limited number of assemblages from MIS 5 (ca. 128– ka or MIS 5c-a (Avery et al. 2008) is a flake industry in mainly 75 ka; Carto et al. 2009) have both reliable dates and recently silcrete and calcrete with a few blades. The formal tool com- published descriptions. Spatial and chronological patterning ponent consists of denticulates (Wurz 2012). The nearby remains vague, but this may have more to do with the ana- Hoedjies Punt industry of the last interglacial age resembles lytical approaches followed and intensity of research focus YFT 1 technology closely (Will et al. 2013) and that from Sea than a true absence of cultural patterning. For a considerable Harvest (Volman 1978). The three Western Cape techno-com- period of time lithic analysts were guided by a typological plexes differ from the Klasies River occurrence, and there is paradigm to describe MSA retouched artefacts, blades, con- a strong possibility that they are younger (Will et al. 2013). vergent flakes (or points), and cores in terms of typometrical The Mossel Bay from Klasies River dating to between attributes (e.g., Kaplan 1990; Singer and Wymer 1982; Thack- ∼100,000 and 80,000 years ago is characterized by a unipolar eray 1989, 2000; Thackeray and Kelly 1988; Volman 1981, recurrent Levallois reduction process for points and blades. 1984; Wadley and Harper 1989), but more technological de- The majority of the end products have large, faceted platforms scriptions are emerging (e.g., Porraz et al. 2013b; Will et al. associated with prominent bulbs of percussion and straight 2013; Wurz 2002). The MIS 5 occurrence from Klasies River profiles (Wurz 2000, 2002). A similar assemblage comes from dates to between ca. 115,000 and 80,000 years ago (Wurz Diepkloof (Porraz et al. 2013b) and Cape St. Blaize (Sampson 2002) and consists of over 10 m of deposits. It constitutes 1974; Thompson and Marean 2008). The Pinnacle Point Cave one of the largest collection of artefacts from this time range 13B assemblage is typologically but not necessarily techno- in South Africa, sourced from both older and modern ex- logically similar (Thompson, Williams, and Minichillo 2010). cavation techniques, and this provides an opportunity to in- Commonalities between the stratified sequences from Cave vestigate technological trends through time. It is subdivided of Hearths (not dated) from the north of South Africa and into the Light Brown Sand and Shell and Sand Members Klasies River have been noted (Sampson 1974; Sinclair and (Deacon and Geleijnse 1988) and two corresponding cultural McNabb 2005; Volman 1984). In Bed 4, Cave of Hearths, units, the MSA I and the MSA II (Singer and Wymer 1982), long flake-blades and convergent points occur, while the over- or MSA 2a and 2b (Volman 1981). The morphometric sim- lying levels comprise a Levallois industry with blades, radial ilarity of the MSA I and II quartzite-dominated assemblages flakes, and, especially noticeable, convergent points. The Cave at Klasies River has been interpreted as evidence for tech- of Hearth assemblage served as a comparative basis for the nological continuity (Singer and Wymer 1982; Thackeray many surface occurrences found in the Free State (Sampson 1989), but end product profile, platform details, debitage 1968). The South African interior contains a multitude of types, core geometry, and scar patterning (after Chazan 1997; surface sites (Beaumont and Morris 1990; Sampson 1968, Van Peer 1992) indicate observable technological change 1974). For example, in an area of 216 square miles in the (Wurz 2000, 2002). Multivariate statistical analyses support middle reaches of the floodplains of the Orange River, 290 this contention and reveal significant differences between the MSA well-preserved occurrences, though undated, are iden- end products of the two taxons, especially in terms of platform tified (Sampson 1968). Sampson comments on their similarity thickness and length (Wurz et al. 2003). For this reason dif- to the pre–Howiesons Poort Klasies River and Cave of Hearths ferent terminology for the MSA I and MSA II at Klasies River assemblages. One important difference, however, is a much is suggested. “Klasies River” is the name given for the MSA larger extent of bifacial and unifacial flaking at the Cave of I because it is the first more detailed description for this type Hearths, Orange River, and also Border Cave assemblages in of occurrence (Wurz 2000), and “Mossel Bay” is the name what has been termed the “Pietersburg” Industry (Mason given for the MSA II because this name has historical pre- 1962; Sampson 1974). It would be interesting to discover how cedence for assemblages from the southern Cape that fit this these bifacials and unifacials compare with those of other time description (Sampson 1974). periods and areas. This will provide the context for under- The Klasies River techno-complex is characterized in part standing technological innovations involving invasive uni- by a recurrent blade reduction strategy. Some of the end prod- and bifacial retouch. The blades and points are probably mul- ucts have the same characteristics as those of the Howiesons tifunctional tools. Some triangular pointed flakes and blades Poort—curved blades and points with diffused bulbs of per- have damage consistent with use in a longitudinal cutting cussion and platforms showing rubbing and removal of small action (Brink and Henderson 2001; Henderson 2001; Kuman flakes from the dorsal edge (Wurz 2000, 2002). Blade and 1989). The unretouched MIS 5 points from Cave 13B at Pin- point production similar to that of the MSA II (Mossel Bay) nacle Point (Bird, Minichillo, and Marean 2007; Schoville Wurz Technological Trends in the Middle Stone Age of South Africa S309

2010) were used in a similar way. It is likely that both the by Villa et al. (2009) on the largest available collection (352 retouched and unretouched points of this time period served bifacial elements) in South Africa, from Blombos Cave (Hen- as spear tips (Lombard and Phillipson 2010; McBrearty and shilwood 2012). Direct internal percussion (with a hard ham- Brooks 2000; Wilkins et al. 2012). The populations of MIS 5 mer) and shaping by marginal percussion (with a soft or soft were efficient hunters (see Clark and Kandel 2013) and gath- stone hammer) were used in the production process of sil- erers, and integrating technological and foraging behavior crete, quartzite, and quartz bifacials. Initially, it was felt that more extensively is a priority for future research (d’Errico and pressure flaking and heat treatment (Brown 2011; Brown et Banks 2013). al. 2009) were not part of the bifacial production process, but subsequent experimentation (Mourre, Villa, and Henshil- wood 2010) finds evidence for both pressure flaking and heat The Still Bay and Howiesons Poort treatment on 4% of the silcrete bifacial points from Blombos Still Bay. The Still Bay techno-complex has fascinated re- Cave. Mourre replicated bifacial points on heat treated silcrete searchers and collectors since the first discovery of bifacial using a bone tool for pressure flaking. New diagnostic traits lanceolate points, the type artefact of the Still Bay, on the for this technique are described, and heat treatment is re- Cape Flats in 1870 (Henshilwood 2012; Minichillo 2005). A garded as a prerequisite for pressure flaking on silcrete. number of Still Bay sites occur in stratified contexts—for Mourre, Villa, and Henshilwood (2010) regard pressure flak- example, Dale Rose Parlour (Trappieskop), Peers Cave, Diep- ing used for the Still Bay bifacials as the first occurrence of kloof, Hollow Rock Shelter, Blombos Cave in the Western its kind anywhere in the world, thus an invention or inno- Cape, Sibudu Cave and Umhlatuzana in KwaZulu Natal, and vation. Intentionally heated silcrete artefacts also occur in the Apollo 11 in Namibia—in addition to several surface occur- Diepkloof SB (Porraz et al. 2013b; Schmidt et al. 2013). Some rences (Minichillo 2005; Steele et al. 2012; Wadley 2007). Most bifacials at Hollow Rock Shelter appear to have been pressure of the current dating evidence suggests that the Still Bay spans flaked with a fine-tipped small tool without heat treatment the end of MIS 5 and the beginning of MIS 4 (72–60 ka; (Ho¨gberg and Larsson 2011), a technique purportedly dif- Blome et al. 2012). There are relatively few dates, but optically ferent from the more advanced pressure-flaking techniques stimulated luminescence (OSL) dating assays from Blombos found in Europe and North America. The Diepkloof Still Bay Cave, Sibudu, Apollo 11, and Diepkloof indicate a duration bifacial elements (n p 74 ) are not pressure flaked (Porraz et for the Still Bay period of around 7,700 years, from 75,500 al. 2013b), and this is also true for the bifacials from Sibudu to 67,800 years ago (Henshilwood 2012; Jacobs et al. 2008, (Soriano et al. 2007; Wadley 2013), a feature that is partly 2012), while Ho¨gberg and Larsson report preliminary OSL related to the raw materials used. dates of ca. 72,000 and 80,000 BP. Thermoluminescence dates The majority of bifacial points in the Still Bay are in local place the Still Bay industry from Diepkloof much earlier, with or near-local materials. At Blombos Cave, for example, 72% a mean age of 109,000 years ago (Tribolo et al. 2009, 2013). of the bifacials are in silcrete, which probably originated 20– This implies that the Still Bay may have a much longer du- 30 km from the site. At Diepkloof most bifacials are in local ration in South Africa than previously thought (Porraz et al. quartzite (80%), and at Hollow Rock Shelter and Sibudu Cave 2013a, 2013b; Tribolo et al. 2013). about half are in local quartzite and dolerite, respectively. It The Still Bay is characterized by bifacially retouched foliate is likely that the bifacials are multifunctional tools (Henshil- points with lenticular cross-sections. Their shapes vary from wood 2012)—residue patterns on complete bifacials suggest narrowly elliptic to lanceolate, and they either have a wide- that they functioned as knives as well as projectile points angled pointed butt or two pointed apices (Henshilwood (Lombard 2006; Wadley 2007). One proposition is that they 2012; Minichillo 2005; Villa et al. 2009; Wadley 2007). The were used exclusively as parts of hand-delivered spears be- shape of Still Bay points may be unique in the MSA (cf. Porraz cause their morphometric attributes and macrofractures are et al. 2013b). An intersite comparative study involving bifacial similar to Paleo-Indian points (Villa and Soriano 2010; Villa points from Dale Rose Parlour, Hollow Rock Shelter, Peers et al. 2009). Another is that they may have functioned as darts Cave, Blombos Sands, Kleinjongensfontein, and Cape Hang- (Shea 2009). Because the bifacials from Diepkloof display rare klip (fig. 1) demonstrates that they occur in a wide variety impact-like fractures and frequent resharpening of the lateral of sizes: they vary from 34 mm to more than 120 mm in edges, they may have primarily functioned as cutting imple- length (Minichillo 2005). The technology of blank production ments (see also Minichillo 2005). in the Still Bay is relatively unknown and consists of three The onset of the Still Bay is correlated with climatic changes rather short production sequences—a unifacial flake reduc- that occurred at the end of MIS 5 (Thackeray 2009; Ziegler tion sequence and two versions of a bifacial block chaıˆne et al. 2013). This may have pressured populations to adopt ope´ratoire—at Hollow Rock shelter (Ho¨gberg and Larsson bifacial reduction as a strategy to economize lithic raw ma- 2011). At Diepkloof a reduction sequence for the production terial and maintain tools more intensively in the context of of laminar blanks and flakes is present but awaits further long-distance residential moves, a pattern inferred for bifacial description (Porraz et al. 2013b). industries from other parts of the world as well (McCall and An extensive attribute analysis of bifacial points is presented Thomas 2012). A period of severe climatic instability did S310 Current Anthropology Volume 54, Supplement 8, December 2013 occur ∼72,000 years ago as indicated by, for example, the Harper 1997; Kaplan 1990; Minichillo 2005; Sampson 1974; Crevice Cave carbon isotopic record (Bar-Matthews et al. Singer and Wymer 1982; Thackeray 1992; Volman 1984; Wurz 2010), the EPICA EDML curve (EPICA Community Members 1999). The blades originate from a recurrent blade production 2006), and core CD154-17-17K (Ziegler et al. 2013). For Bar- system using a soft hammer, with preparation of the lateral Matthews et al. (2010) and Ziegler et al. (2013), the Still Bay and distal convexities by elongated de´bordantes or sometimes signals a technological innovation in response to this punc- flat centripetal removals. The small blades—with high-angled, tuated environmental event, a scenario that becomes unlikely sometimes extremely small, off-center platforms with rubbed in the face of a longer duration of the Still Bay. dorsal edges—are typical of the HP (Wurz 2000). Recent de- scriptions from a strict French chaıˆne ope´ratoire perspective Howiesons Poort. The Howiesons Poort (HP) is the most (Porraz et al. 2013b; Soriano, Villa, and Wadley 2007; Villa widespread MSA industry recorded in southern Africa and is et al. 2010) provide more details on the blade reduction strat- known from more than 20 sites south of the Zambezi (Hen- egy followed at Klasies River, Rose Cottage Cave, and Diep- shilwood 2012; Lombard 2005, 2009), and new occurrences kloof. It is described as non-Levallois (Porraz et al. 2013b; were reported recently (e.g., Kandel and Conard 2012; Steele Villa et al. 2010) because the intersection of the debitage et al. 2012). Thanks to the distinctiveness of the geometric- surface with the platform and nonproduction surface is not backed artefacts, the industry has served as horizon marker a plane, and convexities are more pronounced at the begin- for the MSA of South Africa (Deacon 1989, 1992) for decades. ning of the reduction. Technological continuity may exist be- Howiesons Poort sites occur at, for example, the HP name tween this system and the strategies followed in Middle Pleis- site, Boomplaas, Border Cave, Diepkloof, Klasies River, Klein tocene Kathu Pan and the 115,000-year-old Klasies River Kliphuis, Rose Cottage Cave, Sibudu, and Um- (MSA I) industries. In both cases the cores are described as hlatuzana in South Africa; at Melikane and Ntloana Tsoana “Levallois-like,” with very convex upper surfaces (Wilkins and in Lesotho; and at Apollo 11 in Namibia and a number of Chazan 2012; Wurz 2002, 2010). The backed artefacts are surface sites. The HP is sometimes considered to be a relatively mostly on blade blanks, but at some sites, such as Diepkloof, short-lived entity occurring within well-understood time lim- they may be on flakes (Mackay 2008b; Porraz et al. 2013b). its because eight sites have single-grain OSL dates of between Experimental knapping by S. Soriano and G. Porraz indicate ca. 64,800 and 59,500 years ago (Cochrane, Doelman, and marginal percussive gestures with a soft stone hammer (Porraz Wadley 2013; Jacobs et al. 2008). et al. 2008; Soriano, Villa, and Wadley 2007; Villa et al. 2010). There are a number of younger and older dates for the HP. It is not clear whether local raw materials were used, but this At Klasies River, for example, the HP is associated with elec- is essential in replicative experiments. Villa et al. (2010) sug- tron spin resonance and thermoluminescence dates of be- gest that the HP is the first known occurrence of small blades tween 50,000 and 60,000 years ago (Eggins et al. 2005; Feathers produced by marginal percussion and used as blanks for the 2002; Tribolo, Mercier, and Valladas 2005). The HP from manufacture of backed pieces. A noteworthy departure from Border Cave dates to around 75,000 years ago (Gru¨n and the chaıˆne ope´ratoire–like methodologies to investigate HP Beaumont 2001; Gru¨n, Beaumont, and Stringer 1990), and variability is the materialist approach in which categories are this is somewhat similar to dates from Pinnacle Point and constructed without notions of “ideal forms, the mental pre- Diepkloof. Unit SADBS at Pinnacle Point 5–6 (Brown 2011; disposition of the maker, or the presumed goals of the artisan” Brown et al. 2012) contains small blades, notched pieces, and (Hiscock 2007:201). From this perspective the HP from Diep- backed artefacts as well as quartzite blades and points. This kloof and Klein Kliphuis is interpreted as a time period in industry is interpreted as transitional to the HP (Brown et which core mass was transformed into flake length more ef- al. 2012), but in most respects it falls within the techno- ficiently than in preceding or subsequent technological sys- typological range of variability of the HP (see also Porraz et tems (Mackay 2008a, 2009). The ratio of edge length to mass al. 2013b). The Diepkloof TL dates are between ∼109,000 and of complete flakes is used as a proxy for flaking efficiency. ∼52,000 years ago (Porraz et al. 2013b; Tribolo et al. 2013). The HP is not a static entity, and changes through time in It is suggested that the single-grain OSL dates for the early the proportions of backed tools and notched artefacts occur HP (previously interpreted as Still Bay) are erroneous because (e.g., Harper 1994; Mackay 2011; Porraz et al. 2013b; Singer of methodological problems (Gue´rin et al. 2013; Tribolo et and Wymer 1982; Soriano, Villa, and Wadley 2007; Volman al. 2013). Whether the TL dating from Diepkloof demon- 1984; Wadley and Mohapi 2008). Bifacially flaked pieces do strates that the HP is not a horizon marker and is of much occur in some HP assemblages (e.g., Deacon 1995; Wurz 2000: longer duration than previously thought (Porraz et al. 2013a, 91), and their presence does not necessarily indicate an early 2013b; Tribolo et al. 2013) is still open to debate. phase of the HP (Porraz et al. 2013a, 2013b). Changes in raw Despite the increased research focus on the HP, its primary material proportions, especially quartz and silcrete, are also elements remained unchanged. It is a small-blade industry noted (Mackay 2011; Porraz et al. 2013b; Wurz 2000). with backed and notched artefacts and lesser proportions of The HP consists of different phases at sites such as Um- pieces esquille´es, scrapers, unifacial points, and partly bifacial hlatuzana, Rose Cottage Cave, Sibudu Cave, Diepkloof, and points (Deacon 1995; Goodwin and Van Riet Lowe 1929; Klasies River (Porraz et al. 2013b; Soriano, Villa, and Wadley Wurz Technological Trends in the Middle Stone Age of South Africa S311

2007; Villa et al. 2010; Wadley and Mohapi 2008). At Klasies chaeological sites (e.g., Klein 1976; Van Pletzen 2000). A num- River, for example, a flake production strategy occurs in the ber of proxies (e.g., EPICA Community Members 2004; North middle of the sequence continuing into the upper levels, and Greenland Ice Core Project Members 2004; Pahnke et al. 2003; this is associated with changes in the blade parameters and Ziegler et al. 2013) indicate the colder conditions of MIS 4. percussion technique. Despite these more subtle changes and The speleothem carbon and oxygen isotope ratios from Crev- morphometric differences in the cores (Clarkson 2010), the ice Cave, close to Pinnacle Point, are interpreted as evidence HP technological signal is similar at sites as far apart as Rose for more summer rain and more grass cover in this time Cottage Cave, Klasies River, and Border Cave (Villa et al. period (Bar-Matthews et al. 2010). The grassier habitats in 2010). The perception that the selection of fine-grained “high the southern Cape in MIS 4 may be linked to wetter con- quality, exotic” (Singer and Wymer 1982) raw material oc- ditions due to the shift of the position of the winter rainfall curred in HP assemblages (e.g., McCall and Thomas 2012) belt during glacial advances (Chase and Meadows 2007). Mi- needs to be qualified. There is in fact considerable variability crofaunal evidence (e.g., Avery 1982, 1992) from the southern in the raw material composition of different sites, and the Cape points to colder and moister conditions during this time sources are often local (Lombard 2005; Minichillo 2006; Wad- period. Similar conditions occurred at Sibudu Cave, where ley 2008; Wadley and Mohapi 2008; Wurz 2000). What is the fauna (Clark 2011; Clark and Plug 2008) and charcoal more relevant is that fine-grained rocks were preferentially (Allott 2006; Hall, Woodborne, and Scholes 2008) indicate a selected. cool, moist environment with evergreen forest but with a Experimental, use-wear, and trace analyses of the HP woodland/savannah habitat close by. At Sibudu and a few backed artefacts indicate that they were inserts in composite other HP sites, a marked increase in the exploitation of small projectile weapons and that hafting arrangements and the raw mammals and the smallest ungulates in MIS 4 occur, indi- material of choice for hafts changed through time (Lombard cating closed environments and an expansion of diet breadth 2008; Wadley and Mohapi 2008). A complex chain of actions (Clark and Kandel 2013). It could be expected that a mosaic was followed to create compound adhesives, with red ochre of open and closed environments occurred in MIS 4. and plant gum among the ingredients for hafting of the The HP has been interpreted as an adaptation to fluctuating backed artefacts (Wadley 2013; Wadley, Hodgkiss, and Grant MIS 4 environments in the context of residential mobility and 2009). The impact fractures on backed artefacts imply that risk reduction through investment in technology (e.g., Am- they functioned as spear points (Villa and Soriano 2010). brose 2002; McCall 2007) and time-dependent foraging (Min- Small quartz segments from Sibudu Cave may represent the ichillo 2006). Relatively more time and energy had been ex- first bow-and-arrow technology, a crucial invention (Hen- pended to produce the geometric-backed artefacts for shilwood 2012; Lombard 2011; Lombard and Phillipson composite tools (Bird and O’Connell 2006; Elston and Bran- 2010). Several lines of evidence suggest that the quartz seg- tingham 2002). Another interpretation is that the HP and Still ments tipped arrows in a transverse position: their morpho- Bay reflect a response to the expansion and isolation of mod- metric dimensions (Wadley and Mohapi 2008), experiments ern human populations between 80,000 and 60,000 years ago by Lombard and Pargeter (2008), impact fractures (Lombard (Jacobs and Roberts 2009), as may be implied by some genetic 2011), microresidues (Lombard 2007, 2008), and use-trace studies (Atkinson, Gray, and Drummond 2009; Behar et al. evidence (Lombard 2011). It is likely that different types of 2008; Quintana-Murci et. al. 2008; Scheinfeldt, Sameer, and backed artefacts were used in different kinds of hunting ac- Tishkoff 2010; Tishkoff et al. 2007; see also Lombard, Schle- tivities. They were used to perform a variety of functions, busch, and Soodyall, forthcoming). The interplay between including cutting (Igreja and Porraz 2013; McBrearty and climate, demography, mobility, and technological responses Brooks 2000; Wadley and Mohapi 2008). A bone point from is a complex topic (d’Errico and Banks 2013) only touched the Sibudu HP layers at Sibudu Cave, similar to unpoisoned on here because the focus is on technological change in the bone arrow points from much later contexts, may also have MSA (for recent reviews see Brown 2011; McCall and Thomas been used in bow-and-arrow technology (Backwell, d’Errico, 2012). To investigate more productively whether climate has and Wadley 2008; d’Errico, Backwell, and Wadley 2012). It been a motor of behavioral change in the HP, less coarse tools seems that bow-and-arrow technology disappeared by the end need to be developed to integrate climatic data with cultural of the HP (Wadley and Mohapi 2008). change (d’Errico and Henshilwood 2007). The relationship between climatic change and HP cultural expressions is a much-discussed topic (Cochrane, Doelman, After the Howiesons Poort and Wadley 2013). The colder conditions during this glacial period may be reflected by the angular spall in HP levels at In his 2008 review of MIS 3 MSA, Mitchell (2008) notes that Border Cave, Rose Cottage Cave, and Boomplaas Cave (as this period is relatively unexplored in comparison with MIS reviewed in Chase 2010). The colder conditions are related 4, and this remains true. The transition to the post-HP to more open environments (Bar-Matthews et al. 2010; Dea- ∼58,000 years ago preceded a large-scale change in climate con and Lancaster 1988; Deacon et al. 1984), an inference ∼57,000 years ago (Cochrane 2008). Occupational hiatuses supported by the presence of grazers in southern Cape ar- occur in some southern Cape sites in MIS 3 (∼50–20 ka), S312 Current Anthropology Volume 54, Supplement 8, December 2013 perhaps due to hyperarid conditions (Deacon and Thackeray South Africa. Organic artefacts—such as bone arrowheads, 1984; Steele and Klein 2009), but otherwise MIS 3 was marked notched bones for notational purposes, and digging sticks and by fluctuating climatic conditions that included very wet pe- bone awls similar to those used by ethnographically known riods (Deacon and Lancaster 1988; Mitchell 2008). Proxies San—are identified (d’Errico et al. 2012) and interpreted as from Tswaing Crater, Wonderkrater, Elands Bay, and the Cape the emergence of adaptations that are typical of the San. These Flats show that rainfall was sometimes less but still compa- new results do not clarify the considerable ambiguity sur- rable to levels experienced by the South African Highveld rounding the MSA-LSA transition appreciably as the contin- today. The occurrence of more than 25 MSA sites dating to ued presence of typical MSA stone tool assemblages until this time period in southern Africa (Mitchell 2008) is testi- 22,000 years ago (Deacon and Deacon 1999) still prevents mony to successful adaptation in the post-HP time period. understanding the technological difference between the MSA MIS 3 assemblages are associated with a number of labels, and LSA in terms of a clear-cut Rubicon (Clark 1999; Mitchell including post-HP, MSA 3, MSA III, or informal late MSA 2012; Wadley 1993). (Lombard et al. 2012). In most assemblages there is a return to the larger blades and points that characterize the MSA, but continuities between the HP and post-HP at Klasies River and Concluding Discussion Rose Cottage Cave do occur (Clark 1999; Lombard and Par- In South Africa, as in other areas of Africa, the transition sons 2010; Soriano, Villa, and Wadley 2007; Villa et al. 2010; between the ESA and MSA is not well understood. The tran- Wadley and Harper 1989; Wurz 2000). Technological trends sition was between ∼500,000 and 285,000 years ago, and dur- are much less apparent than in MIS 4, even though the same ing this time period MSA and Acheulean elements occur to- variety of analytical approaches has been applied for the post- gether. The technological antecedents of the MSA are in HP. Many sites dating to between ∼58,000 and 45,000 years prepared-core technology associated with Acheulean assem- ago contain points, often unifacially retouched, for example, blages and Fauresmith blade and point technologies. In Early at Border Cave, Klein Kliphuis, Sibudu Cave, Diepkloof, Kla- MSA assemblages (300,000–130,000 years ago), blade tech- sies River, Umhlatuzana, and Rose Cottage Cave in South nology, Levallois flake and point technologies, bladelets, and Africa and Melikane, Ntloana Tsoana, and Sehonghong in retouched types such as scrapers, denticulates, and retouched Lesotho (Lombard et al. 2012). The most extensive analysis points form part of the stone tool inventory. In MIS 5 the of the post-HP is from Sibudu Cave. The technology is not same diversity of technical elements continues. There is much very elaborate, with a minimal degree of predetermination of potential in further investigating the spatiochronological ex- shapes (Villa, Delagnes, and Wadley 2005), and the retouched tent and details of the technological trends documented by component, mainly consisting of unifacial points, is well de- earlier syntheses (e.g., Sampson 1974; Volman 1984). Cur- scribed (Cochrane 2006; Conard, Porraz, and Wadley 2012; Mohapi 2012; Villa, Delagnes, and Wadley 2005; Wadley rently it is only in MIS 4 that the technological trends are 2005). These points exhibit unique attributes: they have fac- clear, but this may simply be the result of the more intensive eted platforms and are relatively broader, thicker, and longer analytical scrutiny of this period. Components of the Still Bay than the other point types in the Sibudu sequence (Mohapi bifacials and their production processes are thoroughly de- 2012). It is suggested that post-HP assemblages in South Af- scribed, but the debitage processes await further consider- rica dating to between 58,000 and 45,000 years ago be termed ation. The Howiesons Poort techno-complex is described to the “Sibudu” (Lombard et al. 2012). It has already stimulated a fuller extent than that for the Still Bay. It encompasses a debate on whether this label is appropriate for all post-HP characteristic blade and bladelet associated with different pro- sites in this time range (e.g., Conard, Porraz, and Wadley portions of flakes, backed geometric shapes, notched pieces, 2012; Porraz et al. 2013b). and other retouched classes. The Still Bay and Howiesons Final MSA assemblages, dating to between 45,000 and Poort are widely considered as distinct technological pulses 20,000 years ago (Lombard et al. 2012), are associated with of a relatively short duration each, but the ambiguity in dating a wide variety of point types, such as hollow-based points results may mean that this idea needs revisiting. Post–How- and bifacial and unifacial points. Beaumont (1978) suggests iesons Poort technologies occurring between 58,000 and that a ∼40 ka industry at Border Cave with small, irregular 45,000 years ago are united by the presence of unifacial points microlithic flakes in quartz—frequently produced by bipolar in what has been termed the Sibudu techno-complex. The reduction and a few retouched tools, including outils e´cail- LSA may be heralded by an ∼42,000-year-old assemblage from le´es—represents an “Early Later Stone Age” (ELSA). In sup- Border Cave with bipolar technology, small quartz flakes and port of this hypothesis, Villa et al. (2012) describe the tech- bladelets, and San-like bone tools. This assemblage is the only nology of ELSA and preceding industries and also microliths occurrence of its kind in South Africa at this time. It may be (here referring to bladelet fragments and small flakes) hafted an indication that regionally distinct technological trajectories with pitch from Podocarpus elongates bark. Like Beaumont, existed around 40,000 years ago. Hypotheses that are con- they propose that the ∼43,000-year-old assemblage at Border structed to investigate how the MSA and LSA contrast hand- Cave marks the beginning of the Later Stone Age (LSA) in icap understanding of cultural trends (Mitchell 2002) as it Wurz Technological Trends in the Middle Stone Age of South Africa S313 masks continuities and the complex mosaic of cultural choice requires identifying local antecedents. Historical factors change. structure the adaptive configuration of populations as ex- Several claims of “firsts” or innovations are associated with plained by, for example, Kuhn (2006) in the “rugged fitness the Still Bay and Howiesons Poort. These are pressure flaking landscape” model. Populations would have drawn on partic- in combination with heat treatment for the production of Still ular technical expressions that they were already familiar with. Bay silcrete bifacial points, marginal percussion with a soft When seen from this perspective, the “innovativeness” of the stone hammer and the transformation of blades into geo- Howiesons Poort and Still Bay is not revolutionary. Artefact metrics in the Howiesons Poort, and the use of small How- production systems in MIS 4 are different in detail and not iesons Poort quartz-backed artefacts as transverse arrowheads in kind from those of other MSA industries. Pre–Still Bay in what may represent the earliest known bow-and-arrow MSA populations were able to use most of the stone tool technology. Noticeably more objects—such as beads (d’Errico, techniques that flourished in MIS 4. For example, the blade Vanhaeren, and Wadley 2008; d’Errico et al. 2005; Henshil- technology of the Howiesons Poort was present in the wood et al. 2004), engraved ochre (Henshilwood and d’Errico ∼110,000-year-old Klasies River assemblage; bladelet produc- 2011; Henshilwood, d’Errico, and Watts 2009; Henshilwood tion is ancient; heat treatment of stone occurs already 162,000 et al. 2002; Mackay and Welz 2008), engraved ostrich eggshell years ago; and bifacial flaking occurred in MIS 5. Several (Texier et al. 2010, 2013), and formal bone tools (d’Errico researchers have noted that fluctuating backed artefact pro- and Henshilwood 2007; d’Errico, Moreno, and Rifkin 2012; duction characterizes the MSA from at least 230,000 years Henshilwood et al. 2001)—occur within the MIS 4 techno- ago (Barham 2002; Hiscock and O’Connor 2006; Wadley and complexes. These innovations are interpreted as evidence for Mohapi 2008). The same argument can be made for organic symbolically mediated behavior (Henshilwood 2012; Hen- artefacts. The enigmatic remains of what was possibly part of shilwood and Dubreuil 2011 [but see comments on the latter]; a 121,000-year-old throwing stick is described from Florisbad Henshilwood and Marean 2003). The innovative aspects of (Bamforth and Henderson 2003), and bone tools occur in behavior of interest in developing explanatory models of MSA the ∼100,000-year-old levels from Klasies River (Wurz 2000). behavior are listed by d’Errico and Banks (2013). Among these Climatic and paleoenvironmental reconstruction plays an are the ochre tool kits from Blombos Cave dating to ∼100,000 important role in developing explanatory theory for the MSA, years ago (Henshilwood et al. 2011) and pre–MIS 4 engraved but the relationship between climatic and technological ochres (d’Errico, Moreno, and Rifkin 2012; Henshilwood, change is indirect and multifaceted (Cochrane, Doelman, and d’Errico, and Watts 2009; Watts 2010). This places incidences Wadley 2013; Jacobs and Roberts 2009). In the MSA, as in of “symbolic behavior” (Henshilwood and Dubreuil 2011) the LSA, climate and environment structured the available and complex cognition (Wadley 2010, 2013) earlier than MIS subsistence choice but had no deterministic effect on tech- 4. Insights into the cognitive capabilities of MSA people are nology. LSA technological changes are described as filtering further explored by other frameworks (e.g., Lombard and through the social web of interactions in a complex way (Dea- Haidle 2012; Wynn and Coolidge 2011), including those that con 1984:285). Farther afield, in Europe, for example, Ne- cast the theoretical net wider by incorporating evolutionary anderthal technological changes in MIS 9–7 occur indepen- theory (Dubreuil 2011; Ellis 2011; Gontier 2012). Further dent of climatic change (Moncel et al. 2011). research on how MSA technological trends covary with trends Although this review focuses on the MSA of South Africa, in foraging (Clark and Kandel 2013) and other kinds of be- the area south of Zambezi and the Kunene forms an ecolog- havior (d’Errico and Banks 2013; Lombard 2012) is needed ical, cultural, and archaeological unit (Mitchell 2002). It is to develop integrated models of cultural development in the thus relevant to compare technological trends in the South MSA. African MSA with those of countries such as Botswana, Zim- The search for innovation drives research on the MSA, babwe, Lesotho, southern Mozambique, and Swaziland. On explicitly so during the past decade. What is considered “in- a superficial level the technological patterning of South Africa novative” technologically is often sourced from the Upper is different from what is described for these countries, al- Paleolithic from Europe (but see Wadley 2013). Combining though the same technical elements, such as retouched points approaches developed for the European Paleolithic with local and backed artefacts, occur (Barham and Mitchell 2008; ones to analyze MSA assemblages has led to more detailed McBrearty and Brooks 2000). Nevertheless, the absence of descriptions of technologies, but it is pointless to describe adequate chronometric control and the lack of a unified de- assemblages as “Middle” or “Upper Paleolithic” in character scriptive taxonomy to compare assemblages impede a thor- (e.g., Soriano, Villa, and Wadley 2007; Villa, Delagnes, and ough understanding of how the areas relate. The spatio- Wadley 2005; Villa et al. 2010). Eurocentric interpretation of chronological integrity of the South African MSA is mostly the MSA is not new, and already in 1928 Goodwin counseled insufficient to identify clear trends, and it is far from “sorted against seeing the South African Stone Age through “Euro- fully” (Goodwin and Van Riet Lowe 1929:7; Thackeray 1992: pean spectacles” (Goodwin 1928:29). Technological change in 399). There are adequate data to postulate that the MSA in the MSA is best interpreted within the context of local his- South Africa was a period characterized by fluctuating tech- torical trajectories. Inferring the origins of any technological nological “fashions” (cf. Thackeray 1989), but the chrono- S314 Current Anthropology Volume 54, Supplement 8, December 2013 logical and regional details remain largely obscure. One hy- Herries’ “A chronological perspective on the Acheulian and its transition to the Middle Stone Age in Southern Africa: the question of the Fauresmith” pothesis is that similar cycles of technological change occur (2011). International Journal of Evolutionary Biology (2012):1–5. over the subcontinent (Volman 1984; Wurz 2012). Volman Barham, Lawrence, and Peter Mitchell. 2008. The first Africans: African ar- (1984:194) notes “a similar pattern of assemblage change chaeology from the earliest tool makers to most recent foragers. Cambridge: Cambridge University Press. through time occurs in MSA sequences south of the Lim- Bar-Matthews, Miryam, Curtis W. Marean, Zenobia Jacobs, Panagiotis Kar- popo,” and Sampson (1974) suggests notable chronological kanas, Erich C. Fisher, Andy I. R. Herries, Kyle Brown, et al. 2010. A high and regional patterning for the South African MSA. This level resolution and continuous isotopic speleothem record of paleoclimate and paleoenvironment from 90 to 53 ka from Pinnacle Point on the south coast of change can be most clearly observed for the Still Bay and of South Africa. Quaternary Science Reviews 29:2131–2145. Howiesons Poort techno-complexes. The lack of recently Beaumont, Peter B. 1978. Border Cave. PhD thesis, University of Cape Town. demonstrated patterning on a similar scale for the rest of the Beaumont, Peter B., and David Morris. 1990. Guide to archaeological sites in the Northern Cape. Kimberly, South Africa: McGregor Museum. South African MSA may be an artefact of research intensity Beaumont, Peter B., and John C. Vogel. 2006. On a timescale for the past or may indicate an actual absence of subcontinental scale million years of human history in central South Africa. South African Journal patterning. Technological patterning in MIS 3, for example, of Science 102:217–228. Behar, Doron M., Richard Villems, Himla Soodyall, Jason Blue-Smith, Luisa reveals that regionally variable technological trajectories ex- Pereira, Ene Metspalu, Rosaria Scozzari, et al. 2008. The dawn of human isted at times in the prehistory of South Africa (Mitchell matrilineal diversity. American Journal of Human Genetics 82:1130–1140. 2012). Progress in this regard would be optimal if new in- Bird, Catherine, Tom Minichillo, and Curtis W. Marean. 2007. Edge damage distribution at the assemblage level on Middle Stone Age lithics: an image- vestigations identify trends and construct hypotheses in the based GIS approach. Journal of Archaeological Science 34:771–780. context of data generated by previous generations. Bird, Douglas W., and James F. O’Connell. 2006. Behavioral ecology and archaeology. Journal of Archaeological Research 14:143–188. Blome, Margaret W., Andrew S. Cohen, Christian A. Tryon, Alison S. Brooks, and Joellen Russell. 2012. The environmental context for the origins of modern human diversity: a synthesis of regional variability in African cli- Acknowledgments mate 150,000–30,000 years ago. Journal of Human Evolution 62:563–592. Blum, Michael G. B., and Mattias Jakobsson. 2011. Deep divergences of human My thanks go to Erella Hovers and Steve Kuhn for the in- gene trees and models of human origins. Molecular Biology and Evolution vitation to take part in this conference and to them and Laurie 28:889–898. Obbink and Leslie Aiello for organizing the conference. Two Brink, James S. 2005. The evolution of the black wildebeest (Connochaetes gnou) and modern large mammal faunas of central southern Africa. PhD reviewers are thanked for their helpful comments on the man- thesis, University of Stellenbosch. uscript. I also thank the African Origins Platform of the Na- Brink, James S., and Zoe¨ L. Henderson. 2001. A high-resolution last interglacial tional Research Foundation of South Africa for funding my MSA horizon at Florisbad in the context of other open-air occurrences in the central interior of southern Africa: an interim statement. In Middle research. Palaeolithic and Middle Stone Age settlement systems. Nicholas J. Conard, ed. Pp. 1–20. Tu¨bingen: Kerns. Brink, James S., and Julia Lee-Thorp. 1992. The feeding niche of an extinct References Cited springbok, Antidorcas bondi (Antilopini, Bovidae), and its palaeoenviron- Allott, Lucy. 2006. Archaeological charcoal as a window on the palaeovege- mental meaning. South African Journal of Science 88:227–229. tation and wood use during the Middle Stone Age at Sibudu Cave. South Brown, Kyle S. 2011. The sword in the stone: lithic raw material exploitation African Humanities 18:173–201. in the Middle Stone Age at Pinnacle Point Site 5–6, southern Cape, South Ambrose, Stanley H. 2002. Early microlithic industries in sub-Saharan Africa. Africa. PhD dissertation, University of Cape Town. In Thinking small: global perspectives on microlithization. Robert G. Elston Brown, Kyle S., Curtis W. Marean, Andy I. R. Herries, Zenobia Jacobs, Chantal and Steven L. Kuhn, eds. Pp. 9–29. Archeological Papers of the American Tribolo, David Braun, David L. Roberts, et al. 2009. Fire as an engineering Anthropological Association, no. 12. Arlington, VA: American Anthropo- tool of early modern humans. Science 325:859–862. logical Association. Brown, Kyle S., Curtis W. Marean, Zenobia Jacobs, Benjamin J. Schoville, Atkinson, Quentin D., Russel D. Gray, and Alexei J. Drummond. 2009. Bayes- Simen Oestmo, Erich C. Fisher, Jocelyn Bernatchez, et al. 2012. An early ian coalescent inference of major human mitochondrial DNA haplogroup and enduring advanced technology originating 71,000 years ago in South expansions in Africa. Proceedings of the Royal Society B 276:367–373. Africa. Nature 491:590–593. Avery, Graham, David Halkett, Jayson Orton, Teresa Steele, Madelon Tusenius, Carto, Shannon L., Andrew J. Weaver, Renee´ Hetherington, Yin Lam, and and Richard Klein. 2008. The Ysterfontein 1 Middle Stone Age rock shelter Edward C. Wiebe. 2009. Out of Africa and into an ice age: on the role of and the evolution of coastal foraging. South African Archaeological Society global climate change in the late Pleistocene migration of early moderns Goodwin Series 10:66–89. Out of Africa. Journal of Human Evolution 56:139–151. Avery, Margaret D. 1982. Micromammals as palaeoenvironmental indicators Chase, Brian M. 2010. South African palaeoenvironments during marine ox- and an interpretation of the late Quaternary in the southern Cape Province, ygen isotope stage 4: a context for the Howiesons Poort and Still Bay South Africa. Annals of the South African Museum 85:183–377. industries. Journal of Archaeological Science 37:1359–1366. ———. 1992. The environment of early modern humans at Border Cave, Chase, Brian M., and Michael E. Meadows. 2007. Late Quaternary dynamics South Africa: micromammalian evidence. Palaeogeography, Palaeoclimatol- of southern Africa’s winter rainfall zone. Earth Science Reviews 84:103–138. ogy, Palaeoecology 91:71–87. Chazan, Michael. 1997. Lead review: redefining Levallois. Journal of Human Backwell, Lucinda, Francesco d’Errico, and Lyn Wadley. 2008. Middle Stone Evolution 33:719–735. Age bone tools from the Howiesons Poort layers, Sibudu Cave, South Africa. Clark, Amelia M. B. 1999. Late Pleistocene technology at Rose Cottage Cave: Journal of Archaeological Science 35:1566–1580. a search for modern behavior in a Middle Stone Age context. African Bamforth, Marion K., and Zoe¨L. Henderson. 2003. A reassessment of the Archaeological Review 16:93–119. wooden fragment from Florisbad, South Africa. Journal of Archaeological Clark, Jamie L. 2011. The evolution of human culture during the later Pleis- Science 30:637–651. tocene: using fauna to test models on the emergence and nature of ‘‘mod- Barham, Lawrence S. 2002. Backed tools in the Middle Pleistocene central ern” human behavior. Journal of Anthropological Archaeology 30:273–291. Africa and their evolutionary significance. Journal of Human Evolution 43: Clark, Jamie L., and Andrew W. Kandel. 2013. The evolutionary implications 585–603. of variation in human hunting strategies and diet breadth during the Middle ———. 2012. Letter to the editor: clarifying some fundamental errors in Stone Age of southern Africa. Current Anthropology 54(suppl. 8):S269–S287. Wurz Technological Trends in the Middle Stone Age of South Africa S315

Clark, Jamie L., and Ina Plug. 2008. Animal exploitation strategies during the Dubreuil, Benoıˆt. 2011. The other middle-range theories: mapping behaviour South African Middle Stone Age: Howiesons Poort and post–Howiesons and the evolution of the mind. In Homo symbolicus: the dawn of language, Poort fauna from Sibudu Cave. Journal of Human Evolution 54:886–898. imagination and spirituality. Christopher S. Henshilwood and Francesco Clarkson, Chris. 2010. Regional diversity within the core technology of the d’Errico, eds. Pp. 185–204. Amsterdam: Benjamins. Howiesons Poort techno-complex. In New perspectives on old stones: ana- Dusseldorp, Gerrit, Marlize Lombard, and Sarah Wurz. 2013. Pleistocene lytical approaches to Palaeolithic technologies. Stephen Lycett and Parth Chau- Homo and the updated Stone Age sequence of South Africa. South African han, eds. Pp. 43–60. New York: Springer. Journal of Science 109(5/6):1–7. Cochrane, Grant W. G. 2006. An analysis of lithic artefacts from the ∼60 ka Eggins, Stephen M., Rainer Gru¨n, Malcolm T. McCulloch, Alistair W. G. Pike, layers of Sibudu Cave. Southern African Humanities 18:69–88. John Chappell, Les Kinsley, Graham Mortimer, et al. 2005. In situ U-series ———. 2008. The transition from Howiesons Poort to post–Howiesons Poort dating by laser-ablation multi-collector ICPMS: new prospects for Quater- industries in southern Africa. South African Archaeological Society Goodwin nary geochronology. Quaternary Science Reviews 24:2523–2538. Series 10:157–167. Ellis, George F. R. 2011. Biology and mechanisms related to the dawn of Cochrane, Grant W. G., Trudy Doelman, and Lyn Wadley. 2013. Another language. In Homo symbolicus: the dawn of language, imagination and spir- dating revolution for prehistoric archaeology? Journal of Archaeological ituality. Christopher S. Henshilwood and Francesco d’Errico, eds. Pp. 163– Method and Theory 20:42–60. 184. Amsterdam: Benjamins. Compton, John S. 2011. Pleistocene sea-level fluctuations and human evo- Elston, Robert G., and P. Jeffrey Brantingham. 2002. Microlithic technology lution on the southern coastal plain of South Africa. Quaternary Science in northern Asia: a risk-minimizing strategy of the Late Paleolithic and Reviews 30:506–527. Early Holocene. In Thinking small: global perspectives on microlithization. Conard, Nicholas J., Guillaume Porraz, and Lyn Wadley. 2012. What is in a Robert G. Elston and Steven L. Kuhn, eds. Pp. 104–117. Washington: Amer- name? characterizing the “post–Howieson’s Poort” at Sibudu. South African ican Anthropological Association. Archaeological Bulletin 67 (196):180–199. EPICA Community Members. 2004. Eight glacial cycles from an Antarctic ice Deacon, Hilary J. 1989. Late Pleistocene palaeoecology and archaeology in core. Nature 429:623–628. the southern Cape, South Africa. In The human revolution: behavioural and ———. 2006. One-to-one coupling of glacial climate variability in Greenland biological perspectives on the origins of modern humans. Paul Mellars and and Antarctica. Nature 444:195–198. Chris Stringer, eds. Pp. 547–564. Edinburgh: Edinburgh University Press. Fagundes, Nelson J. R., Nicolas Ray, Mark Beaumont, Samuel Neuenschwan- ———. 1992. Southern Africa and modern human origins. Transactions of der, Francisco M. Salzano, Sandro L. Bonatto, and Laurent Excoffier. 2007. the Royal Society B 337:177–183. Statistical evaluation of alternative models of human evolution. Proceedings Deacon, Hilary J., and Janette Deacon. 1999. Human beginnings in South of the National Academy of Sciences of the USA 104:17614–17619. Africa: uncovering the secrets of the Stone Age. Cape Town: David Philip. Feathers, James K. 2002. Luminescence dating in less than ideal conditions: Deacon, Hilary J., Janette Deacon, Anton Scholtz, J. Francis Thackeray, James case studies from Klasies River main site and Duinefontein, South Africa. S. Brink, and John C. Vogel. 1984. Correlation of palaeoenvironmental data Journal of Archaeological Science 29(2):177–194. from the Late Pleistocene and Holocene deposits at Boomplaas Cave, south- Gonder, Mary K., Holly M. Mortensen, Floyd A. Reed, Alexandra de Sousa, ern Cape. In Late Cenozoic palaeoclimates of the Southern Hemisphere. John and Sarah A. Tishkoff. 2007. Whole-mtDNA genome sequence analysis of C. Vogel, ed. Pp. 339–352. Rotterdam: Balkema. ancient African lineages. Molecular Biology and Evolution 24:757–768. Deacon, Hilary J., and Vera B. Geleijnse. 1988. The stratigraphy and sedi- Gontier, Nathalie. 2012. Selectionist approaches in evolutionary linguistics: mentology of the main site sequence, Klasies River, South Africa. South an epistemological analysis. International Studies in the Philosophy of Science African Archaeological Bulletin 43:5–14. 26(1):67–95. Deacon, Hilary J., and Francis Thackeray. 1984. Late Pleistocene environ- Goodwin, A. John H. 1928. An introduction to the Middle Stone Age of South mental changes and implications for the archaeological record in southern Africa. South African Journal of Science 25:410–418. Africa. In Late Cenozoic palaeoclimates of the Southern Hemisphere.JohnC. Goodwin, A. John H., and Clarence Van Riet Lowe. 1929. The Stone Age Vogel, ed. Pp. 375–390. Rotterdam: Balkema. cultures of South Africa. Annals of the South African Museum 27:9–51. Deacon, Janette. 1984. The Later Stone Age of southernmost Africa. Cambridge Gru¨n, Rainer, and Peter B. Beaumont. 2001. Border Cave revisited: a revised Monographs in African Archaeology 12, BAR International Series 213. Ox- ESR chronology. Journal of Human Evolution 40:467–482. ford: BAR. Gru¨n, Rainer, Peter B. Beaumont, and Chris B. Stringer. 1990. Electron spin ———. 1995. An unsolved mystery at the Howiesons Poort name site. South resonance dating evidence for early modern humans at Border Cave in African Archaeological Bulletin 50:110–120. South Africa. Nature 344(6266):537–539. Deacon, Janette, and Nick Lancaster. 1988. Late Quaternary Palaeoenviron- Gru¨ n, Rainer, James S. Brink, Nigel A. Spooner, Lois Taylor, Chris B. Stringer, ments of southern Africa. Oxford: Clarendon. Robert G. Franciscus, and Andrew S. Murray. 1996. Direct dating of Flor- d’Errico, Francesco, Lucinda Backwell, Paola Villa, Ilaria Degano, Jeannette isbad hominid. Nature 382:500–501. J. Lucejko, Marion K. Bamford, Thomas F. G. Higham, Maria Perla Col- Gue´rin, Guillaume, Andrew S. Murray, Mayank Jain, Kristina J. Thomsen, ombini, and Peter B. Beaumont. 2012. Early evidence of San material culture and Norbert Mercier. 2013. How confident are we in the chronology of represented by organic artifacts from Border Cave, South Africa. Proceedings the transition between Howieson’s Poort and Still Bay? Journal of Human of the National Academy of Sciences of the USA 109(33):13214–13219. Evolution 64:314–317. d’Errico, Francesco, Lucinda Backwell, and Lyn Wadley. 2012. Identifying Hall, Grant, Stephan Woodborne, and M. Scholes. 2008. Stable carbon isotope regional variability in Middle Stone Age bone technology: the case of Sibudu ratios from archaeological charcoal as palaeoenvironmental indicators. Cave. Journal of Archaeological Science 39(7):2479–2495. Chemical Geology 247:384–400. d’Errico, Francesco, and William E. Banks. 2013. Identifying mechanisms Harper, Philip T. N. 1994. The Middle Stone Age sequence at Rose Cottage behind Middle Paleolithic and Middle Stone Age cultural trajectories. Cur- Cave: a search for continuity and discontinuity. Master’s thesis, University rent Anthropology 54(suppl. 8):S371–S387. of the Witwatersrand, Johannesburg. d’Errico, Francesco, and Christopher S. Henshilwood. 2007. Additional evi- ———. 1997. The Middle Stone Age sequence at Rose Cottage Cave: a search dence for bone technology in the southern African Middle Stone Age. for continuity and discontinuity. South African Journal of Science 93(10): Journal of Human Evolution 52:142–163. 470–475. d’Errico, Francesco, Christopher S. Henshilwood, Marian Vanhaeren, and Henderson, Zoe¨. 2001. Spatial patterning at southern African Middle Pleis- Karen van Niekerk. 2005. Nassarius kraussianus shell beads from Blombos tocene open-air sites: Florisbad, Duinefontein 2/2 and Mwanganda’s Village. Cave: evidence for symbolic behaviour in the Middle Stone Age. Journal of PhD thesis, University of Cambridge. Human Evolution 48:3–24. Henshilwood, Christopher S. 2012. Late Pleistocene techno-traditions in d’Errico, Francesco, Renata G. Moreno, and Riaan F. Rifkin. 2012. Techno- southern Africa: a review of the Still Bay and Howiesons Poort, c. 75–59 logical, elemental and colorimetric analysis of an engraved ochre fragment ka. Journal of World Prehistory 25:205–237. from the Middle Stone Age levels of Klasies River Cave 1, South Africa. Henshilwood, Christopher S., and Francesco d’Errico. 2011. Middle Stone Age Journal of Archaeological Science 39:942–952. engravings and their significance to the debate on the emergence of symbolic d’Errico, Francesco, Marian Vanhaeren, and Lyn Wadley. 2008. Possible shell material culture. In Homo symbolicus: the dawn of language, imagination beads from the Middle Stone Age layers of Sibudu Cave, South Africa. and spirituality. Christopher S. Henshilwood and Francesco d’Errico, eds. Journal of Archaeological Science 35(10):2675–2685. Pp. 75–96. Amsterdam: Benjamins. S316 Current Anthropology Volume 54, Supplement 8, December 2013

Henshilwood, Christopher S., Francesco d’Errico, Curtis W. Marean, Richard In Transitions before the transition: evolution and stability in the Middle G. Milo, and Royden Yates. 2001. An early bone tool industry from the Paleolithic and Middle Stone Age. Erella Hovers and Steven L. Kuhn, eds. Middle Stone Age at Blombos Cave, South Africa: implications for the Pp. 109–120. New York: Springer. origins of modern human behaviour, symbolism and language. Journal of Kuman, Kathleen. 1989. Florisbad and (Gi: the contribution of open air Human Evolution 41:631–678. sites to study of the Middle Stone Age in southern Africa. PhD dissertation, Henshilwood, Christopher S., Francesco d’Errico, Marian Vanhaeren, Karen University of Pennsylvania, Philadelphia. van Niekerk, and Zenobia Jacobs. 2004. Middle Stone Age shell beads from ———. 2007. The Earlier Stone Age in South Africa: site context and the South Africa. Science 304:404. influence of cave studies. In Breathing life fossils: taphonomic studies in honor Henshilwood, Christopher S., Francesco d’Errico, Karen L. van Niekerk, Yvan of C.K. (Bob) Brain. Travis Rayne Pickering, Kathy Schick, and Nicholas Coquinot, Zenobia Jacobs, Stein-Erik Lauritzen, Michel Menu, and Renata Toth, eds. Pp. 181–198. Bloomington IN: Stone Age Institute Press. Garcı´a-Moreno. 2011. A 100,000-year-old ochre-processing workshop at Kuman, Kathleen, Moshe Inbar, and Ronald J. Clark. 1999. Palaeoenviron- Blombos Cave, South Africa. Science 334(6053):219–222. ments and cultural sequence of the Florisbad Middle Stone Age hominid Henshilwood, Christopher S., Francesco d’Errico, and Ian Watts. 2009. En- site, South Africa. Journal of Archaeological Science 26:1409–1425. graved ochres from the Middle Stone Age levels at Blombos, South Africa. Lombard, Marlize. 2005. The Howiesons Poort of South Africa: what we know, Journal of Human Evolution 57:27–47. what we think we know, and what we need to know. South African Hu- Henshilwood, Christopher S., Francesco d’Errico, Royden Yates, Zenobia Ja- manities 17:33–55. cobs, Chantal Tribolo, Geoff A. T. Duller, Norbert Mercier, et al. 2002. ———. 2006. First impressions on the functions and hafting technology of Emergence of modern human behavior: Middle Stone Age engravings from Still Bay pointed artefacts from Sibudu Cave. South African Humanities 18: South Africa. Science 295:1278–1280. 27–41. Henshilwood, Christopher S., and Benoıˆt Dubreuil. 2011. The Still Bay and ———. 2007. The gripping nature of ochre: the association of ochre with Howiesons Poort, 77–59 ka: symbolic material culture and the evolution Howiesons Poort adhesives and Later Stone Age mastics from South Africa. of the mind during the African Middle Stone Age. Current Anthropology Journal of Human Evolution 53:406–419. 52(3):361–400. ———. 2008. Finding resolution for the Howiesons Poort through the mi- Henshilwood, Christopher S., and Curtis W. Marean. 2003. The origin of croscope: micro-residue analysis of segments from Sibudu Cave, South modern human behavior: a critique of the models and their test implica- Africa. Journal of Archaeological Science 35:26–41. tions. Current Anthropology 44:627–651. ———. 2009. The Howiesons Poort of South Africa amplified. South African Herries, Andy I. R. 2011. A chronological perspective on the Acheulian and Archaeological Bulletin 189:4–12. its transition to the Middle Stone Age in southern Africa: the question of ———. 2011. Quartz-tipped arrows older than 60 ka: further use-trace evi- the Fauresmith. International Journal of Evolutionary Biology 2011:1–25. dence from Sibudu, KwaZulu-Natal, South Africa. Journal of Archaeological Hiscock, Peter. 2007. Looking the other way: a materialist/technological ap- Science 38:1918–1930. proach to classifying tools and implements, cores and retouched flakes. In ———. 2012. Thinking through the Middle Stone Age of sub-Saharan Africa. Tools versus cores? alternative approaches to stone tool analysis. Shannon P. Quaternary International 270:140–155. McPherron, ed. Pp. 98–222. Newcastle: Cambridge Scholars. Lombard, Marlize, and Miriam N. Haidle. 2012. Thinking a bow-and-arrow: Hiscock, Peter, and Sue O’Connor. 2006. An Australian perspective on modern cognitive implications of Middle Stone Age bow and stone-tipped arrow behaviour and artefact assemblages. Before Farming 2:1–10. technology. Cambridge Archaeological Journal 22(2):237–264. Ho¨gberg, Anders, and Lars Larsson. 2011. Lithic technology and behavioural Lombard, Marlize, and Justin Pargeter. 2008. Hunting with Howiesons Poort modernity: new results from the Still Bay site, Hollow Rock Shelter, Western segments: pilot experimental study and the functional interpretation of Cape Province, South Africa. Journal of Human Evolution 61:133–155. archaeological tools. Journal of Archaeological Science 35:26–41. Hovers, Erella, and Anna Belfer-Cohen. 2013. On variability and complexity: Lombard, Marlize, and Isabelle Parsons. 2010. Fact or fiction? behavioural lessons from the Levantine Middle Paleolithic record. Current Anthropology and technological reversal after 60 ka in southern Africa. South African 54(suppl. 8):S337–S357. Archaeological Bulletin 65:224–228. Igreja, Marina, and Guillaume Porraz. 2013. Functional insights into the in- Lombard, Marlize, and Laurel Phillipson. 2010. Indications of bow and stone- novative Early Howiesons Poort technology at Diepkloof Rock Shelter tipped arrow use 64,000 years ago in KwaZulu-Natal. Antiquity 84:635– (Western Cape, South Africa). Journal of Archaeological Science 40:3475– 648. 3491. Lombard, Marlize, Carina Schlebusch, and Himla Soodyall. Forthcoming. Jacobs, Zenobia. 2010. An OSL chronology for the sedimentary deposits from Early Homo sapiens in southern Africa (88–160 ka): an overview of the Pinnacle Point Cave 13B: a punctuated presence. Journal of Human Evo- most recent DNA, fossil and archaeological evidence. South African Journal lution 59(3/4):289–305. of Science. Jacobs, Zenobia, Elspeth H. Hayes, Richard G. Roberts, Rex F. Galbraith, and Lombard, Marlize, Lyn Wadley, Janette Deacon, Sarah Wurz, Isabelle Parsons, Christopher S. Henshilwood. 2012. An improved OSL chronology for the Moleboheng Mohapi, Joane Swart, and Peter Mitchell. 2012. South African Still Bay layers at Blombos Cave, South Africa: further tests of single-grain and Lesotho Stone Age sequence updated. South African Archaeological dating procedures and a re-evaluation of the timing of the Still Bay industry Bulletin 67(195):123–144. across southern Africa. Journal of Archaeological Science 40:579–594. Loock, Johan C., and N. Johann Grobler. 1988. The regional geology of Flor- Jacobs, Zenobia, and Richard G. Roberts. 2009. Were environmental or de- isbad. Navorsinge van die Nasionale Museum, Bloemfontein 5:489–497. mographic factors the driving force behind Middle Stone Age innovations Lycett, Stephen J. 2009. Are Victoria West cores ‘‘proto-Levallois”? a phylo- in southern Africa? South African Journal of Science 105:333–334. genetic assessment. Journal of Human Evolution 56:175–191. Jacobs, Zenobia, Richard G. Roberts, Rex F. Galbraith, Hilary J. Deacon, Rainer ———. 2011. “Most beautiful and most wonderful”: those endless stone tool Gru¨n, Alex Mackay, Peter Mitchell, Ralf Vogelsang, and Lyn Wadley. 2008. forms. Journal of Evolutionary Psychology 9(2):143–171. Ages for the Middle Stone Age of southern Africa: implications for human Mackay, Alex. 2008a. A method for estimating edge length from flake di- behavior and dispersal. Science 322:733–735. mensions: use and implications for technological change in the southern Kandel, Andrew W., and Nicholas J. Conard. 2012. Settlement patterns during African MSA. Journal of Archaeological Science 35:614–622. the Earlier and Middle Stone Age around Langebaan Lagoon, Western Cape ———. 2008b. On the production of blades and its relationship to backed (South Africa). Quaternary International 270:15–29. artefacts in the Howiesons Poort at Diepkloof, South Africa. Lithic Tech- Kaplan, Jonathan. 1990. The Umhlatuzana rock shelter sequence: 100,000 nology 33(1):87–99. years of Stone Age history. Natal Museum Journal of Humanities 2:1–94. ———. 2009. History and selection in the late Pleistocene archaeology of the Klein, Richard G. 1976. The mammalian fauna of the Klasies River Mouth Western Cape, South Africa. PhD thesis, Australian National University, sites, southern Cape Province, South Africa. South African Archaeological Canberra. Bulletin 31:75–98. ———. 2011. Potentially stylistic differences between backed artefacts from Klein, Richard G., Graham Avery, Kathryn Cruz-Uribe, David Halkett, Tim two nearby sites occupied 60,000 years before present in South Africa. Hart, Richard G. Milo, and Thomas P. Volman. 1999. Duinefontein 2: an Journal of Anthropological Archaeology 30(2):235–245. Acheulean site in the Western Cape Province of South Africa. Journal of Mackay, Alex, and Aara Welz. 2008. Engraved ochre from a Middle Stone Age Human Evolution 37:153–190. context at Klein Kliphuis in the Western Cape of South Africa. Journal of Kuhn, Steven L. 2006. Trajectories of change in the Middle Paleolithic in Italy. Archaeological Science 35:1521–1532. Wurz Technological Trends in the Middle Stone Age of South Africa S317

Manengold, Albrecht, and James S. Brink. 2011. Descriptions and palaeo- industry from Kathu Pan, southern Africa: implications for the Earlier to ecological implications of bird remains from the Middle Pleistocene of Middle Stone Age transition. Journal of Archaeological Science 37(2):269– Florisbad, South Africa. Pala¨ontologische Zeitschrift 85:19–32. 283. Marean, Curtis W. 2010. Pinnacle Point Cave 13B (Western Cape Province, Porraz, Guillaume, John E. Parkington, Jean-Philippe Rigaud, Christopher M. South Africa) in context: the Cape floral kingdom, shellfish, and modern Miller, Cedric Poggenpoel, Chantal Tribolo, Will Archer, et al. 2013a.The human origins. Journal of Human Evolution 59:425–443. MSA sequence of Diepkloof and the history of southern African Late Pleis- ———. 2011. Coastal South Africa and the co-evolution of the modern hu- tocene populations. Journal of Archaeological Science 40:3542–3552. man lineage and the coastal adaptation. In Trekking the shore: changing Porraz, Guillaume, Pierre-Jean Texier, Will Archer, Michel Piboule, Jean-Phi- coastlines and the antiquity of coastal settlement. Nuno F. Bicho, Jonathan lippe Rigaud, and Chantal Tribolo. 2013b. Technological successions in the A. Haws, and Loren G. Davis, eds. Pp. 421–440. New York: Springer. Middle Stone Age sequence of Diepkloof Rock Shelter, Western Cape, South Marean, Curtis W., Miryam Bar-Matthews, Jocelyn Bernatchez, Erich Fisher, Africa. Journal of Archaeological Science 40:3376–3400. Paul Goldberg, Andy I. R. Herries, Zenobia Jacobs, et al. 2007. Early human Porraz, Guillaume, Pierre-Jean Texier, Jean-Philippe Rigaud, John Parkington, use of marine resources and pigment in South Africa during the Middle Cedric Poggenpoel, and Dave Roberts. 2008. Preliminary characterization Pleistocene. Nature 449:905–908. of a Middle Stone Age lithic assemblage preceding the “classic” Howiesons Mason, Revil J. 1962. Prehistory of the Transvaal: a record of human activity. Poort complex at Diepkloof Rock Shelter, Western Cape Province, South Johannesburg: Witwatersrand University Press. Africa). South African Archaeological Society Goodwin Series 10:105–121. McBrearty, Sally, and Alison S. Brooks. 2000. The revolution that wasn’t: a Quintana-Murci, Lluis, He´le`ne Quach, Christine Harmant, Francesca Luca, new interpretation of the origin of modern human behavior. Journal of Blandine Massonnet, Etienne Patin, Lucas Sica, et al. 2008. Maternal traces Human Evolution 39:453–563. of deep common ancestry and asymmetric gene flow between Pygmy McCall, Grant S. 2007. Behavioral ecological models of lithic technological hunter-gatherers and Bantu-speaking farmers. Proceedings of the National change during the later Middle Stone Age of South Africa. Journal of Ar- Academy of Sciences of the USA 105:1596–1601. chaeological Science 34:1738–1751. Sampson, C. Garth. 1968. The Stone Age industries of the Orange River scheme McCall, Grant S., and Jonathan T. Thomas. 2012. Still Bay and Howiesons and South Africa. National Museum, Bloemfontein Memoir, no. 6. Bloem- Poort foraging strategies: recent research and models of culture change. fontein, South Africa: National Museum. African Archaeological Review 29:7–50. ———. 1974. The Stone Age archaeology of southern Africa. New York: Aca- McNabb, John, and Peter Beaumont. 2011. A report on the archaeological demic Press. assemblages from excavations by Peter Beaumont at Canteen Koppie, Northern Scheinfeldt, Laura B., Soi Sameer, and Sarah A. Tishkoff. 2010. Working toward Cape, South Africa. University of Southampton Series in Archaeology 4. a synthesis of archaeological, linguistic, and genetic data for inferring Af- Oxford: Archaeopress. rican population history. Proceedings of the National Academy of Sciences of ———. 2012. Excavations in the Acheulean levels at the Earlier Stone Age the USA 107(suppl. 2):8931–8938. site of Canteen Koppie, Northern Province, South Africa. Proceedings of the Schlebusch, Carina M., Pontus Skoglund, Per Sjodin, Lucie M. Gattepaille, Prehistoric Society 78:51–71. Dena Hernandez, Flora Jay, Sen Li, et al. 2012. Genomic variation in seven Mellars, Paul. 2006. Why did modern human populations disperse from Africa Khoe-San groups reveals adaptation and complex African history. Science ca. 60,000 years ago? a new model. Proceedings of the National Academy of 338:374–379. Sciences of the USA 103:9381–9386. Schmidt, Patrick, Guillaume Porraz, Aneta Slodczyk, Ludovic Bellot-Gurlet, Minichillo, Tom. 2005. Middle Stone Age lithic study, South Africa: an ex- William Archer, and Christopher E. Miller. 2013. Heat treatment in the amination of modern human origin. PhD dissertation, University of Wash- South African Middle Stone Age: temperature induced transformations of ington, Seattle. silcrete and their technological implications. Journal of Archaeological Science ———. 2006. Raw material use and behavioural modernity: Howiesons Poort 40:3519–3531. lithic foraging strategies. Journal of Human Evolution 50:359–364. Schoville, Benjamin J. 2010. Frequency and distribution of edge damage on Mitchell, Peter. 2002. Hunter-gatherer archaeology in southern Africa: recent Middle Stone Age lithic points, Pinnacle Point 13B, South Africa. Journal research, future trends. Before Farming 1(3):1–36. of Human Evolution 59:378–391. ———. 2008. Developing the archaeology of marine isotope stage 3. South Scott, Louis, and James S. Brink. 1992. Quaternary palaeoenvironments of African Archaeological Society Goodwin Series 10:52–65. pans in central South Africa: palynological and palaeontological evidence. ———. 2012. San origins and transition to the Later Stone Age: new research Suid-Afrikaanse Geograaf 19:22–34. from Border Cave, South Africa. South African Journal of Science 108(11/ Scott, Louis, and Lloyd Rossouw. 2005. Reassessment of botanical evidence 12):1–2. for palaeoenvironments at Florisbad, South Africa. South African Archae- Mohapi, M. 2012. Point morphology and the Middle Stone Age cultural ological Bulletin 60:96–102. sequence of Sibudu Cave, KwaZulu-Natal, South Africa. South African Ar- Shea, John J. 2009. The impact of projectile weaponry on Late Pleistocene chaeological Bulletin 67(195):5–15. hominin evolution. In The evolution of hominin diets: integrating approaches Moncel, Marie-He´le`ne, Anne-Marie Moigne, Youssef Sam, and Jean Combier. to the study of Palaeolithic subsistence. Jean-Jacques Hublin and Michael 2011. The emergence of Neanderthal technical behavior: new evidence from Richards, eds. Pp. 89–199. Dordrecht: Springer. Orgnac 3 (Level 1, MIS 8), southeastern France. Current Anthropology 52(1): Sinclair, Anthony, and John McNabb. 2005. All in a day’s work: Middle Pleis- 37–75. tocene individuals, materiality and the lifespace at Makapansgat, South Mourre, Vincent, Paola Villa, and Christopher S. Henshilwood. 2010. Early Africa. In The individual hominid in context: archaeological investigations of use of pressure flaking on lithic artifacts at Blombos Cave, South Africa. Lower and Middle Palaeolithic landscapes, locales and artefacts. Clive Gamble Science 330:659–662. and Martin Porr, eds. Pp. 176–196. London: Routledge. North Greenland Ice Core Project Members. 2004. High-resolution record of Singer, Ronald, and John Wymer. 1982. The Middle Stone Age at Klasies River Northern Hemisphere climate extending into the last interglacial period. Mouth in South Africa. Chicago: Chicago University Press. Nature 431:147–151. Sjo¨din, Per, Agne`s E. Sjo¨strand, Mattias Jakobsson, and Michael G. B. Blum. Pahnke, Katharina, Rainer Zahn, Harry Elderfield, and Michael Schulz. 2003. 2012. Resequencing data provide no evidence for a human bottleneck in 340,000-year centennial-scale marine record of Southern Hemisphere cli- Africa during the penultimate glacial period. Molecular Biology and Evolution matic oscillation. Science 301:948–952. 2012:1–30. Petit, Jean-Robert, Jean Jouzel, Dominique Raynaud, Narcisse I. Barkov, Jean- Soriano, Sylvain, Paola Villa, and Lyn Wadley. 2007. Blade technology and Marc Barnola, Isabelle Basile, Michael Bender, et al. 1999. Climate and tool forms in the Middle Stone Age of South Africa: the Howiesons Poort atmospheric history of the past 420,000 years from the Vostok ice core, and post–Howiesons Poort at Rose Cottage Cave. Journal of Archaeological Antarctica. Nature 399:429–436. Science 34:681–703. Pickrell, Joseph K., Nick Patterson, Chiara Barbieri, Falko Berthold, Linda Steele, Teresa E., and Richard G. Klein. 2009. Late Pleistocene subsistence Gerlach, Tom Gu¨ldemann, Blesswell Cure, et al. 2012. The genetic pre- strategies and resource intensification in Africa. In The evolution of hominin history of southern Africa. Nature Communications 3:1143, doi:10.1038 diets: integrating approaches to the study of Palaeolithic subsistence. Jean- /ncomms2140. Jacques Hublin and Michael P. Richards, eds. Pp. 113–126. New York: Porat, Naomi, Michael Chazan, Rainer Gru¨n, Maxime Aubert, Vera Eisen- Springer. mann, and Liora K. Horwitz. 2010. New radiometric ages for the Fauresmith Steele, Teresa E., Alex Mackay, Jayson Orton, and Steve Schwortz. 2012. Var- S318 Current Anthropology Volume 54, Supplement 8, December 2013

sche Rivier 003, a new Middle Stone Age site in southern Namaqualand, Villa, Paola, Sylvain Soriano, Tsenka Tsanova, Ilaria Degano, Thomas F. G. South Africa. South African Archaeological Bulletin 67(195):108–119. Higham, Francesco d’Errico, Lucinda Backwell, Jeannette J. Lucejko, Maria Texier, Pierre-Jean, Guillaume Porraz, John Parkington, Jean-Philippe Rigaud, Perla Colombini, and Peter B. Beaumont. 2012. Border Cave and the be- Cedric Poggenpoel, Christopher Miller, Chantal Tribolo, et al. 2010. A How- ginning of the Later Stone Age in South Africa. Proceedings of the National iesons Poort tradition of engraved ostrich eggshell containers dated to Academy of Sciences of the USA 109(33):13208–13213. 60,000 years ago at Diepkloof Rock Shelter, South Africa. Proceedings of the Visser, Johan N. J., and Alta Joubert. 1991. Cyclicity in the Late Pleistocene National Academy of Sciences of the USA 107:6180–6185. to Holocene spring and lacustrine deposits at Florisbad, Orange Free State. Texier, Pierre-Jean, Guillaume Porraz, John Parkington, Jean-Philippe Rigaud, South African Journal of Geology 94:123–131. Cedric Poggenpoel, and Chantal Tribolo. 2013. The context, form and Volman, Tom. 1978. Early archaeological evidence for shellfish collecting. Sci- significance of the MSA engraved ostrich eggshell collection from Diepkloof ence 201:911–913. Rock Shelter, Western Cape, South Africa. Journal of Archaeological Science ———. 1981. The Middle Stone Age in the southern Cape. PhD dissertation, 40:3412–3431. University of Chicago. Thackeray, Anne I. 1989. Changing fashions in the Middle Stone Age: the ———. 1984. Early prehistory of Southern Africa. In Southern African pre- stone artefact sequence from Klasies River main site, South Africa. African history and paleoenvironments. Richard G. Klein ed. Pp. 169–220. Rotter- Archaeological Review 7:33–57. dam: Balkema. ———. 1992. The Middle Stone Age south of the Limpopo River. Journal of Wadley, Lyn. 1993. The Pleistocene Later Stone Age south of the Limpopo World Prehistory 6(4):385–431. River. Journal of World Prehistory 7:243–296. ———. 2000. Middle Stone Age artefacts from the 1993 and 1995 excavations ———. 2005. A typological study of the final Middle Stone Age stone tools of Die Kelders Cave 1, South Africa. Journal of Human Evolution 38:147– from Sibudu Cave, KwaZulu-Natal. South African Archaeological Bulletin 168. 60:51–63. Thackeray, Anne I., and Alison J. Kelly. 1988. A technological and typological ———. 2007. Announcing a Still Bay Industry at Sibudu Cave, South Africa. analysis of Middle Stone Age assemblages antecedent to the Howiesons Journal of Human Evolution 52:681–689. Poort at Klasies River main site. South African Archaeological Bulletin 43: ———. 2008. The Howiesons Poort Industry of Sibudu Cave. South African 15–26. Archaeological Society Goodwin Series 10:122–132. Thackeray, J. Francis. 2009. Chronology, climate and technological innovation ———. 2010. Compound-adhesive manufacture as a behavioral proxy for associated with the Howiesons Poort and Still Bay industries in South Africa. complex cognition in the Middle Stone Age. Current Anthropology 51(suppl. South African Journal of Science 105:90. 1):S111–S119. Thompson, Erin, and Curtis Marean. 2008. The Mossel Bay lithic variant: ———. 2013. Recognizing complex cognition through innovative technology 120 years of Middle Stone Age research from Cape St Blaize Cave to Pinnacle in Stone Age and Palaeolithic sites. Cambridge Archeological Journal 23(2): Point. South African Archaeological Society Goodwin Series 10:90–104. 163–183. Thompson, Erin, Hope M. Williams, and Tom Minichillo. 2010. Middle and Wadley, Lyn, and Philip T. N. Harper. 1989. Rose Cottage revisited: Malan’s late Pleistocene Middle Stone Age lithic technology from Pinnacle Point Middle Stone Age collection. South African Archaeological Bulletin 44:23– 13B (Mossel Bay, Western Cape Province, South Africa). Journal of Human 32. Evolution 59:358–377. Wadley, Lyn, Tamaryn Hodgskiss, and Michael Grant. 2009. Implications for Tishkoff, Sarah A., Mary K. Gonder, Brenna M. Henn, Holly Mortensen, Alec complex cognition from the hafting of tools with compound adhesives in Knight, Christopher Gignoux, Neil Fernandopulle, et al. 2007. History of the Middle Stone Age, South Africa. Proceedings of the National Academy click-speaking populations of Africa inferred from mtDNA and Y chro- of Sciences of the USA 106(24):9590–9594. mosome genetic variation. Molecular Biology and Evolution 24:2180–2195. Wadley, Lyn, and Moleboheng Mohapi. 2008. A segment is not a monolith: Tribolo, Chantal, Norbert Mercier, E. Douville, J.-L. Joron, D. Reyss, N. Rufer, evidence from the Howiesons Poort of Sibudu, South Africa. Journal of N. Cantin, et al. 2013. OSL and TL dating of the Middle Stone Age sequence Archaeological Science 35:2594–2605. at Diepkloof Rock Shelter (South Africa): a clarification. Journal of Ar- Watts, Ian. 2010. The pigments from Pinnacle Point Cave 13B, Western Cape, chaeological Science 40:3401–3411. South Africa. Journal of Human Evolution 59:392–411. Tribolo, Chantal, Norbert Mercier, and He´le`ne Valladas. 2005. Chronology of White, Mark, Nick Ashton, and Beccy Scott. 2011. The emergence, diversity the Howiesons Poort and Still Bay technocomplexes: assessment and new and significance of Mode 3 (prepared core) technologies. In The ancient data from luminescence. In From tools to symbols: from early hominids to human occupation of Britain. Nick Ashton, Simon Lewis, and Chris Stringer, modern humans. Francesco d’Errico and Lucinda Backwell, eds. Pp. 493– eds. Pp. 53–65. Amsterdam: Elsevier. 512. Johannesburg: Witwatersrand University Press. Wilkins, Jayne, and Michael Chazan. 2012. Blade production ∼ 500 thousand Tribolo, Chantal, Norbert Mercier, He´le`ne Valladas, Jean-Louis Joron, Pierre years ago at Kathu Pan 1, South Africa: support for a multiple origins Guibert, Yannick Lefrais, Madeleine Selo, et al. 2009. Thermoluminescence hypothesis for Early Middle Pleistocene blade technologies. Journal of Ar- dating of a Stillbay–Howiesons Poort sequence at Diepkloof Rock Shelter chaeological Science 39:1883–1900. (Western Cape, South Africa). Journal of Archaeological Science 36:730–739. Wilkins, Jayne, Luca Pollarolo, and Kathleen Kuman. 2010. Prepared core Tryon, Christian A., and J. Tyler Faith. 2013. Variability in the Middle Stone reduction at the site of Kudu Koppie in northern South Africa: temporal Age of eastern Africa. Current Anthropology 54(suppl. 8):S234–S254. patterns across the Earlier and Middle Stone Age boundary. Journal of Van Peer, P. 1992. The Levallois reduction strategy. Monographs in World Archaeological Science 37:1279–1292. Archaeology, no. 13. Madison, WI: Prehistory Press. Wilkins, Jayne, Benjamin J. Schoville, Kyle S. Brown, and Michael Chazan. Van Pletzen, Vos L. 2000. The large mammal fauna from Klasies River. Master’s 2012. Evidence for early hafted hunting technology. Science 338:942–946. thesis, University of Stellenbosch. Will, Manuel, John E. Parkington, Andrew W. Kandel, and Nicholas J. Conard. Van Riet Lowe, Clarence. 1945. The evolution of the Levallois technique in 2013. Coastal adaptations and the Middle Stone Age lithic assemblages from South Africa. Man 45:49–59. Hoedjiespunt 1 in the Western Cape, South Africa. Journal of Human Evo- Van Zinderen Bakker, Eduard M. 1989. Middle Stone Age palaeoenvironments lution 64:518–537. at Florisbad (South Africa). Palaeoecology of Africa 20:133–154. Wurz, Sarah. 1999. The Howiesons Poort at Klasies River: an argument for Villa, Paola, Anne Delagnes, and Lyn Wadley. 2005. A Late Middle Stone Age symbolic behaviour. South African Archaeological Bulletin 54(69):38–50. artifact assemblage from Sibudu (KwaZulu-Natal): comparisons with the ———. 2000. The Middle Stone Age sequence at Klasies River, South Africa. European Middle Paleolithic. Journal of Archaeological Science 32:399–422. PhD dissertation, University of Stellenbosch. Villa, Paola, Marie Soressi, Christopher S. Henshilwood, and Vincent Mourre. ———. 2002. Variability in the Middle Stone Age lithic sequence, 115,000– 2009. The Still Bay points of Blombos Cave (South Africa). Journal of 60,000 years ago at Klasies River, South Africa. Journal of Archaeological Archaeological Science 36:441–460. Science 29:1001–1015. Villa, Paola, and Sylvain Soriano. 2010. Hunting weapons of Neanderthals ———. 2008. Modern human behaviour at Klasies River. South African Ar- and early modern humans in South Africa. Journal of Anthropological Re- chaeological Society Goodwin Series 10:150–156. search 66:5–38. ———. 2010. Middle Stone Age tools from Klasies River main site: conven- Villa, Paola, Sylvain Soriano, Nicolas Teyssandier, and Sarah Wurz. 2010. The tions and symbolic cognition. In The cutting edge: stone tools and the evo- Howiesons Poort and MSA III at Klasies River Mouth, Cave 1A. Journal lution of cognition. April Nowell and Iain Davidson, eds. Pp. 135–158. of Archaeological Science 37:630–655. Boulder: Colorado University Press. Wurz Technological Trends in the Middle Stone Age of South Africa S319

———. 2012. The significance of MIS 5 shell middens on the Cape coast: Wynn, Thomas, and Frederick L. Coolidge. 2011. The implications of the lithic perspective from Klasies River and Ysterfontein 1. Quaternary Inter- working memory model for the evolution of modern cognition. Interna- national 270:61–69. tional Journal of Evolutionary Biology 2011:1–12. Wurz, Sarah, Niel J. le Roux, Sugnet Gardner, and Hilary J. Deacon. 2003. Ziegler, Martin, Margit H. Simon, Ian R. Hall, Stephen Barker, Chris Stringer, Discriminating between the end products of the Earlier Middle Stone Age and Rainer Zahn. 2013. Development of Middle Stone Age innovation sub-stages at Klasies River using biplot methodology. Journal of Archaeo- linked to rapid climate change. Nature Communications 4:1905, doi:10.1038 logical Science 30:1107–1126. /ncomms2897. S320 Current Anthropology Volume 54, Supplement 8, December 2013

Change and Stasis in the Iberian Middle Paleolithic Considerations on the Significance of Mousterian Technological Variability

by Ignacio de la Torre, Jorge Martı´nez-Moreno, and Rafael Mora

CAϩ Online-Only Material: Supplements A and B

The European Mousterian has traditionally been portrayed as a long period of technological stasis as opposed to the technotypological dynamism of Upper Paleolithic cultures. The classic debate on Mousterian variability explained interassemblage differences either by ethnic, cultural, functional, and chronological or by paleoenvironmental causes, but variability was based on typological considerations. Recently, technological factors have been introduced in discussions over time trends and geographic differences in the Mousterian. This paper will address the topic by reviewing technological strategies in the Iberian Middle Paleolithic. Three sites from northeastern Spain are chosen as a case study to address the existence of directional patterns in the Iberian Mousterian. We conclude that albeit diachronic variability exists, it does not show patterning, which suggests stochastic variation rather than directional change in the technological strategies of Iberian Neanderthals.

The notion of variability is at the heart of the study of the novative emphasis on the role of ecological factors in tech- Mousterian. Spanning ca. 300 kyr and present in hundreds nological behavior (Kuhn 1995) added a novel dimension to of sites across western Europe, the search for explanations of the study of Middle Paleolithic variability. interassemblage variability is consubstantial to Middle Pale- In recent years, generalization of technological perspectives olithic research. The classic debate on the interpretation of in most of western European Middle Paleolithic studies and the Mousterian facies privileged cultural (Bordes 1972), func- the dramatic improvement of dating techniques for chro- tional (Binford 1973), diachronic (Mellars 1969), or climatic nologies beyond the accelerator mass spectrometry (AMS) 14C (Laville 1973) explanations, but it was strictly typological; that method have provided new tools to address the question of is, it was based on the assumption that Bordes’s tool types variability and time depth, now with an emphasis on the were meaningful categories. Since the 1980s, the advent of understanding of knapping techniques rather than tool types. technological approaches in Mousterian research (e.g., Boe¨da Interest in evolution, stasis, and rhythms of change during 1986; Geneste 1985) and the consideration of tool types as the Middle Paleolithic (e.g., Kuhn and Hovers 2006) has shed the result of different stages of reduction (Dibble 1987) led new light in Neanderthal cultural adaptations, with pioneering to explanations of most variability by raw material constraints studies focused on the technological characterization of and functional or settlement dynamics (e.g., Dibble and Rol- Mousterian diachronic variability (Delagnes and Meignen land 1992). New attempts to understand the Mousterian facies 2006) and its functional/ecological correlation (Delagnes and from a functional perspective (e.g., Beyries 1988) and an in- Rendu 2011). The aim of this paper is to apply some of these perspectives to the Mousterian of the Iberian Peninsula with the specific Ignacio de la Torre is Reader in Palaeolithic Archaeology at the purpose of addressing the potential significance of diachronic Institute of Archaeology of University College London (31–34 patterns in lithic technology. A long history of research, abun- Gordon Square, WC1H 0PY London, United Kingdom [i.torre@ dance of karstic systems susceptible of preserving archaeo- ucl.ac.uk]). Jorge Martı´nez-Moreno is Research Associate and Rafael Mora is Professor of Prehistory at the Centre d’Estudis del Patrimoni logical deposits, and relatively milder conditions than in most Arqueologic de la Prehistoria of the Universitat Auto´noma de of Europe during the Pleistocene explains the substantial Barcelona (Facultat de Lletres, 08193 Bellaterra, Spain). This paper number of Mousterian sites known in Spain and Portugal. was submitted 3 VII 13, accepted 27 VIII 13, and electronically However, problems on the chronostratigraphic correlation of published 20 XII 13. assemblages plus the variety of nomenclatures and analytical

᭧ 2013 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved. 0011-3204/2013/54S8-0012$10.00. DOI: 10.1086/673861 de la Torre, Martı´nez-Moreno, and Mora Change and Stasis in the Iberian Middle Paleolithic S321 perspectives employed make it difficult to produce overall the most recent Middle Paleolithic sites in northern Iberia, assessments such as those currently available for adjacent perhaps as late as 33–32 kyr (Martı´nez-Moreno, Mora, and regions (Delagnes and Meignen 2006; Jaubert 2011) and the de la Torre 2010). Near East (Goren-Inbar and Belfer-Cohen 1998; Hovers and The three sites share a number of features enabling inter- Belfer-Cohen 2013). assemblage comparisons. Both Roca dels Bous and Trago´are In order to overcome the sample disparity derived from situated over two main river valleys that are natural corridors such methodological and empirical pitfalls, three Iberian sites between the Ebro plain and the first ranges of the Pyrenees excavated and studied within the same research program (Ca- (fig. A1), while Cova Gran is located in a small subsidiary sanova et al. 2009; Martı´nez-Moreno et al. 2010; Mora et al. valley about 10 km north from Roca dels Bous. Pleistocene 2011) in the area of La Noguera (northeast Spain) are used biotopes should have been similar in the three sites, with in this paper to search for temporal trends in the Middle variations in the diversity of game according to climatic pulses Paleolithic. Our results suggest that although diachronic var- but comparable density and availability of biotic resources. iation is detected, it does not follow any particular pattern. Availability of raw material was also similar; flint of poor This case study from La Noguera will then be discussed within quality is present in the immediate surroundings of Cova Gran the context of the Iberian Mousterian, in which a lack of time and Trago´but not in the proximity of Roca dels Bous. Good trends also prevails. Instead, the observed technological var- quality flint was not locally available in any of the three sites. iability could be associated with intrasite specifics and, po- Quartzite and other metamorphic cobbles were readily avail- tentially, regional idiosyncrasies. This apparent lack of tem- able from river beds in the three sites, particularly in Roca poral patterns in the Iberian Middle Paleolithic contrasts with dels Bous and Trago´. neighboring areas such as France, where chronological trends In order to evaluate the possible existence of temporal seem to exist (e.g., Delagnes and Meignen 2006). By com- trends, in this paper the Roca dels Bous (Martı´nez-Moreno, bining different units of analysis (from particular case studies Mora, and de la Torre 2010; Mora, de la Torre, and Martı´nez- in northeast Spain to the regional scale of Iberia and to its Moreno 2004), Cova Gran (Martı´nez-Moreno, Mora, and de contextualization within the sequence of southwestern Eu- la Torre 2010), and Trago´ (Casanova 2009) assemblages will rope), the purpose of this paper is to contribute to the debate be organized diachronically; Trago´levels cover oxygen isotope on technological change and stasis during the Middle Pale- stage (OIS) 5e (UA3), OIS 5/4 (UA2), and OIS 3 (UA1). Roca olithic and the nature and causes of Mousterian technological dels Bous levels are probably within the same chronological variability. range as UA1, but for the sake of comparison, N12 and N10 have been placed after the sequence of Trago´. Cova Gran levels S1D, S1C, and S1B have yielded consistent dates post 40 kyr and therefore are considered here to belong to the latter part The Middle Paleolithic of La Noguera: of OIS 3. A Case Study on the Analysis of The artefacts (N p 59,001 ) analyzed from the three sites Technological Diachronic Patterns are distributed unevenly, with 23,557 lithics from Trago´, 19,569 from Roca dels Bous, and 15,875 from Cova Gran Trago´, Roca dels Bous, and Cova Gran are three rockshelters (CAϩ Supplement B: table B2). Relative frequencies of main in the region of La Noguera (Lleida, Catalunya) located within stone tool types are shown in figure 1. The percentage of cores a radius of less than 20 km at the Marginal Exterior Sierras does not indicate any clear patterning; while cores are con- of the Eastern Pre-Pyrenees in the northeast of the Iberian sistently less abundant in Cova Gran than in the two older Peninsula (CAϩ Online Supplement A: fig. A1). Trago´has sites, N10 at Roca dels Bous yields the highest proportion of yielded eight archaeological levels, all attributed to the Mous- cores of all levels. Therefore, the comparative paucity of cores terian. Thermoluminescence dating (CAϩ Online Supple- in the Cova Gran levels is better explained by site-specific ment B: table B1) situates the bottom layer (UA3) at ap- economic variables rather than by temporal trends. In con- proximately 126 kyr and the uppermost (UA1) level of the trast, frequency of flakes seems to decrease steadily through- main excavation area at ca. 42 kyr (Casanova et al. 2009). out the sequence, with the exception of S1C in Cova Gran Ongoing excavations in Roca dels Bous have yet to reach the (fig. 1B). The meaning of such consistent reduction on the bedrock, and so far four main levels (R3, N10, N12, and N14) frequency of flakes is obscure and could tentatively be related have been unearthed (Martı´nez-Moreno et al. 2010; Mora, to recurrent export of flakes off site. A divide seems to exist de la Torre, and Martı´nez-Moreno 2004), all corresponding in the sequence with regard to the frequency of retouched -kyr, while tools; while the four older assemblages yield very low fre 1.2 ע to the Mousterian. R3 was dated to 38.8 underlying levels seem to be placed beyond the range of AMS quencies, all levels from N10 to S1B contain percentages of 14C (see table B1). Cova Gran is a huge (12,500 m2) rockshelter retouched tools over 7% (fig. 1C). Even though no cumulative with Upper Pleistocene and early Holocene deposits (Mora increase is observed across the four younger levels, the fact et al. 2011). Four Mousterian levels (S1B, S1C, S1D, and S1E) that a higher frequency of retouched tools is present in two have been documented so far, situating Cova Gran among sites (Roca dels Bous and Cova Gran) rules out the possibility Figure 1. Frequency and size of main categories in Trago´(Ua3, Ua2, Ua1), Roca dels Bous (N12, N10), and Cova Gran (S1D, S1C, S1B). A, Percentage of cores. B, Percentage of whole flakes. C, Percentage of retouched tools. D, Average length of cores. E, Elongation (length divided by width) of whole flakes. F, Mean length of retouched tools. G, Mean length of whole flakes. H, Average length of cores, whole flakes, and retouched tools. All data from CAϩ Online Supplement B: table B2. A color version of this figure is available in the online edition of Current Anthropology. de la Torre, Martı´nez-Moreno, and Mora Change and Stasis in the Iberian Middle Paleolithic S323 that this pattern was site-specific and could suggest some ually (see also CAϩ Supplement B: table B5). There seem to intensification in the shaping of blanks among latest Pre- be some differences between Trago´ and the other two sites; Pyrenean Neanderthals. while in Trago´ nearly all shaped tools are either denticulates Metric analysis of the main categories (fig. 1; CAϩ Sup- or sidescrapers, both in Roca dels Bous and Cova Gran, higher plement B: table B3) reflects consistency on lithic dimensions typological variability is attested, with points, endscrapers, and throughout the sequence. Average length of cores and re- other tool types present in all levels. Trago´ also consistently touched tools is remarkably similar from UA3 to S1B (fig. yields higher proportions of sidescrapers than Roca dels Bous 1D,1F), and even though flakes are larger in the oldest as- and Cova Gran. However, it is unclear whether such a pattern semblages (fig. 1G), the length/width ratio (fig. 1E) of Trago´ responds to temporal trends or whether it could be explained flakes resembles that of Cova Gran, which yields the most instead by settlement dynamics specific to the earlier site of recent layers. The widely documented pattern of larger blank Trago´. selection for retouching during the Mousterian (e.g., Dibble It remains to assess the role of raw materials in discerning and Rolland 1992; Geneste 1985) is also verified in each and potential time trends. Figure 2D suggests an overall preference every one of the assemblages analyzed here, which show con- for flint across the sequence in all levels apart from N12. sistent larger size of retouched tools over unmodified blanks. Interestingly, the four more recent layers show clear preference In general, data from figure 1 (see also table B3) do not for flint, whereas the earlier assemblages of Trago´yield a more indicate any kind of temporal trend but rather remarkably mixed procurement of flint and quartzite (see also CAϩ Sup- homogeneous size range for the main lithic categories across plement B: table B6). However, the exception of N12 in Roca all assemblages. The only possible divergence is shown by dels Bous, where the raw material procurement strategy is N10 at Roca dels Bous, whose cores and flakes are noticeably completely reversed to that of the later levels, precludes es- smaller than those in the rest of the levels. However, the short- tablishing a clear diachronic pattern throughout the assem- term nature of this occupation (Martı´nez-Moreno, Mora, and blages. The same applies to lithic categories with particular de la Torre 2004; Mora, de la Torre, and Martı´nez-Moreno technological relevance; figure 2E indicates that with the ex- 2004) rather than any chronological pattern explains partic- ception of N12, flint cores prevail in all assemblages. Similarly, ularities of N10 more satisfactorily. flint was more often selected for structured knapping than In order to explore the possible presence/existence of quartzite. No temporal trend can be observed, either, in the diachronic patterns on knapping methods, nearly 500 cores selection of raw material for retouched tools; while the youn- were considered in our analysis (CAϩ Supplement B: table ger levels of Cova Gran reflect a strong preference for flint B4). Levallois cores are poorly represented, with only N12 over quartzite, such preferential selection is exacerbated in and S1C yielding relevant percentages. Bifacial hierarchical UA1 at Trago´, one of the earlier assemblages. Although centripetal (sensu Casanova et al. 2009; de la Torre and Mora quartzite retouched tools in N12 are more abundant than 2004) and unifacial methods are the most common in several flint in absolute terms, flint was preferentially used for re- assemblages, whereas discoid cores sensu stricto are rare. Fig- touching, which is consistent with the pattern observed in the ure 2A suggests that neither Levallois nor any other flaking rest of the sequence (fig. 2F). methods show temporal trends. To avoid subjectivities derived from flaking methods classification, a comparison (fig. 2B) was made between expedient cores—those that bear only a Chronological Patterns of the Iberian few scars (e.g., unifacial) and which underwent limited and/ Middle Paleolithic or unstandardized (e.g., multifacial) reduction—versus struc- tured cores—those including several stages of preparation Although better known for putatively yielding the last Ne- and/or production (e.g., Levallois, bifacial hierarchical cen- anderthal traces anywhere in Europe (e.g., Finlayson et al tripetal and discoid). Albeit expedient methods are rarely 2006; Zilha˜o 2008), the Iberian record also contains a number mentioned in the literature, they are common in the Pre- of Middle and early Upper Pleistocene sequences. In principle, Pyrenean Mousterian and indicate opportunistic reduction of this should provide the time depth required to evaluate part of the raw material stocks. However, once again no tem- diachronic trends in the Iberian Mousterian. Unfortunately, poral pattern is discerned in the variation between expedient the wealth of Middle Paleolithic sites in Iberia (fig. 3) is not and structured methods: while structured methods tend to accompanied by a solid chronostratigraphic framework; in prevail (as expected from any Mousterian assemblage), ex- recent years, research agendas have targeted dating of the last pedient flaking is common at the beginning (UA3), middle Mousterian and the transition to the Upper Paleolithic (N10), and end (S1B) of the sequence. (d’Errico and Sa´nchez-Gon˜i 2003; Maroto et al 2012; Mar- Low variability of retouched tool types is well attested across tı´nez-Moreno, Mora, and de la Torre 2010; Zilha˜o 2006), the northeastern Iberian Mousterian sites (Mora 1988) and whereas older sequences are poorly constrained or simply lack is confirmed in the Pre-Pyrenees sequence. Figure 2C shows any radiometric dating. Thus, figure 4 must be seen as a very that sidescrapers and denticulates predominate, while points, preliminary attempt to order chronologically some of the rel- endscrapers, and other retouched tools are represented resid- evant sites in the Iberian Mousterian. S324 Current Anthropology Volume 54, Supplement 8, December 2013

Figure 2. Temporal trends in the technology and raw materials of Trago´, Roca dels Bous, and Cova Gran. LEV p Levallois; DIS p discoid; BHC p bifacial hierarchical centripetal; UF p unifacial; MF p multifacial. A, Relative frequency of main core reduction methods. B, Relative frequency of structured (LEV, BHC, DIS) versus expedient (bipolar, MF, UF) reduction methods. C, Relative frequency of main retouched types. All data from CAϩ Online Supplement B: tables B4, B5. D, Breakdown of raw materials per assemblage. E, Raw material breakdown of quartzite and flint cores. F, Raw material breakdown of quartzite and flint retouched tools. All data from CAϩ Online Supplement B: table B6. A color version of this figure is available in the online edition of Current Anthropology. kyr, assemblages, but a robust radiometric frame has yet to be 29 ע The level TD 10.1 at Atapuerca, dated to337 could represent the earliest evidence of Middle Paleolithic developed. At present, the best documented cultural succes- technology in Iberia (Rodrı´guez 2004). Its large lithic and sion for the latter part of the Middle Pleistocene is that from fossil assemblage shares some elements of continuity with the Bolomor, in eastern Spain. Here, radiometric dates bracket Acheulean (e.g., handaxes), but cores and retouched flakes between OIS 9 and OIS 5e more than a dozen archaeological indicate more standardized systems typical of the Mousterian units in which denticulate and sidescraper-rich layers alternate (Olle´ et al 2013). Cueva Hora´and Cueva del A´ ngel, both in and no handaxes are recorded (Ferna´ndez-Peris 2007). The southern Spain, may potentially contain very old Mousterian earliest levels of Bolomor (XVII–XV) are positioned between de la Torre, Martı´nez-Moreno, and Mora Change and Stasis in the Iberian Middle Paleolithic S325

Figure 3. Geographic locations of the main Middle Paleolithic sites in Iberia. A color version of this figure is available in the online edition of Current Anthropology.

347 and 242 kyr and are considered as early Middle Paleolithic Other pre-OIS 5e sequences in caves/rockshelters present (Ferna´ndez-Peris et al. 2008), which is in agreement with data similar dating problems. Lezetxiki in northern Spain has from Atapuerca TD 10.1. yielded dates 1240 kyr for level VII (Falgue`res, Yokoyama, Both the Bolomor sequence and Atapuerca TD 10.1 point and Arrizabalaga 2005), but most of the other radiometric to an emergence of the Middle Paleolithic earlier than 300,000 ages are inconsistent. Cueva de los Aviones, Cueva de las BP. This would be in agreement with Mousterian-like features Grajas, and Cueva Hora´, all in the southern half of Spain, seen in late Acheulean open-air sites such as Ambrona AS6, contain Mousterian (and in the case of Cueva Hora´ also dated over 350 kyr (Santonja and Villa 2006), but it is at odds Acheulean) layers attributed to the Middle Pleistocene, but with the chronology of other late Acheulean sites in the Man- no reliable radiometric dates have been published. It is likely zanares (e.g., Arriaga, Oxı´geno), Guadiana, and Duero valleys, that several other caves from figure 4 contain pre-OIS 5e for which terminal Middle Pleistocene ages have been pro- Mousterian deposits, but at present, apart from Bolomor, only posed. This calls for either a long coexistence of Acheulean a few yield radiometric dates. Two of those caves are Carihuela and Mousterian technologies between 300 and 100 kyr, a and Trago´ (southern and northeastern Spain, respectively); gradual disappearance of handaxes in early Middle Paleolithic lower deposits from Carihuela range between 146 and 117 sequences as recorded elsewhere (e.g., Moncel et al 2011), kyr (Vega et al. 1997) and show clear Mousterian features, and/or for a need to reconsider the chronology of some of while the bottom of the Middle Paleolithic sequence in Trago´ the central Iberia terraces, which is in reality poorly con- is radiometrically dated to the beginning of the last interglacial strained. (Casanova et al. 2009). Figure 4. Time span of relevant Mousterian sites in Iberia (isotopic curve adapted from Gamble 1986). Given the inconsistencies of radiometric results in a number of assemblages and the lack of dates in several others, this graphic should be considered only as a very preliminary attempt to organize temporally the Iberian Middle Paleolithic. de la Torre, Martı´nez-Moreno, and Mora Change and Stasis in the Iberian Middle Paleolithic S327

Figure 4 could lead us to believe that the majority of Iberian Mousterian become even more difficult for several other rea- cave sites have no Mousterian occupation for most of OIS 5. sons. First, many of the classic studies of the Iberian Middle However, that is likely to be an artefact of dating methods Paleolithic were conducted within the typological approach, (in many sites no techniques other that 14C have been at- and therefore technological data are only available from recent tempted), and research program contingencies (often the bot- publications. Even when technological studies are available, tom of stratigraphy has not yet been reached, and lower, there is a focus on the late Mousterian and its comparison unreported levels might be present). Focusing on the few sites with the early Upper Paleolithic. In consequence, available for which OIS 5 levels are published, some of those already information is biased to the detriment of older, pre-OIS 3 mentioned contain Early Glacial deposits, and several others Mousterian assemblages, for which technological data are of- can also be considered (see fig. 4). Among the latter, El Castillo ten very scarce. There is also a tendency toward qualitative and Cova Negra have yielded relatively consistent radiometric descriptions against quantitative data that sometimes makes dates across thick sequences; in El Castillo, the classic Mous- statistical comparisons unfeasible. More importantly, there is terian is capped between 70 kyr (the age of level 22) and 39 a generalized lack of consistency on the methodology em- kyr (dating proposed for the Early Aurignacian of level 18; ployed to classify knapping methods and technological pat- Rink et al. 1997). Cova Negra contains a thick Mousterian terns; different conceptions exist on the meaning of discoid, stratigraphy spanning the whole of the Early Glacial (OIS Levallois, other centripetal cores, expedient flaking, and so 5d-a), OIS 4, and part of OIS 3. Although radiometrically not forth, and therefore similar reduction methods are named as robust as Cova Negra, the Mousterian sequence of Cueva with different terms. The opposite also occurs, with terms Bajondillo is also constrained between OIS 5 and OIS 3 (Cor- such as “discoid” used to classify cores that could be consid- te´s Sa´nchez et al. 2007), and so is the thick stratigraphy of ered as recurrent Levallois or alike according to recent tech- Covalejos, dated between 100 and 40 kyr (Montes and San- nological conceptions. All of this makes it difficult to con- guino 2005), and Oliveira, dated between 170 and 35 kyr textualize the Pre-Pyrenean diachronic patterns within the (Angelucci and Zilha˜o 2009). Iberian Middle Paleolithic sequence and constrains the num- According to the available dates, most of Mousterian sites ber of case studies that provide comparable data. Further- in Iberia would correspond to the latest part of the Middle more, available data from other case studies elsewhere in Ibe- Paleolithic. While the possibility of a sudden proliferation of ria do not necessarily comprehend every technological aspect, Neanderthal sites after 45 kyr cannot be excluded, it is also which precludes a unified assessment of all analytical criteria very likely that the radiocarbon dating threshold rather than (e.g., raw material selection, tool types, core reduction tech- an actual gap in the archaeological record explains the absence niques, etc.) across the Iberian chronostratigraphic sequence. of longer spans in the OIS 3 assemblages plotted in figure 4. Hence, the patchy character of the available data determines Similarly, the very recent radiocarbon dates obtained in a the comparisons below, which are built on a composite view number of Iberian sites may be due to problems of the limits of a number of Iberian case studies. of AMS 14C; therefore, many of the late Mousterian dates At present, typological classifications of the Iberian Middle from Iberian sites could be seen as minimum ages (e.g., Mar- Paleolithic are unusual, and so the Bordesian taxonomy of tı´nez-Moreno, Mora, and de la Torre 2010). table 1 is limited to data from some 1960s–1980s studies. As Leaving aside chronometric problems, the Iberian record expected, no diachronic patterns are observed in the variation contains some firmly dated sequences that account for con- of Mousterian facies; in Cova Negra, one of the few dated tinuous occupation throughout the first half of OIS 3. Prob- Middle Paleolithic sequences, Quina-type Charentian was rec- ably the most outstanding is Abric Romanı´, which yields well- ognized at the bottom (level XIV: OIS 5d-b) and middle constrained Mousterian levels between 60 and 40 kyr (XIII–XII, X–IX, VII–VI: OIS 4) of the stratigraphy, and the (Bischoff, Julia`, and Mora 1988). Apart from several of the same occurs with the para-Charentian/typical Mousterian, sites mentioned above, Esquilleu, El Salt, Vanguard Cave, and identified both in OIS 4 (level XI, VIII) and at the beginning a few others (see fig. 4) also yield levels 145 kyr overlaid by of OIS 3 (Villaverde 1984). While Cova Negra can be used late Mousterian assemblages. They all attest to thick sequences as an instance of nondirectional temporal variability of Mous- of Middle Paleolithic and, in principle, should enable assessing terian facies, Axlor provides an example of homogeneity of diachronic patterns in Mousterian technology, as discussed in tool types through time; here, all levels are classified as typical the following section. or Quina Charentian (Baldeo´n 1999) despite the thick stra- tigraphy of the rockshelter. Despite reservations derived from the absence of radiometric dates, Axlor and several other Technological Trends in the sequences from table 1 can be placed at the onset of OIS 3 Mousterian of Iberia (see fig. 4). Given the variety of Mousterian facies represented in this time interval, it may be concluded that no temporal If chronostratigraphic correlations are weak because of a lack patterning exists. of radiometric dates (especially for levels beyond the range The relative typological homogeneity of the Iberian Mous- of AMS 14C), interassemblage comparisons of the Iberian terian in comparison with the neighboring sequence of France S328 Current Anthropology Volume 54, Supplement 8, December 2013

Table 1. Mousterian facies in some classic Spanish assemblages

Denticulate Mousterian Charentian Typical Mousterian MTA Mousterian with cleavers Conde 6 Castillo Beta (level 22) Casares Cova Negra V Morı´n 13/14, 15–17 Morı´n 17b2, 11–5 Hornos de la Pen˜a Cova Negra XI Abauntz Castillo Alfa (level 20) La Flecha Za´jara Cova Negra IV–I Pendo XIII Pendo 4 Cova Negra XIV–XII Lezetxiki VII–V Amalda Romanı´ Cova Negra X–IX Amalda VII Lezetxiki V–VI Cova Negra VII–VI Morı´n 17a–17b1, 16–13 Gatzarria Lezetxiki IV Pendo 12–6 Pen˜a Miel Mollet Cueva Milla´n La Ermita Morı´n 12 Pendo 5 Axlor Ermita Eudoviges Sources. Baldeo´n 1993, 1999; Butzer 1981; Cabrera 1984; Freeman 1966; Ripoll and de Lumley 1965; and Villaverde 1984. Note. MTA p Mousterian of Acheulean tradition. has sometimes been explained by limited availability of good ite, the most commonly used rock (table B7), varies through- raw materials in Spain and Portugal. While flint is ubiquitous out the sequence, but again no particular trend is observed around a number of sites in Andalusia and in some parts of (fig. 5). Most of the rocks were procured from a stream bed the Mediterranean, quartzite and other metamorphic rocks 200 m from the cave, and 99% of raw materials were found predominate in large parts of western, central, and northern within a 5-km radius (Manzano et al. 2005), although some Iberia. Despite this unequal distribution of raw materials, we stone tools are reported to come from more distant sources can discuss whether any diachronic patterns exist in the Mid- (Carrio´n et al. 2008). dle Paleolithic procurement strategies. Again, data are not Cova Gran and Trago´present an analogous pattern to the available for all assemblages, so here we selected case studies sites mentioned above; flint was readily available in Cova Gran in which both raw materials and their availability in the land- and Trago´ and consistently preferred in both sequences. The scape have been investigated (CAϩ Supplement B: table B7). case of Trago´ is particularly enlightening: UA3 and UA1 are In the early Middle Paleolithic of TD 10.1, local flint is the ca. 80 kyr apart, and yet raw material procurement for each predominant raw material, but a particular type of exotic flint is remarkably similar (see table B7). Among the case studies is also documented for the first time in the Atapuerca se- selected for figure 5, only Roca dels Bous and Axlor suggest quence (Olle´ et al. 2013), providing yet another element of some divergence; in the case of Roca dels Bous, N10 presents differentiation between the previous Acheulean and TD 10.1. opposite trends to N12 in the use of local (quartzite) versus The thick sequence of Bolomor shows variations in raw ma- imported (flint) rocks. However, the current lack of data for terial procurement (see fig. 5), but there seems to be no tem- underlying levels makes it difficult to ascertain whether such poral patterning, and changes are related to climatic pulses reversal in the use of raw material was episodic or instead during the Middle Pleistocene; small marine flint pebbles were corresponds to a temporal trend in the sequence. With regard preferentially selected when locally available during sea trans- to Axlor, Rı´os (2008) situates most of the flint sources between gressions in detriment of other raw materials such as quartzite 15 and 30 km from the site, whereas other raw material— (accessible in the early part of the sequence) and especially such as quartz, lutite, and so forth—were local (!10 km). limestone, an immediately available rock (Ferna´ndez Peris et Based on data from Baldeo´n (1999), it could then be stated al. 2008). that the sequence of Axlor sees a steady increase of exotic In the case of Abric Romanı´, the location of raw material raw materials (fig. 5). This diachronic pattern could, poten- sources was more fixed, with flint available in a radius of 5– tially, be linked to increasingly larger foraging ranges and/or 10 km and limestone and quartz in the immediate (1 km) more demanding manufacture processes that required higher surrounding of the rockshelter. Although from farther dis- quality raw materials. Time trends have also been detected in tances, flint was consistently preferred in most of the Romanı´ some Portuguese Mousterian sites, but they point in the op- levels (Vaquero 1999). Raw material variation is documented posite direction; according to Zilha˜o (2001), in the sequence throughout the sequence, but figure 5 shows no directionality of Gruta da Oliveira, there is a steady decrease in the use of toward any particular rock type. The Mousterian sequence flint, and Gruta do Caldeira˜o shows a similar trend, which is from El Esquilleu probably spans more than 20 kyr, but sub- related to mobility patterns focused on the exploitation of stantial uniformity in raw material procurement is found local quartzite sources. across the levels (Manzano et al. 2005); percentage of quartz- With some exceptions (e.g., Rı´os 2008; Vaquero et al. 2012), Figure 5. Raw material percentages in selected Iberian Middle Paleolithic assemblages. All data from CAϩ Online Supplement B: table B7. A color version of this figure is available in the online edition of Current Anthropology. S330 Current Anthropology Volume 54, Supplement 8, December 2013 relationships between raw material procurement and other Most of the data on knapping systems correspond to OIS aspects of Neanderthal ecology (e.g., Geneste 1985; Kuhn 4 and especially OIS 3. In the central Mediterranean region, 1995) are yet to be explored in the Iberian Middle Paleolithic, Ferna´ndez-Peris et al. (2008) state that Mousterian assem- so it is not easy to assess the role of other subsistence activities blages of this chronology show predominance of Levallois as in raw material acquisition and how specific foraging strat- opposed to earlier periods. An example is the long sequence egies may affect temporal patterns. Nonetheless, with this of El Salt, dated between 60 and 40 kyr, which shows prev- caution in mind, time trends seem to be an exception rather alence of recurrent centripetal Levallois methods (Galva´n et than the norm; most of the case studies discussed here show al. 2006). A different pattern is reported in Cueva Bajondillo conservative patterns in the acquisition of raw materials, in (southern Spain), where discoid flaking is better represented which fluctuations do exist (not unexpectedly, for in several in the upper levels, albeit Levallois is common across the instances we are dealing with sequences spanning many sequence (Corte´s Sa´nchez 2008). thousands of years), but such variations may be explained by Detailed reports of flaking techniques are now available for environmental constraints (e.g., Bolomor) or contingent set- a number of north Iberian OIS 4/3 assemblages. In Catalunya, tlement dynamics (Roca dels Bous, Romanı´) rather than by discoid and discoid-like (e.g., BHC) methods are omnipresent directional trends toward local or exotic raw materials. in the Middle Paleolithic as opposed to lower frequencies of After excluding the existence of diachronic trends in ty- Levallois (Mora 1988). Although Cantabria follows a similar pological and raw material patterns, we shall now discuss pattern, this region shows higher variability than Catalunya, whether patterns in the variability of flaking methods can be which is probably explained by the larger number of sites traced. As mentioned above, this issue is particularly difficult documented. Carrio´n et al. (2008) state that although Leval- to address; one reason is that disparity in the conceptions of lois is not abundant, it is present in most of Cantabrian sites, Levallois, discoid, and other centripetal methods obscure po- usually in the recurrent centripetal modality. Discoid cores tential interassemblage comparison of structured flaking predominate and are often made on flakes (Carrio´n et al. methods. On the other hand, a fairly large number of available 2008), and Kombewa flaking is also attested (e.g., Rı´os 2008). studies focus on such structured techniques, and simpler, Quina flaking is now identified in El Esquilleu (Carrio´n et al. more expedient flaking solutions are frequently excluded from 2008), Axlor, Gatzarri, Lezetxiki, and Amalda (Rı´os 2008), the analysis. While consideration of Levallois and discoid be- but bearing in mind the recentness of the definition of this comes on occasion sophistic and might not help to discern core reduction technique (Bourguignon 1997), it will be un- general patterns (de la Torre 2009), differentiation between surprising if over the next few years Quina flaking is docu- structured/long reduction sequences and expedient/short se- mented elsewhere. Bladelet production is suggested at Cueva ries cores (Casanova et al. 2009) may provide a more effective Morı´n, El Castillo, and Covalejos (Bernaldo de Quiro´s,Sa´n- ground for time-depth comparisons. Such a comparative chez-Ferna´ndez, and Maı´llo 2010) within Mousterian levels frame, nonetheless, is only available for a few sequences, so where no traces of Upper Paleolithic are reported. discussion will be restrained here to some particular case stud- Indistinctive Quina and discoid flaking are documented in ies in which comparable technological data exist. quartzite and flint, while Levallois is consistently made in the Discoid-like methods are known in Iberia since the Lower highest quality raw materials available (Carrio´n et al. 2008). Pleistocene; Vaquero and Carbonell (2003) consider the re- Beyond recurrence of this pattern, and despite the wealth of current bifacial centripetal method of Atapuerca TD 6 to be Mousterian assemblages in the Cantabrian region, no clear fairly similar to discoid flaking and report an increase of this correlations are yet available between knapping methods and technique throughout the sequence. According to Olle´etal. other contextual elements. A particularly telling recent ex- (2013), centripetal strategies progressively become more stan- ample illustrates the impenetrability of the meaning of flaking dardized, leading to the Levallois-like technique from Upper methods variability; at El Sidro´n, the lithic assemblage was TD 10.1 (OIS 9), in which morphometrical predetermination aimed at one very specific activity, that is, butchering other is observed. The roughly contemporary level XVII of Bolomor Neanderthals. Yet both discoid and Levallois cores were flaked contains poor evidence of Levallois, while in upper levels such in this single and task-specific event (Santamarı´a et al. 2010). as XII–VII (OIS 6), hierarchization of cores (recurrent cen- While acknowledging pitfalls derived from the disparity of tripetal Levallois and/or hierarchical discoid) is attested (Fer- core classification systems and chronometric problems of as- na´ndez Peris et al. 2008). In OIS 5e, Bolomor levels VI–I semblages, we have compiled a few of the available case studies contain discoid and Levallois flaking accompanied by trifacial in order to assess diachronically north Iberia Mousterian and Kombewa cores (Ferna´ndez Peris et al. 2008). Trago´UA3, knapping methods (CAϩ Supplement B: table B8). Some with a similar chronology, shows predominance of expedient sequences such as Gabasa indicate no change in primary flak- methods followed by bifacial hierarchical centripetal (BHC) ing techniques. Abric Romanı´does show diachronic pattern- reduction, the latter being potentially similar to the recurrent ing toward higher frequency of Levallois in the upper levels, bifacial centripetal method of Atapuerca (Vaquero and Car- whereas variations in El Esquilleu are nondirectional: discoid bonell 2003) and to many of the cores described as discoid dominates the bottom and the top of the sequence, while and Levallois in Bolomor. Quina and Levallois methods are the commonest in some of de la Torre, Martı´nez-Moreno, and Mora Change and Stasis in the Iberian Middle Paleolithic S331 the middle levels. Therefore, no clear patterns emerge from Technological Variability of the Iberian the diachronic assessment of intrasite variation. Middle Paleolithic in the Context of With regard to intersite variability, figure 6 attempts to Southwestern Europe order chronologically the main flaking systems in the OIS 4– The case study of the Middle Paleolithic at La Noguera region OIS 3 case studies (see also table B8). Late OIS 4/early OIS has been presented above to examine possible time trends in 3 assemblages such as the lower levels of El Esquilleu, Abric the technology of the Iberian Mousterian. Our results suggest Romanı´, and (perhaps) Gabasa show predominance of discoid that albeit changes occur, they do not show directional pat- methods. If we were to assume that dates at the limit of the terns. Framing the three Pre-Pyrenean case studies within the radiocarbon method are correct, then the trend would be wider context of the Iberian Mousterian, although hindered more variable in the 45–39 kyr interval; Levallois predomi- by methodological and empirical constraints, seems to suggest nates in El Castillo level 20e, top levels of Abric Romanı´, that the lack of diachronic patterning in technological strat- Axlor N, and Arrillor level emj. Nonetheless, in a similar egies is applicable to most of the Portuguese and Spanish chronological span, discoid and discoid-like (e.g., BHC) tech- Middle Paleolithic. In truth, this is hardly surprising and niques are the primarily flaking methods in Gabasa, Trago´ comes to confirm on technological grounds what typological UA1, Roca dels Bous level N12, and Morı´n levels 11 and 13, approaches (e.g., Bordes 1972) had long reported; that is, a while Quina predominates in El Esquilleu levels XIII–XVI and substantial part of western European Middle Paleolithic as- Axlor B–D. The pattern is not clearer in post-39-kyr assem- semblages show no defined time trends. blages; Quina, Levallois, and discoid follow one another in However, it is important to stress that absence of temporal the El Esquilleu top levels, while BHC flaking predominates directionality does not mean randomness of variation. In- in Cova Gran S1D–S1C and unifacial methods in Cova Gran trasite variability in Roca dels Bous (Martı´nez-Moreno, Mora, S1B and Roca dels Bous N10. In short, figure 6 certifies for and de la Torre 2004; Mora, de la Torre, and Martı´nez-Moreno technical systems the same pattern (or rather, the lack of any) 2004), Bolomor (Ferna´ndez Peris 2007), Romanı´ (Vaquero discussed above for typological facies and raw material pro- 1999), Axlor (Rı´os 2008), El Esquilleu (Baena et al. 2005), curement: an absence of directional changes or temporal and others may respond to alternative settlement dynamics trends. potentially related to distinct foraging strategies by Neander-

Figure 6. Chronostratigraphic position of flaking techniques in the OIS 4/3 assemblages from CAϩ Online Supplement B: table B8. S332 Current Anthropology Volume 54, Supplement 8, December 2013 thal groups. A challenge in current studies of intrasite vari- Pyrenees (Isturitz, Abri Olha, Calavante´, and Noisetier) ability is to decipher why changing technological behaviors Mousterian assemblages. Although subsequent studies (e.g., are documented in contexts where raw material sources were Cabrera, Pike-Tay, and Bernaldo de Quiro´s 2004; Freeman fixed and biotic resources would have been similar through 1966) excluded Bordes’s original proposal of the Vasconian time; that will surely help explain the apparently random as a distinct facies, new hypotheses (e.g., Thie´bault et al. 2012) changes detected in some archaeological sequences. somehow rescue the original idea and propose a specific tech- Once methodological disparities are overcome and the se- nological entity for the group of Mousterian sites with cleav- rious chronometric problems are addressed, interassemblage ers. Also, denticulates predominate in some Cantabrian and comparisons may also play a more important role in modern many Catalonian assemblages, which is not a common pattern analysis of Iberian Middle Paleolithic technological strategies elsewhere in Iberia. This prevalence of denticulate assem- and in the recognition of temporal trends. The Iberian record blages, especially in Catalonian sites, has been known for a has the required time depth to explore more than 250 kyr of long time (Mora 1988; Ripoll and de Lumley 1965) and has Mousterian history, and it is unfortunate that substantial parts also recently been proposed as sharing traits with French Py- of this research focus only on the Neanderthal extinction renean assemblages (Thie´bault et al. 2012). Albeit further cor- interval. Also, the absence of time trends and the attribution relations and interassemblage comparative work is required, of intrasite variability to contingent settlement strategies do these examples seem to suggest that regional variability in not preclude the existence of other variables of differentiation Iberia could exist. in the Mousterian of Iberia with respect to the adjacent region For the moment, however, it is difficult to compare data of France and within Iberia itself. Traditionally attributed to directly on temporal and regional variation from Iberia with raw material constraints, the scarcity of preferential flake Le- neighboring regions such as France. In the latter, the high vallois cores in Iberia is noticeable; they certainly appear in density of sites, development of a reliable radiometric frame- some Middle Paleolithic sites, but they are much less common work, and consistency of technological study and publication than Levallois recurrent centripetal methods. Predominance of site reports has enabled researchers to construct a solid of recurrent centripetal techniques is a feature shared with chronostratigraphic sequence where time trends in technology the Mousterian of the Aquitaine basin, but while in the French can be evaluated more precisely. Technological reviews began sequence some evolution within Levallois methods is docu- in the wake of the 1960s–1980s typological debate on the mented (Turq 2000), such variation is not identified in Iberia. meaning of Middle Paleolithic variability, and even in recent Turq (2000) also reports numerous Kombewa flakes both for years (e.g., Delagnes, Jaubert, and Meignen 2007) attempts making handaxes and cleavers and as core blanks. While have been made to adequate the Mousterian facies to the Kombewa is not yet much reported in Iberian assemblages, currently prevalent technological reading of assemblages. the role of flakes as cores is increasingly recognized; as detailed Thus, a meaningful association is proposed to exist between by Bourguignon, Faivre, and Turq (2004) across French sites, the typological facies of La Ferrassie Mousterian and the Le- such “ramification” of chaıˆnes ope´ratoires may explain the vallois method, the Denticulate Mousterian and discoid flak- remarkably small size of stone tools in some Mousterian as- ing, the Quina Mousterian facies and Quina reduction, and semblages from both the north (Rı´os 2008) and south (Corte´s the typical Mousterian of Levallois facies and Levallois re- Sa´nchez 2007) ends of the Iberian Peninsula. It is unlikely, current flaking (Delagnes, Jaubert, and Meignen 2007). however, that this so-called ramification explains all instances Since the beginning of the chaıˆne ope´ratoire approach in of small-sized assemblages, for in some cases, such as Cuesta the 1980s, certain diachronic patterns in technology were dis- de la Bajada (Santonja et al 2000) and Bolomor (Ferna´ndez cerned, particularly with regard to the evolution of the Le- Peris 2007), raw material constraints may be responsible, vallois technique (Geneste 1990). In the Aquitaine basin, Turq while in others (e.g., Roca dels Bous) the small size of artefacts (2000) reports a change from unidirectional and bidirectional is due to intentional extreme exhaustion of nodular cores recurrent Levallois in pre-OIS 5 assemblages to centripetal (Mora, de la Torre, and Martı´nez-Moreno 2004). recurrent Levallois in Last Glacial sites. He also states that Regional variability of tool types should also be investi- from OIS 5, centripetal methods dominated alongside Kom- gated. In general, the diversity of retouched tools seems to bewa and Quina flaking. According to Delagnes and Meignen be lower in Iberia than in southwestern France. Levallois and (2006), preferential Levallois is consistently older and com- Mousterian points, backed knives, and others are present in moner in the north than in southern France, and centripetal a number of Iberian Mousterian sites but show even lower recurrent Levallois became dominant after OIS 5. The Rhone frequencies than in southwestern France. Sidescrapers dom- Valley (Moncel and Daujeard 2012) offers a different picture, inate most Iberian sites apart from Catalunya and some Can- however, in which Levallois is mostly centripetal during OIS tabrian sites, and such sidescraper-rich assemblages are often 8 while unidirectional and bidirectional Levallois cores pre- considered as “typical” Mousterian. Cantabria and Catalunya dominate in OIS 4 and 3. Actually, the OIS 8–OIS 3 sequence seem to present some idiosyncrasies. The presence of cleavers at the Rhone Valley (Moncel and Daujeard 2012:113) some- is well documented in some Cantabrian coast (El Castillo, what resembles the random variation of flaking systems and Morı´n, Lezetxiki, Pendo, Amalda, and Gatzarria) and French tool-type frequencies discussed above for Iberia, showing sto- de la Torre, Martı´nez-Moreno, and Mora Change and Stasis in the Iberian Middle Paleolithic S333 chastic predominance of Levallois and discoid methods and ability of blanks and low mobility patterns, link the MTA with nondirectional changes in the percentage of scrapers and high mobility Neanderthal groups, and see discoid/denticu- points. late-dominated assemblages as the result of multipurpose and Other technological indicators may potentially bear clearer highly versatile tool kits. Scott and Ashton (2011), however, chronological meaning. In northern France, blade production propose for early Levallois assemblages exactly the opposite is limited to the early OIS 5 (Delagnes and Meignen 2006) and consider that Levallois represents increased transport of and does not appear in the Rhone Valley before the last in- blanks and extended curation. Likewise, whereas some pro- terglacial, although in southeastern France, blade assemblages pose that MTA bifaces were multipurpose tools (e.g., Soressi continues after the early OIS 5 (Moncel and Daujeard 2012). 2004), others see these as single-tasked artefacts related to Blade and/or bladelet production seem also to be character- butchery made by highly mobile Neanderthals (Delagnes and istic of some late Mousterian assemblages; elongated blanks Rendu 2011). in the southwestern France Mousterian of Acheulean tradition Beyond interpretive problems of interassemblage variabil- (MTA) could be a precursor for the Chatelperronian blade ity, it remains clear that the French record contains a solid technology (Pelegrin and Soressi 2007), while in the southeast and reliable chronostratigraphic record and that such a record the Charentian would evolve into blade and bladelet industries could indicate some kind of time patterning in Mousterian with a marked regional character (Slimak 2008). technology (Delagnes, Jaubert, and Meignen 2007; Delagnes In summary, evidence seems to suggest that some time and Meignen 2006; Delagnes and Rendu 2011). For the mo- trends exist in the Middle Paleolithic technology of France. ment, such trends are particularly conspicuous in the later Recent overviews of the Aquitaine record report a succession part of the Mousterian and could be related to a process of of the predominance of flaking systems, with prevalence of regionalization, particularly in the south of France (Slimak Levallois and laminar reduction in the OIS 7–OIS 5 sites 2008). Regionalization phenomena are seen in adjacent followed by Quina, MTA, and discoid/denticulate assemblages regions of Europe at the end of the Middle Paleolithic not from OIS 4 until the transitional industries to the Upper only on typological grounds but also according to techno- Paleolithic (Delagnes and Rendu 2011). Changes within the logical indicators, such as in Italy (Kuhn 2006). However, this MTA are also given a chronological significance, with MTA temporal trend has yet to be discerned in the Iberian Pen- type A assemblages being consistently older than the MTA insula, where a number of challenges discussed in this paper type B (Soressi 2004), and !50 kyr sites from southeast France confound any potential patterns. When compared with the are reported to undergo a process of microlithization and neighboring area of France, the Iberian Middle Paleolithic typological specialization (Slimak 2008). record shows more technotypological homogeneity, both re- Although the transitional industries to the Upper Paleo- gionally and diachronically. In order to ascertain whether this lithic are beyond the scope of this paper, it is worth noting lack of variability in the Iberian Mousterian is real or an that most indicators of Mousterian variability in France are artefact of empirical and methodological problems, further clustered in the later Middle Paleolithic. Delagnes and Meig- efforts are required to develop a chronostratigraphic and tech- nen (2006) report higher diversification of flaking systems in nological data set framework comparable with that available later stages of the Mousterian in which production of blanks in other parts of western Europe and the Near East. with a low degree of predetermination predominates. Idio- syncratic industries such as the MTA seem to be mostly con- Conclusions strained to post-50-kyr sequences (Soressi 2004), and a similar time trend applies to the appearance of new tool types and The debate on change versus stasis is inseparable from the blade/bladelet production in the Rhone Valley (Slimak 2008). assessment of Mousterian technology. Two questions can be Therefore, there seems to be some kind of “shift of gear” that asked here (Kuhn 2006). First, did major technological in- accelerates technological change in the !50-kyr Mousterian novations occur during the Middle Paleolithic? Many would French sequence, resulting in a more conspicuous interas- agree with Kuhn (2006) that the whole package of techno- semblage variability. Such variability certainly bears chrono- logical features documented in the latest Mousterian is already logical connotations, potentially regional differentiation, and present during the early Middle Paleolithic. Second, and a putatively also cultural evolutionary implications, given the different issue, does Mousterian variability shows directional temporal proximity to the early Upper Paleolithic. trends? With some exceptions (e.g., Mellars 1969), the ty- Causes of these general temporal trends have also been pological approach failed to detect such temporal patterns in investigated. For example, recent studies have aimed to es- the Middle Paleolithic repertoire. tablish links between particular knapping systems and sub- In recent years, technological perspectives have also ex- sistence strategies, providing an explanatory cause for chro- plored the existence of diachronic trends but from the view nological patterns. But so far results are contradictory; of the knapping systems. For example, Scott and Ashton assuming an association between flaking methods and mo- (2011) argue that most of early Middle Paleolithic sites in bility patterns, Delagnes and Rendu (2011) relate predomi- Northern Europe show predominance of preferential Levallois nance of Levallois and blade production with low transport- methods that are not so conspicuous in later assemblages. In S334 Current Anthropology Volume 54, Supplement 8, December 2013 the same vein, laminar technologies are reported to be more References Cited common in earlier Middle Paleolithic assemblages (Bar-Yosef Angelucci, Diego E., and Joao Zilha˜o. 2009. Stratigraphy and formation pro- and Kuhn 1999). In general, the French sequence seems to cesses of the Upper Pleistocene deposit at Gruta da Oliveira, Almonda present at the end of the Mousterian greater diversity of flak- Karstic System, Torres Novas, Portugal. Geoarchaeology 24:277–310. Baena, Javier, E. Carrio´n, B. Ruiz, B. Ellwood, C. Sese´, J. Yravedra, J. Jorda´, ing methods coexisting at the same time and larger frequency et al. 2005. Paleoecologı´a y comportamiento humano durante el Pleistoceno of lower predetermination systems (Delagnes and Meignen Superior en la comarca de Lie´bana: la secuencia de la Cueva del Esquilleu, 2006). Occidente de Cantabria, Espan˜a. In Neandertales canta´bricos: estado de la cuestio´n. Jose A. Lasheras and Ramon Montes Barquı´n, eds. Pp. 461–487. Can we apply such observations to the Iberian record? This Monografı´as Museo de Altamira 20. Madrid: Ministerio de Cultura, Sub- paper has argued that according to the evidence currently direccio´n General de Publicaciones, Informacio´n y Documentacio´n. available, time trends either do not exist or are not discernible Baldeo´n, Amelia. 1993. El yacimiento de Lezetxiki (Gipuzkoa, Paı´s Vasco): los niveles musterienses. Munibe 45:3–97. because of a lack of comparable data. We acknowledge that ———. 1999. El abrigo de Axlor (Bizkaia, Paı´s Vasco): las industrias lı´ticas the La Noguera Middle Paleolithic does not provide enough de sus niveles Musterienses. Munibe 51:9–121. Bar-Yosef, Ofer, and Steven L. Kuhn. 1999. The big deal about blades: laminar time depth and sufficient sites to build up a solid sample. It technologies and human evolution. American Anthropologist 10:322–338. should also be remembered that chronostratigraphic and Bernaldo de Quiro´s, Federico, Granada Sa´nchez-Ferna´ndez, and Jose M. methodological problems are too acute to allow drawing firm Maı´llo. 2010. Technological characteristics at the end of the Mousterian in Cantabria: the El Cı´astillo and Cueva Morı´n (Spain). In The Upper Pa- conclusions on interassemblage correlations for the whole of laeolithic revolution in global perspective: papers in honour of Sir Paul Mellars. Iberia. Therefore, this paper should be seen as an attempt to Katherine V. Boyle, Clive Gamble, and Ofer Bar-Yosef, eds. Pp. 153–160. speculate about the diachronic potential of the Iberian Middle Cambridge: McDonald Institute. Beyries, Sylvia. 1988. Functional variability of lithics sets in the Middle Pa- Paleolithic rather than a categorical exclusion of the existence leolithic. In Upper Pleistocene prehistory of western Eurasia. Harold L. Dibble of the time patterns documented elsewhere in western Europe. and Anta Montet-White, eds. Pp. 213–223. Philadelphia: University Mu- seum, University of Philadelphia. As stated in an earlier section, it would be unsurprising if Binford, Lewis R. 1973. Interassemblage variability: the Mousterian and the such patterns are detected once the Iberian record is organized functional argument. In The explanation of culture change: models in pre- in a more reliable chronological framework and technological history. Colin Renfrew, ed. Pp. 227–254. London: Duckworth. Bischoff, James L., Ramon Julia`, and Rafael Mora. 1988. Uranium series dating studies become readily available. of the Mousterian occupation at Abric Romanı´, Spain. Nature 332:68–70. For now, however, directionality of assemblage variability Boe¨da, Eric. 1986. Approche technologique du concept Levallois et e´valuation is not visible in the Iberian Middle Paleolithic. Quoting Kuhn de son champ d’application. PhD dissertation, Universite´Paris X Nanterre. Bordes, Franc¸ois. 1972. A tale of two caves. New York: Harper & Row. (2006:110), “While the Mousterian may have changed over Bourguignon, Laurence. 1997. Le Mouste´rien de type Quina: nouvelle de´fin- time it was not going anywhere in particular,” Iberia included. ition d’une entite´technique. PhD dissertation, Universite´Paris X Nanterre. Bourguignon, Laurence, Jean-Philippe Faivre, and A. Turq. 2004. Ramification This does not mean that it is unworthy to keep trying to des chaıˆnes ope´ratoires: une spe´cificite´ du Mouste´rien? Pale´o 16:37–48. disentangle the long historical trajectory of the Mousterian. Butzer, Karl W. 1981. Cave Sediments, Upper Pleistocene stratigraphy and As pointed out above, most of current efforts in Mousterian Mousterian facies in Cantabrian Spain. Journal of Archaeological Science 8: 133–183. research in Spain and Portugal are privileging only the tran- Cabrera, Victoria. 1984. El yacimiento de la cueva de “El Castillo” (Puente sition to the Upper Paleolithic. This is definitely not unique Viesgo, Santander). Bibliotheca Praehistorica Hispana, vol. 22. Madrid: to Iberia, and it has been argued that also elsewhere “what is Consejo Superior de Investigaciones Cientı´ficas, Instituto Espan˜ol de Pre- historia. almost never addressed is what was going on earlier in the Cabrera, Victoria, A. Pike-Tay, and F. Bernaldo de Quiro´s. 2004. Trends in Middle Paleolithic, before modern humans and the Upper Middle Paleolithic settlement in Cantabrian Spain: the Late Mousterian at Castillo Cave. In Settlement dynamics of the Middle Paleolithic and Middle Paleolithic came on the scene” (Kuhn and Hovers 2006:3). Stone Age, vol. 2. Nicholas J. Conard, ed. Pp. 437–460. Tu¨bingen: Kerns. Only further research can help to address this problem. Carrio´n, Elena, J. Baena, C. Conde, F. Cuartero, and M. Roca. 2008. Varia- bilidad tecnolo´gica en el musteriense de Cantabria. In Variabilidad te´cnica del Paleolı´tico Medio en el sudoeste de Europa, vol. 14. Rafael Mora, Jorge Martı´nez-Moreno, Ignacio de la Torre, and Joel Casanova, eds. Pp. 279– 318. Barcelona: Treballs d’Arqueologia. Casanova, Joel. 2009. Estrategias tecnolo´gicas de los neandertales en la ver- tiente sur del Pirineo oriental. PhD dissertation, Universidad Autonoma Acknowledgments de Barcelona. Casanova, Joel, R. Mora, J. Martı´nez-Moreno, and I. de la Torre. 2009. Stra- We thank Erella Hovers, Steven Kuhn, and Leslie Aiello for te´gies techniques dans le Pale´olithique Moyen du sud-est des Pyre´ne´es. the invitation to participate in the “Alternative Pathways to L’Anthropologie 113:313–340. Complexity” Wenner-Gren symposium and to contribute to Corte´s Sa´nchez, Miguel, ed. 2007. Cueva Bajondillo (Torremolinos): secuencia cronocultural y paleoambiental del Cuaternario reciente en la Bahı´a de Ma´laga. this special issue of Current Anthropology. Comments by the Ma´laga: Diputacio´n de Ma´laga. guest editors and two anonymous reviewers on an earlier draft ———. 2008. Variabilidad tecnolo´gica en el Paleolı´tico Medio meridional of this paper are gratefully acknowledged. Excavations in the ibe´rico: un punto de partida. In Variabilidad te´cnica del Paleolı´tico Medio en el sudoeste de Europa, vol. 14. Rafael Mora, Jorge Martı´nez-Moreno, Pre-Pyrenees Middle Paleolithic are funded by the Spanish Ignacio de la Torre, and Joel Casanova, eds. Pp. 105–119. Barcelona: Treballs Ministerio de Ciencia e Innovacion (HAR2010-15002). R. d’Arqueologia. Delagnes, Anne, Jacques Jaubert, and Liliane Meignen. 2007. Les technocom- Mora is grateful for the support of the Institucio´ Catalana de plexes du Pale´olithique moyen en Europe occidentale dans leur cadre diach- Recerca i Estudis Avanc¸ats. ronique et ge´ographique. In Les ne´andertaliens, biologie et cultures. Bernard de la Torre, Martı´nez-Moreno, and Mora Change and Stasis in the Iberian Middle Paleolithic S335

Vandermeersch and Bruno Maureille, eds. Pp. 213–229. Paris: Editions du dertals and modern humans in western Asia. Takeru Akazawa, Kenichi Aoki, Comite´ des Travaux Historiques et Scientifiques. and Ofer Bar-Yosef, eds. Pp. 205–221. New York: Plenum. Delagnes, Anne, and Liliane Meignen. 2006. Diversity of lithic production Hovers, Erella, and Anna Belfer-Cohen. 2013. On variability and complexity: systems during the Middle Paleolithic in France: are there any chronological lessons from the Levantine Middle Paleolithic record. Current Anthropology trends? In Transitions before the transition: evolution and stability in the 54(suppl. 8):S337–S357. Middle Paleolithic and Middle Stone Age. Erella Hovers and Steven. L. Kuhn, Jaubert, Jacques. 2011. Arche´ose´quences du Pale´olithique moyen du sud-ouest eds. Pp. 85–107. New York: Springer. de la France: quel bilan un quart de sie`cle apre`s Franc¸ois Bordes? In Franc¸ois Delagnes, Anne, and Will Rendu. 2011. Shifts in Neandertal mobility, tech- Bordes et la pre´histoire . Franc¸ois Delpech and Jacques Jaubert, eds. Pp. 235– nology and subsistence strategies in western France. Journal of Archaeological 253. Paris: Editions du Comite´ des Travaux Historiques et Scientifiques. Science 38:1771–1783. Kuhn, Steven L. 1995. Mousterian lithic technology: an ecological perspective. de la Torre, Ignacio. 2009. Technological Strategies in the Lower Pleistocene Princeton, NJ: Princeton University Press. at Peninj (west of Lake Natron, Tanzania). In The cutting edge: new ap- ———. 2006. Trajectories of change in the Middle Paleolithic of Italy. In proaches to the archaeology of human origins. Kathy Schick and Nicholas Transitions before the transition: evolution and stability in the Middle Pale- Toth, eds. Pp. 93–113. Bloomington, IN: Stone Age Institute. olithic and Middle Stone Age. Erella Hovers and Steven L. Kuhn, eds. Pp. de la Torre, Ignacio, and Rafael Mora. 2004. El Olduvayense de la seccio´n tipo 109–120. New York: Springer. de Peninj (Lago Natron, Tanzania). Monografies del CEPAP 1. Barcelona: Kuhn, Steven L., and Erella Hovers. 2006. General introduction. In Transitions Centre d’estudis del patrimoni arqueolo´gic de la prehistoria. before the transition: evolution and stability in the Middle Paleolithic and d’Errico, Francesco, and Maria F. Sa´nchez-Gon˜i. 2003. Neandertal extinction Middle Stone Age. Erella Hovers and Steven L. Kuhn, eds. Pp. 1–11. New and the millennial scale climatic variability of OIS 3. Quaternary Science York: Springer. Reviews 22:769–788. Laville, Henri. 1973. The relative position of Mousterian industries in the Dibble, Harold L. 1987. The interpretation of Middle Paleolithic scraper mor- climatic chronology of the Early Wurm in the Perigord. World Archaeology phology. American Antiquity 52:109–117. 4:323–329. Dibble, Harold L., and Nicholas Rolland. 1992. On assemblage variability in Manzano, Ivan, J. Baena, A. La´zaro, D. Martı´n, L. Dapena, M. Roca, and E. the Middle Paleolithic of western Europe: history, perspectives, and a new Moreno. 2005. Ana´lisis de los recursos lı´ticos en la Cueva del Esquilleu: synthesis. In The Middle Paleolithic: adaptation, behaviour and variability. gestio´n y comportamiento durante el Musteriense (Comarca de la Lie´bana, Harold L. Dibble and Paul Mellars, eds. Pp. 1–28. Philadelphia: University Occidente de Cantabria). In Neandertales canta´bricos: estado de la cuestio´n. Museum, University of Philadelphia. Jose A. Lasheras and Ramon Montes, eds. Pp. 285–300. Monografı´as Na- Falgue`res, Christophe, Y. Yokoyama, and A´ . Arrizabalaga. 2005. La geocron- cional y Centro de Investigacio´n de Altamira 20. Madrid: Ministerio de ologı´a del yacimiento pleistoce´nico de Lezetxiki (Arrasate, Paı´s Vasco): crı´- Cultura, Subdireccio´n General de Publicaciones, Informacio´n y Documen- tica de las dataciones existentes y algunas nuevas aportaciones. Munibe 57: tacio´n. 93–106. Maroto, Julia, M. Vaquero, A. Arrizabalaga, J. Baena, E. Baquedano, J. Jorda´, Ferna´ndez Peris, Josep. 2007. La Cova del Bolomor (Tavernes de la Valldigna, R. Julia, et al. 2012. Current issues in Late Middle Palaeolithic chronology: new assessments from northern Iberia. Quaternary International 247:15– Valencia): las industrias del Pleistoceno medio en el a´mbito del Mediterra´neo 25. peninsular. Serie de Trabajos Varios, no. 108. Valencia: Servicio de Inves- Martı´nez-Moreno, Jorge, Ignacio de la Torre, Rafael Mora, and Joel Casanova. tigacio´n Prehisto´rica, Diputacio´n Provincial de Vale`ncia. 2010. Technical variability and changes in the pattern of settlement at Roca Ferna´ndez Peris, Josep, V. Barciela, R. Blasco, F. Cuartero, and P. San˜udo. dels Bous (southeastern Pre-Pyrenees, Spain). In Settlement dynamics of the 2008. El Paleolı´tico Medio en el territorio valenciano y la variabilidad tecno- Middle Paleolithic and Middle Stone Age, vol. 3. Nicholas J. Conard and econo´mica de la Cova del Bolomor. In Variabilidad te´cnica del Paleolı´tico Anne Delagnes, eds. Pp. 485–507. Tu¨bingen: Kerns. Medio en el sudoeste de Europa, vol. 14. Rafael Mora, Jorge Martı´nez- Martı´nez-Moreno, Jorge, Rafael Mora, and Ignacio de la Torre. 2004. Meth- Moreno, Ignacio de la Torre, and J. Casanova, eds. Pp. 141–169. Barcelona: odological approach for understanding Middle Palaeolithic settlement dy- Treballs d’Arqueologia. namics at La Roca dels Bous (Noguera, Catalunya, Northeast Spain). In Finlayson, Clive, F. Giles, J. Rodrı´guez-Vidal, D. A. Fa, J. M. Gutie´rrez, A. S. Settlement dynamics of the Middle Paleolithic and Middle Stone Age,vol.2. Pe´rez, G. Finlayson, et al. 2006. Late survival of Neanderthals at the south- Nicholas J. Conard, ed. Pp. 393–413. Tu¨bingen: Kerns. ernmost extreme of Europe. Nature 443:850–853. ———. 2010. The Middle-to-Upper Palaeolithic transition in Cova Gran Freeman, Leslie G. 1966. The nature of Mousterian facies in Cantabrian Spain. (Catalonia, Spain) and the extinction of Neanderthals in the Iberian Pen- American Anthropologist 68:230–237. insula. Journal of Human Evolution 58:211–226. Galva´n, Bertila, C. M. Herna´ndez, M. I. Francisco, and A. Rodrı´guez. 2006. Mellars, Paul. 1969. The chronology of Mousterian industries in the Perigord Datos para la caracterizacio´n del final del musteriense en los valles de Alcoi. region of south-west France. Proceedings of the Prehistoric Society 35:134– In En el centenario de la cueva de El Castillo: el ocaso de los Neandertales. 171. Victoria Cabrera, Federico Bernaldo de Quiro´s, and Jose M. Maı´llo, eds. Moncel, Marie-He´le`ne, and Camille Daujeard. 2012. The variability of the Pp. 127–141. Madrid: Centro Asociado a la Universidad Nacional de Ed- Middle Palaeolithic on the right bank of the Middle Rhoˆne Valley (southeast ucacio´n a Distancia en Cantabria. France): technical traditions or functional choices? Quaternary International Gamble, C. 1986. The Palaeolithic settlement of Europe. Cambridge: Cambridge 247:103–124. University Press. Moncel, Marie-He´le`ne, Anne-Marie Moigne, Youssef Sam, and Jean Combier. Geneste, Jean-Michel. 1985. Analyse lithique d’industries mouste´riennes du 2011. The emergence of Neanderthal technical behavior: new evidence from Pe´rigord: une approche technologique du comportement des groupes hu- Orgnac 3 (level 1, MIS 8), southeastern France. Current Anthropology 52: mains au Pale´olitique Moyen. Bordeaux: PhD dissertation, Universite´de 37–75. Bordeaux I. Montes, Ramon, and Juan Sanguino. 2005. Nuevos datos para el conocimiento ———. 1990. De´veloppement des syte`mes de production lithique au cours del Paleolı´tico Medio en el centro de la Regio´n Canta´brica: la Cueva de du Pale´olithique moyen en Aquitaine septentrionale. In Pale´olithique moyen Covalejos. In Neandertales canta´bricos: estado de la cuestio´n. J. A. Lasheras et re´cent et Pale´olithique supe´rieur ancien en Europe: ruptures et transitions: and R. Montes, eds. Pp. 489–538. Monografı´as Nacional y Centro de In- examen critique des documents arche´ologiques: actes du colloque international vestigacio´n de Altamira 20. Madrid: Ministerio de Cultura, Subdireccio´n de Nemours, 9–10 mai 1988. Catherine Farizy, ed. Pp. 203–213. Nemours: General de Publicaciones, Informacio´n y Documentacio´n. Me´moires du Muse´e de Pre´histoire d’Ile-de-France. Mora, Rafael. 1988. El Paleolı´tico Medio en Catalunya. PhD dissertation, Gonza´lez Urquijo, Jesus E., J. J. Iba´n˜ez, J. Rı´os, and L. Bourguignon. 2006. Universidad de Barcelona. Aportes de las nuevas excavaciones en Axlor sobre el final del Paleolı´tico Mora, Rafael, A. Benito-Calvo, Jorge Martı´nez-Moreno, P. Gonza´lez, and I. Medio. In En el centenario de la cueva de El Castillo: el ocaso de los Nean- de la Torre. 2011. Chrono-stratigraphy of the Upper Pleistocene and Ho- dertales. Victoria Cabrera, Federico Bernaldo de Quiro´s, and Jose M. Maı´llo, locene archaeological sequence in Cova Gran (south-eastern Pre-Pyrenees, eds. Pp. 269–289. Madrid: Centro Asociado a la Universidad Nacional de Iberian Peninsula). Journal of Quaternary Science 26:635–644. Educacio´n a Distancia en Cantabria. Mora, Rafael, Ignacio de la Torre, and Jorge Martı´nez-Moreno. 2004. Middle Goren-Inbar, Naama, and Anna Belfer-Cohen. 1998. The technological abil- Palaeolithic mobility and land use in the southwestern Pyrenees: the ex- ities of the Levantine Mousterians: cultural and mental capacities. In Nean- ample of level 10 in La Roca dels Bous (Noguera, Catalunya, northeast S336 Current Anthropology Volume 54, Supplement 8, December 2013

Spain). In Settlement dynamics of the Middle Paleolithic and Middle Stone ropean context. In The ancient human occupation of Britain. Nick Ashton, Age, vol. 2. Nicholas J. Conard, ed. Pp. 415–435. Tu¨bingen: Kerns. S. G. Lewis, and C. B. Stringer, eds. Pp. 91–112. Amsterdam: Elsevier. Olle´, A., M. Mosquera, X. P. Rodrı´guez, A. de Lombera-Hermida, M. D. Slimak, Ludovic. 2008. Faire des outils entre le 50e`meetle35e`me mille´naire Garcı´a-Anto´n, P. Garcı´a-Medrano, L. Pen˜a, et al. 2013. The early and Middle en France me´diterrane´ene: syste`mes techniques et e´volutions anthropolo- Pleistocene technological record from Sierra de Atapuerca (Burgos, Spain). giques. In Variabilidad te´cnica del Paleolı´tico Medio en el sudoeste de Europa, Quaternary International 295:138–167. vol. 14. Rafael Mora, Jorge Martı´nez-Moreno, Ignacio de la Torre, and Joel Pelegrin, Jacques, and Marie Soressi. 2007. Le Chaˆtelperronien et ses rapports Casanova, eds. Pp. 65–86. Barcelona: Treballs d’Arqueologia. avec le Mouste´rien. In Les Ne´andertaliens, biologie et cultures. Bernard Van- Soressi, Marie. 2004. From the Mousterian of Acheulian tradition type A to dermeersch and Bruno Maureille, eds. Pp. 283–296. Documents pre´his- type B: a change in technical tradition, raw material, task, or settlement toriques 23. Paris: E´ ditions du Comite´ des Travaux Historiques et Scien- dynamic? In Settlement dynamics of the Middle Paleolithic and Middle Stone tifiques. Age, vol. 2. Nicholas J. Conard, ed. Pp. 343–366. Tu¨bingen: Kerns. Rink, W. J., H. P. Schwarcz, H. K. Lee, V. Cabrera Valde´s, F. Bernaldo de Thie´baut, Celine, V. Mourre, P. Chalard, D. Colonge, A. Coudenneau, M. Quiro´s, and M. Hoyos. 1997. ESR dating of Mousterian levels at El Castillo Deschamps, and A. Sacco-Sonador. 2012. Lithic technology of the final Cave, Cantabria, Spain. Journal of Archaeological Science 24:593–600. Mousterian on both sides of the Pyrenees. Quaternary International 247: Rı´os, Joseba. 2008. Variabilidad tecnolo´gica en el Paleolı´tico Medio de los 182–198. Pirineos Occidentales: una expresio´n de las dina´micas histo´ricas de las Turq, Alain. 2000. Le Mouste´rien de tradition Acheule´enne. In Pale´olithique sociedades neandertales. In Variabilidad te´cnica del Paleolı´tico Medio en el infe´rieur et moyen entre Dordogne et Lot. Special issue, Paleo (suppl. 2):244– sudoeste de Europa, vol. 14. Rafael Mora, Jorge Martı´nez-Moreno, Ignacio 273. de la Torre, and Joel Casanova, eds. Pp. 171–194. Barcelona: Treballs Vaquero, Manuel. 1999. Intrasite spatial organization of lithic production in d’Arqueologia. the Middle Palaeolithic: the evidence of the Abric Romanı´ (Capellades, Ripoll, Eduardo, and Henry de Lumley. 1965. El Paleolı´tico medio en Catalun˜a. Spain). Antiquity 73:493–504. Ampurias 26/27:1–67. Vaquero, Manuel, and Eudald Carbonell. 2003. A temporal perspective on Rodrı´guez, Xose P. 2004. Atapuerca y el inicio del Paleolı´tico medio en Europa. the variability of the discoid method in the Iberian Peninsula. In Discoid In Arqueologı´a,vol.4ofMiscela´nea en homenaje a Emiliano Aguirre.Pp. lithic technology: advances and implications. Marco Peresani, ed. Pp. 67–81. 417–431. Zona Arqueologı´a, no. 4. Madrid: Museo Arqueolo´gico Regional. BAR International Series 1120. Oxford: Archaeopress. Santamarı´a, David, J. Fortea, M. d. l. Rasilla, L. Martı´nez, E. Martı´nez, J. C. Vaquero, Manuel, M. G. Chaco´n,M.D.Garcı´a-Anto´n,B.Go´mez de Soler, Can˜averas, S. Sa´nchez-Moral, et al. 2010. The technological and typological K. Martı´nez, and F. Cuartero. 2012. Time and space in the formation of behaviour of a Neanderthal group from El Sidro´n Cave (Asturias, Spain). lithic assemblages: the example of Abric Romanı´level J. Quaternary Inter- Oxford Journal of Archaeology 29:119–148. national 247:162–181. Santamarı´a, David, L. Montes, and P. Utrilla. 2008. Variabilidad te´cnica del Vega, Luis G., P. Cosano, A. Villar, O. Escarpa, and T. Rojas. 1997. Las in- Paleolı´tico Medio en el valle del Ebro: la Cueva de los Moros I de Gabasa dustrias de la interfase Pleistoceno Medio-Superior en la cueva de la Ca- (Peralta de Calasanz, Huesca). In Variabilidad te´cnica del Paleolı´tico Medio rihuela (Pin˜ar, Granada). In II Congreso de Arqueologı´a Peninsular, vol. 1. en el sudoeste de Europa, vol. 14. Rafael Mora, Jorge Martı´nez-Moreno, Rodrigo de Balbı´n and Primitiva Bueno, eds. Pp. 105–118. Zamora, Spain: Ignacio de la Torre, and Joel Casanova, eds. Pp. 319–339. Barcelona: Treballs Fundacio´n Rei Afonso Henriques. d’Arqueologia. Villaverde, Valentin. 1984. La Cova Negra de Xa`tiva y el Musteriense de la Santonja, Manuel, A. Pe´rez-Gonza´lez, P. Villa, C. Sese´, E. Soto, R. Mora, V. regio´n central del Mediterra´neo espan˜ol. Valencia: Diputacio´n Provincial de Eisenmann, and M. Dupre´. 2000. El yacimiento paleolı´tico de Cuesta de Valencia. la Bajada (Teruel). In 3º Congresso de Arqueologia Peninsular: Paleolı´tico da Zilha˜o, Joao. 2001. Middle Paleolithic settlement patterns in Portugal. In penı´nsula ibe´rica, vol. 2. Rodrigo de Balbı´n, N. Bicho, E. Carbonell, B. Settlement dynamics of the Middle Paleolithic and Middle Stone Age,vol.1. Hockett, A. Moure, L. Raposo, M. Santonja, and G. Vega, eds. Pp. 169– Nicholas J. Conard, ed. Pp. 597–608. Tu¨bingen: Kerns. 186. Porto: ADECAP. ———. 2006. Chronostratigraphy of the Middle-to-Upper Paleolithic Tran- Santonja, Manuel, and Paola Villa. 2006. The Acheulean of western Europe. sition in the Iberian Peninsula. Pyrenae 37:7–84. In Axe age: Acheulian toolmaking from quarry to discard. Naama Goren- ———. 2008. The Ebro frontier revisited. In The Mediterranean from 50,000 Inbar and Gonen Sharon, eds. Pp. 429–478. London: Equinox. to 25,000 BP: turning points and new directions. Marta Camps and Carolyn Scott, Becky, and Nick Ashton. 2011. The Early Middle Palaeolithic: the Eu- Szmidt, eds. Pp. 293–311. Oxford: Oxbow. Current Anthropology Volume 54, Supplement 8, December 2013 S337

On Variability and Complexity Lessons from the Levantine Middle Paleolithic Record

by Erella Hovers and Anna Belfer-Cohen

CAϩ Online-Only Material: Supplements A, B

A century of research has led to the recognition of multiple levels of technological variability in the Levantine Middle Paleolithic (MP) that cannot be resolved through single-cause explanatory models. Recent ecological models argue for continual occupation of the region and competitive coexistence of Neanderthal and modern human populations. Current paleogenetic studies underline the feasibility of the latter scenario. The Levantine MP offers a perspective on the interface of historical circumstances and long-term evolutionary mechanisms that structured in-tandem trajectories of technological and behavioral changes as well as insights into the dynamics of nondirectional behavioral complexities in the archaeological record.

Paleoanthropologists dichotomize the global post-Acheulian production of flake-based assemblages (Goren-Inbar 1994), Paleolithic record of 400,000–50,000 yr ago between the Mid- and the proliferation of Levallois flaking systems. At the end dle Paleolithic (MP) of Eurasia/northern Africa and the Mid- of the MP period, the Levallois technology gives way to Upper dle Stone Age (MSA) of sub-Saharan Africa (e.g., Marks Paleolithic (UP) blade-oriented lithic reduction systems cou- 2008). While the MP record (on large temporal and geo- pled with an assortment of less formal flake-oriented tech- graphic scales) has been perceived as relatively monotonous nologies. When examined in detail, the MP record presents and poor in innovations (Kuhn and Stiner 1998), the MSA a mosaic pattern of diversification, loss, and reemergence of record appears studded with technological and symbolic ad- cultural traits throughout its duration (Belfer-Cohen and vances (but see Tryon and Faith 2013). This distinction has Goren-Inbar 1994; Goren-Inbar and Belfer-Cohen 1998; Hov- raised questions about underlying biological, cognitive, de- ers 2009; Hovers and Belfer-Cohen 2006). mographic, and social differences between populations in the A number of reasons make the Levant, a small (ca. 1,000 two metaregions and how those affected the trajectories of km north–south by up to 400 km east–west) geographically cultural evolution in both. Importantly, the record of each circumscribed region (Hovers 2009; Shea 2003), suitable to a high-resolution “dissection” of the MP record (fig. 1).1 A region averages hominin interactions with local social and recurrent problem in environmental reconstructions is that physical environments that had occurred on variable intra- global models, often derived from marine data sources, are and interregional geographic and temporal scales. These incompatible in scale with the size of terrestrial tracts inves- higher-resolution processes should be identified and explained tigated archaeologically and do not take into account the if broader evolutionary trajectories are to be understood effects of local physiography (Rohling et al. 2013). The rich (Hovers 2009:246). Accordingly, we narrow down our dis- history of research on Levantine prehistoric and climatic rec- cussion to the Levantine record of 250,000–50,000 yr ago, ords provides a dense matrix of data and allows for a more which constitutes a discrete MP entity (Bar-Yosef 2006). Our realistic approach to the question of the interaction between aim here is to focus on major traits of the Levantine MP behavior and environmental variability (Goring-Morris, Hov- record as a behavioral system. ers, and Belfer-Cohen 2009; see Hovers 2009 for references). The onset of the MP in the Levant is recognized through Moreover, while Levantine climate corresponded to global the disappearance of the bifacial tools that characterized the climate patterns throughout the Quaternary, the amplitude preceding Lower Paleolithic (LP) complexes, a shift toward of changes has been less dramatic than in Europe, under- mining simple models of climate-driven extinctions or bio- Erella Hovers and Anna Belfer-Cohen are Professors at the Institute geographical movements of hominins and related cultural of Archaeology of the Hebrew University of Jerusalem (Mt. Scopus, Jerusalem 91905, Israel [[email protected]; [email protected] 1. For site-by-site details and dating, see Ahmad (2009), Bisson et al. .ac.il]). This paper was submitted 3 VII 13, accepted 30 VIII 13, and (forthcoming), Conard et al. (2010), Hauck (2011), Hovers (2009, apps. electronically published 20 XII 13. 1, 5), and Sharon et al. (2010).

᭧ 2013 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved. 0011-3204/2013/54S8-0013$10.00. DOI: 10.1086/673880 S338 Current Anthropology Volume 54, Supplement 8, December 2013

Figure 1. Climatic variation shown against site chronologies. The δ18O records from Peqi’in and Soreq caves serve as proxy for the range and magnitude of climatic changes throughout the MP period between 230,000 and 40,000 yr ago (see text). Dated MP sites shown here are placed within their chronological ranges and paleoclimatic contexts as inferred by various dating methods. Where stratigraphic sequences are dated, the arrows show the chronological intervals involved. For the sites of Amud, Kebara, Tabun, Hayonim, and Ein Difla, only the thermoluminescence chronology is shown. Note the millennial scale variations clearly observed in the paleoclimatic record. Paleoclimatic data from Bar-Matthews et al. (2003). See Hovers (2009, app. 6) for detailed chronological information on radiometrically dated MP sites. Reproduced from Hovers (2009, fig. 8.2) with permission from Oxford University Press. changes.2 Furthermore, the arrival of exogenous populations of transmission processes, and are they unique to the Levant into a region is a potential source for cultural variation and within the MP world? change. At least two such events (the respective arrivals of The possibility of behavioral differences between MP mod- modern humans and of Neanderthals) are documented in the erns and Neanderthals is a major issue in elucidating the Levantine MP, providing an opportunity to investigate how Levantine MP, yet we opted to first explore the properties of such historical events are expressed and how they influenced the technological system(s) and only then examine their pos- the local trajectories of cultural evolution. sible correlation with biological distinctions (Hovers 2006; In order to understand how a complex system works, a Lieberman and Bar-Yosef 2005). We premise that the cognitive detailed analysis of the parts and their interactions, followed abilities of MP hominins (and possibly earlier ones; Goren- by an integrated view of why the particular arrangement of Inbar 2011) were comparable, although they may have been parts evolved, is essential (Corning 2002). Accordingly, we expressed and executed differently because of differences in discuss the features that define the Levantine MP, relying mostly on stone tool characteristics. We then identify potential life histories, reproductive success, social organization, or dif- environmental, demographic, and social sources of the be- ferent modes and rates of social transmission (e.g., Belfer- havioral variability associated with the organization of the Cohen and Hovers 2010; Hovers and Belfer-Cohen 2006; lithic technological features. Finally, we turn to questions Langbroek 2012; Rossano 2010; Wynn and Coolidge 2012). about cultural evolution in the context of the Middle and A second premise is that the variety of human responses to Upper Pleistocene in the Levant: can one identify the range shifting and unstable natural and social environments is un- likely to reflect strictly genetically determined responses (Ehr- 2. Almogi-Labin, Bar-Matthews, and Ayalon (2004), Almogi-Labin et lich and Feldman 2003; Pulliam and Dunford 1980). By MP al. (2009), Enzel et al. (2008), Frumkin, Bar-Yosef, and Schwarcz (2011), times, social choices are to be considered as one of the major and Hallin, Schoninger, and Schwarcz (2012) all present detailed research on modern and ancient environments; see summary and discussions in factors in the shaping of a cultural record (Hovers 2009:6–7; Hovers (2009). Sterelny 2012). Hovers and Belfer-Cohen Variability and Complexity S339

The View from the Ground: Characteristics of tions conducive to cost-effective core preparation proce- the Levantine MP dures and successful removals (Gordon 1993; Wojtzak 2011). Levallois flakes were typically detached from broad Lithic Production Systems cores that enabled exploitation of a large surface, and they were relatively unconstrained by the proportions of the Levantine MP lithic assemblages conform to definitions of original raw material. Throughout the MP there was a the Mousterian technocomplex and are often referred to as tendency to select larger blanks (usually Levallois blanks) “Levantine Mousterian.” Those are flake industries charac- for making side scrapers (Hovers 2009 and references terized by (1) absence of “core tools” (a generic name for therein). There are no indications for using fire as an en- bifaces, cleavers, spheroids), and (2) a quantitative dominance gineering tool for lithic raw material manipulation (unlike of Levallois flaking methods (as defined by Boe¨da 1993; in the southern African MSA; Brown et al. 2009; Mourre, Boe¨da, Geneste, and Meignen 1990) used over other formal Villa, and Henshilwood 2010). flaking systems to produce all blank morphotypes (flakes, blades, and points). That said, Levallois flaking systems may co-occur within any assemblage with variable frequencies of blanks and cores from laminar (as defined by Meignen 1998) Extractive Technologies and discoidal production systems. “True” (UP-style) blade Evidence as to the dietary resources used by MP hominins is cores occur in variable frequencies in all published assem- restricted in most cases to animal remains because of their blages. Cores on flakes represent a discrete production system better preservation. Direct evidence for dietary plant exploi- encountered variably in all assemblages, yet the size and shape tation is rare (Henry, Brooks, and Piperno 2011; Henry et al. ranges of its products often fall within those resulting from 2004; Lev, Kislev, and Bar-Yosef 2005; Madella et al. 2002; flaking nodules (Goren-Inbar 1988; Hovers 2007, 2009). Matsutani 1973, 1987), yet it demonstrates that Levantine MP Quina flaking and bifacial shaping are absent, although they populations engaged in the collection and processing (the are known from the earlier Acheulo-Yabrudian (Garrod and latter rarely attested to, but see Lev, Kislev, and Bar-Yosef Bate, 1937; Gopher et al. 2005; Jelinek 1982:3). Flaking sys- 2005) of plant foods, including cereal seeds and lentils. The tems are modular, with preferential and recurrent methods preservation bias has led to an unbalanced focus on tech- of Levallois flaking combined variably with modes of core nologies of meat acquisition (Hovers 1998a), although on the preparation (i.e., centripetal, unidirectional, unidirectional basis of nutritional reasoning (Speth 2010 and references convergent, and bidirectional). Most Levallois morphotypes therein), plants would constitute an important part of the were produced through more than one combination of flaking diet of prehistoric hunter-gatherers in most regions, pending modes and methods (Bar-Yosef and Van Peer 2009; Hovers availability. Even the high metabolic costs of European Ne- 2009; Inizan, Roche, and Tixier 1992; Van Peer 1992). anderthals (Aiello 2003; Churchill 2006; MacDonald, Roe- Changes of flaking modes during the course of reduction are broeks, and Verpoort 2009; Steegman, Cherny, and Holliday often documented, attempting to extend the core’s use-life 2002) may have been met by exploitation of a diverse resource before its discard (Bar-Yosef et al. 1992; Hovers 1998b, 2009; base rather than heavy reliance on large game (Hockett 2012). Nishiaki and Copeland 1992). Only in a few assemblages does Given the mid-low latitude of the Levant, prehistoric hunter- a single Levallois flaking variant account for over half of the gatherers would rely extensively on vegetal resources (Binford Levallois blank population (e.g., Bar-Yosef et al. 1992; Goren- 2001; Cordain et al. 2000; el Zaatari et al. 2011; Hayden, 1981; Inbar 1990; Hauck 2011; Hovers 2009). Kelly 1995). Flint was the nearly exclusive raw material used for all A similar preservation bias against tools made of organic flaking systems. Systematic use of limestone or basalt was materials renders stone tools the main source for understand- reported from only two open-air sites (Gilead 1980, 1988; ing MP extractive technologies. It is assumed that stone ar- Goren-Inbar 1990). Flint is readily available throughout tifacts were used on animal and plant materials, yet direct most areas of the Levant as part of the geological substrate evidence to that effect is uncommon. Analyses of microscopic (Barkai and Gopher 2009; Delage 2007; Druck 2004; Eksh- edge damage and residues of adhesive and mastics on stone tain et al. 2012; Henry 1995b). Procurement was mostly tools provide clues as to the construction of extractive tools. from primary sources, which were exploited selectively ac- Use-wear studies indicate that both unretouched and re- cording to quality, size, and shape suitability for the formal touched blanks were used as multipurpose implements for flaking systems (Levallois or Laminar). Transport distances cutting and scraping of various animal tissues and plant ma- of partially prepared cores from sources to sites ranged terials (Beyries 1988; Bonilauri et al. 2007; Shea 1991). This between !5 and 20 km (typically !10 km), whereas finished is similar to the known technological practices among extant products were rarely transported over distances of 30–40 hunter-gatherers (Oswalt 1976). Modification signs on the km (Delage 2007). base of MP triangular points are consistent with experimental The production of elongated morphotypes may have hafting damage (Shea 1988, 1998), although evidence for thin- promoted selection of nodules of specific size or propor- ning or notching of the proximal ends in preparation for S340 Current Anthropology Volume 54, Supplement 8, December 2013 hafting is scant. Pointed blanks bearing such marks, with hack-Gross et al. 2008; Yeshurun, Bar-Oz, and Weinstein- putative impact fractures at the tip and edge damage,3 were Evron 2007). The different hearth types may have served interpreted as hafted hunting weapons—used either by thrust- different social or economic roles, though empirical data ing (Shea 2006) or thrown over short distances (Boe¨daetal. to this effect are still missing. 1999)—often doubling as cutting tools. The most compelling evidence for hafting comes from Umm el-Tlel, where ca. 70,000 yr ago bitumen was procured Pigments, Shells, and Burials from a distance of 40 km, processed (minimal heating was involved), and used systematically as an adhesive for a large Several sporadic instances of processing and use of “non- number of retouched and unretouched blanks, including but utilitarian” materials are known from Levantine MP sites not exclusively Levallois points (Boe¨da et al. 1998, 1999, 2008; dated to marine isotope stage (MIS) 5e-b. Ochre has been see also Friedman et al. 1994–1995). This practice stopped reported from Skhul and Qafzeh caves (120,000–90,000 yr between 70,000 and 40,000 yr ago even though the production ago). It was collected locally (10–30 km from the sites, contra and use of Levallois points continued. Combined with the Salomon et al. 2012). In both sites pyrotechnology was used low frequencies of MP points with impact fractures (e.g., Shea to transform yellow goethite to red hematite. The operational 2006; Villa et al. 2009), these data imply that pointed forms sequences of preparing ochre powder involved in some in- were not necessarily used as composite hunting weapons stances heating as well as scraping of ochre lumps with sharp (Boe¨da et al. 1999; Plisson and Beyries 1998; Shea 2006). stone tools, abrasion, and grinding (Godfrey-Smith and Ilani There is also sporadic evidence for the use of bitumen in the 2004; Hovers et al. 2003; Salomon et al. 2012). In both in- nearby site of Hummal (Hauck et al. 2013). stances the pigment use was inferred to have been of symbolic Whether hafting occurred and whether it was restricted to importance. specific-function tools are questions of interest because a shift No dietary expansion into marine resources has been doc- from handheld tools (e.g., the Scho¨ningen wooden spears; umented in the Levantine MP, contrary to its European and Thieme 1997) to hafted (composite) tools was a significant African counterparts (Avery et al. 2008; Corte´s-Sa´nchez et al. technological innovation and possibly bears on higher cog- 2011; Finlayson, Barton, and Stringer 2001; Klein and Cruz- nitive capacities (e.g., Greenfield 1991; Wadley, Hodgkiss, and Uribe 1996; Marean et al. 2007; Stiner 1994; Stringer et al. Grant 2009). 2008). Nutritionally insignificant, naturally perforated shells occurring sporadically in Qafzeh and Skhul are interpreted as beads (Bar-Yosef Mayer, Vandermeersch, and Bar-Yosef Hearths and Combustion Features 2009; Vanhaeran et al. 2006; Walter 2003). The shells are similar to those found in Africa during oxygen isotope stage Combustion features have been documented in long cave 5–4 (Nassarius gibbosulus in Skhul) and in European sites sequences from the late LP onward (Henry et al. 2004; Kar- (Glycemerys asurbica in Qafzeh; Bar-Yosef Mayer, Vander- kanas et al. 2007; Meignen, Goldberg, and Bar-Yosef 2007; meersch, and Bar-Yosef 2009; Zilha˜o et al. 2010). The distance Shahack-Gross et al. 2008). The presence of burned lithic of Qafzeh from the Mediterranean shore (∼40 km) suggests debris and artifacts suggests that the lack of clear hearths in selective collection and inland transport (or exchange). stratigraphically shallow open-air sites is potentially a pres- Incised cortices on Levallois cores are known from Qaf- ervation bias (see Gilead 1988 for an exception). zeh Cave, where some of the ochre lumps are also incised, Fireplaces in Levantine MP sites were either flat surface and Quneitra (∼54,000 yr ago; Goren-Inbar 1990; Hovers, (“simple”) or pit hearths (March et al. 2012) constructed Vandermeersch, and Bar-Yosef 1997; Hovers et al. 2003; with wood without evidence for use of auxiliary combus- Marshack 1996). The presence of ochre residues within tibles (Albert, Berna, and Goldberg 2012; Albert et al. 1999, some of the incisions on the Qafzeh item (Nowell, d’Errico, 2003; Madella et al. 2002; Shahack-Gross et al. 2008). Be- and Hovers 2001) suggests that the two types of finds are ginning from the Early MP (EMP, ca. 250,000–170,000 yr linked causally. ago), combustion features occurred as small, thin lenses Several instances of modern (Skhul and Qafzeh) and Ne- representing flat fires directly laid on the soil substrate; as anderthal (Tabun, Kebara, Amud, Dederiyeh) skeletal remains large hearths, sometimes in basin-like shallow structures; are identified as burials based on their specific interment con- and as massive (primary or secondary) accumulations of texts (Belfer-Cohen and Hovers 1992; Hovers, Kimbel, and ashes (Goldberg and Bar-Yosef 1998; Meignen, Goldberg, Rak 2000; Hovers et al. 1995; Nilsson 1998; Tillier 1990). and Bar-Yosef 2007; Rabinovich and Hovers 2004; Sha- Burials are found in thick undifferentiated deposits (Skhul), or within several stratigraphic horizons, interment took place 3. Attempts to identify other diagnostic wear—e.g., Wallner lines, in a given locality over a period of time. When adjusted for which provide evidence about projectile, thrusting, or other manners of using hunting armature (Hutchings 2011)—on pointed items in assem- sediment volume per time (calculated for Qafzeh, Amud, and blages from the whole MP time range have been unsuccessful, possibly Kebara), the behavior seems more common among modern because of the use of flint (K. Hutchings, personal communication, 2009.) humans. Hovers and Belfer-Cohen Variability and Complexity S341

From Data to Behavior Bamforth 1986; Eren et al. 2005; Geneste 1985; Thie´baut et al. 2010; see Nelson 1991).5 Decisions made by hunter-gatherers regarding “the selection and integration of strategies for making, using, transporting, and discarding tools and the material needed for their man- Moving Around: Artifact Use-Lives ufacture and maintenance” (Nelson 1991:57) culminate in technological organization: a repertoire of behavioral strate- Throughout the MP, the characteristic pattern of raw material gies aiming to reduce survival risks by providing technological procurement involved transport of raw material as partly pre- means when needed (Bamforth and Bleed 1997; Bousman pared cores or as blanks over distances typically ranging from 2005; Johnson 1978; Moore 1981; Reynolds 1978; Ugan, 5 to 20 km to caves or open-air sites, transforming them into Bright, and Rogers 2003). It entails variable degrees of mo- temporary raw material sources (Kuhn’s [1995] strategy of 6 bility and raw material curation (recycling and maintenance; “provisioning of places”). Assemblages usually contain prod- ucts from all postdecortication stages of core reduction—core Andrefsky 1994; Binford 1977, 1979, 1989; Bleed 1986; Kuhn preparation, blank removal, core reshaping, blank retouch, 1993, 1994; Marks 1988; Shott 1989) in response to fluctu- and discard. Blank recycling and reshaping in order to extend ations in the temporal and spatial distributions of resources. the artifacts’ use-lives are more common in EMP assemblages Table A1 in CAϩ online supplement A shows archaeological where retouched tools are relatively abundant. In late MP proxies for organizational strategies. assemblages, the final shape of a tool was dictated mainly In the rich Mediterranean ecological zone, the clumped during flaking rather than through retouch. Retouch intensity distributions and geographic seasonality gradients of vegetal (invasiveness on the blank’s surface) and extent (length along resources combined with the small and stable territories of the edge) tend to be low regardless of the production system the main prey animals created nearly yearlong patchy mo- employed; the distance from raw material source is correlated 4 saics. Availability of peak quality resources was segregated with the amount or intensity of retouch only occasionally spatially and temporally. The importance of annuals and (Hovers 2009:209–211). grasses would increase at the expense of trees as one moved Raw material was redistributed from workshop/habitation east and/or south, where plants and animals that are obligate sites across the landscape as tools intended for activities else- drinkers are found near water bodies and could be exploited where. Levallois blanks constituted the mobile component of throughout the year, albeit intermittently. Beyond such focal the knapped stone technology because they were bigger and localities (e.g., the sites of Hummal or Umm el Tlel), plants relatively thinner than other flakes, providing higher utility in the semiarid zone and in hyperarid areas would be clustered ratios per unit mass (Eren and Lycett 2012; Geneste 1985:526; in the winter along ephemeral drainages. Gregarious migra- Hovers 2009:77–80; Kuhn 1994). In the absence of long re- tory species or resident species frequenting water sources fitted sequences (Far’a II and Tor Faraj being exceptions; De- would be a comparatively abundant and predictable food sup- midenko and Usik 2004; Gilead 1988), the best indication for ply even if only for a short term. transport of finished blanks to and from sites (possibly as Ecological considerations (e.g., Dyson-Hudson and Smith individual tool kits; Binford’s [1979] strategy of “personal 1978; Kelly 1983, 1995) suggest that in low and medium-low gear”) are blank/core ratios. Elevated frequencies of pointed latitudes, plant resources play a major role in determining the and elongated blanks in assemblages where suitable cores and core-trimming elements are rare suggest import of prepared intervals, timing, and distances of group mobility because they blanks (Ekshtain et al. 2013; Henry 1995a; Jelinek 1982; Meig- require continuous resource monitoring to ensure accurate nen 1998; Sharon et al. 2010), while the absence of such scheduling of their harvest, and because long-distance trans- port of abundant yet short-lived resources is economically 5. It has been argued that in Europe Neanderthals were also highly wasteful in the absence of storage technology. Such constraints mobile because of their heavy reliance on faunal resources, which are are optimally mitigated by moving consumers to the resource less dense on the landscape (see Kuhn 2013). One of the first adaptations patch (residential mobility; Bar-Yosef and Meadow 1995; Met- of Neanderthals in the Levant would have been reduction in costs of maintaining their metabolically expensive anatomy, which would select calfe and Barlow 1992; O’Shea 1981). Given a patchy resource for a fast organizational response by adjusting their extractive behavior distribution over short distances, MP hominins in the Levant and associated technological organization to the different local conditions. might have prioritized high residential mobility levels and Moreover, if European Neanderthals relied on plants more heavily than opted to curate raw material by artifact recycling (“remaking previously thought (El Zaatari et al. 2011; Henry, Brooks, and Piperno 2011; Hockett 2012), changes from “European” to “Levantine” organi- an implement into a different kind of tool”; Odell 1996:95) zational strategies may not have been so drastic. A biological response and blank maintenance by reshaping (e.g., resharpening; to relaxed environmental pressures, e.g., in the form of changes in the proportions of Levantine Neanderthal long bones (Arensburg and Belfer- Cohen 1998 and references therein) may have lagged behind. 4. Hovers (2009:200–204) presents details and references for the eco- 6. Transport distances to Negev sites are argued to be on the scale of logical conditions inferred from paleoenvironmental data and reconstruc- tens or hundreds of meters (Gilead 1980; Munday 1976), and raw material tions. exploitation in them is more expedient (Hovers 2009:280). S342 Current Anthropology Volume 54, Supplement 8, December 2013 elements from assemblages where appropriate cores do exist from cave occupations. Beyond the trivial observation that implies their removal from the assemblages (e.g., Qafzeh VII, only animals that were available could be hunted, the com- sites D44, D45, and D2 in the Negev, Kebara VIII, and Tabun positions of faunal assemblages reflect complex decision-mak- C; Hovers 2009; Meignen and Bar-Yosef 1992; Munday 1976; ing processes rather than mere availability (Speth 2012). Thus, Shea 1991). lithic raw material management in most open-air localities The patterns of raw material choice and transport to “cen- resembles that of caves: provisioning, on-site knapping, low tral places” observed regionwide are significant in showing investment in blank recycling, and high typological diversity. that (1) raw material was evaluated on the basis of techno- An important difference that sets open-air sites in the Med- logical knowledge rather than exploited opportunistically on iterranean zone apart from caves is the low frequencies of the basis of its availability; (2) in many instances, raw material Levallois blanks and the elevated frequencies of expedient procurement was conducted as a focused activity at distances tools such as notches/denticulates (fig. 2; Hovers 2009, table that may have exceeded daily foraging ranges; (3) throughout 8.2). The exceptions observed at Umm el Tlel and Hummal the MP, lithic technological activities were organized across (Boe¨da, Griggo, and Noe¨l-Soriano 2001; Hauck 2011) may familiar territories; and (4) the differences in core designs be related to their locations in oases, often being used as (e.g., Wallace and Shea 2006) and in frequencies of heavily habitation sites. curated items between the EMP and the later MP suggest Lithic and faunal data converge to show that Levantine MP changes in organizational strategies (table A1). While resi- sites in the Mediterranean zone, in various geographic loca- dential mobility seems to have been prioritized in both, tool tions and in cave and open-air contexts, were usually a “mixed availability was negotiated differently in the EMP compared bag” of organizational practices. Outside of the Mediterranean with the later MP, with different emphases on raw material ecological zone, assemblages are more skewed toward mobile curation/recycling and tool maintainability (in the sense of technological elements (Hauck 2011; Hovers 1997; Shea Bleed 1986), respectively. 1998).

Places of Activity: Caves and Open-Air Sites Places of Activity: Features of Levantine MP Occupations Some open-air sites are located next to discrete resources (e.g., Caves and a few open-air sites (Hummal, Umm el Tlel; quarry sites on raw material sources; hunting stations in prox- Nesher-Ramleh; Zaidner et al., forthcoming) were repeatedly imity to water bodies where fauna could be anticipated). Ac- occupied; accumulated stratigraphic sequences span ca. tivities may have been focused on specific tasks (e.g., Marks 200,000–20,000 yr, yet with stratigraphic and occupation hi- and Freidel 1977; Sharon and Oron 2013), leading to differ- atuses. Other open-air sites represent shorter temporal ranges. ences in technology-assisted activities and therefore in lithic Complex depositional processes and high densities obfuscate assemblage compositions (e.g., Marks and Freidel 1977). Dif- distinctions between isolated occupations of either short or ferences in the compositions of lithic assemblages can be ex- long duration versus some long and intensive occupations as pected between open-air sites, which might have served more well as the estimates of the occupying group sizes. often as task-specific places as compared with cave sites, which Explicit as well as latent spatial patterns show that Levantine were used more often as habitation sites. MP sites are not randomly cluttered debris concentrations. The evidence from Levantine MP caves is generally con- Mainly late MP sites show indications for differential use of sistent with their being central places/habitation sites. The space. Some spatial patterns are site specific while others are lithic assemblages represent variable mixtures of provisioning recognized repeatedly in various sites. The latter include strategies as well as production and maintenance of stone hearth-related knapping activities, ash cleaning into desig- tools on-site. Faunal remains indicate selective transport based nated areas, and evidence for well-defined activity areas ex- on body size, meat utility, and marrow content: large fauna clusively related to animal carcass treatment/discard to the (rhinos, aurochs, equids) are scarce, whereas gazelle- and fal- execution of specific lithic reduction sequences or the disposal low deer-size prey were brought into the caves as nearly com- of human remains. Separation of lithic reduction stages across plete carcasses (Rabinovich and Hovers 2004; Rabinovich and occupation space—for example, core preparation in one area Tchernov 1995; Speth 2012; Speth and Clark 2006; Speth and and advanced knapping carried out near hearths in another Tchernov 2007; Stiner 2005; Yeshurun, Bar-Oz, and Wein- area as well as areas designated for stone stocking—were also stein-Evron 2007). The faunal assemblages from open-air sites observed (Alperson-Afil and Hovers 2005; Garrod and Bate include the remains of large-bodied mammals in addition to 1937:62–63; Gilead 1988; Henry et al. 2004; Hietala 2003; the smaller species known from caves (Davis, Rabinovich, and Hovers et al. 1995, 2011; Meignen, Goldberg, and Bar-Yosef Goren-Inbar 1988; Gilead and Grigson 1984; Griggo 1998; E. 2007; Oron and Goren-Inbar 2013; Rabinovich and Hovers Hovers et al., unpublished manuscript, 2013; Rabinovich and 2004; Schick and Stekelis 1977; Shahack-Gross et al. 2008; Hovers 2004; Sharon et al. 2010; Zaidner et al., forthcoming). Speth 2006; Speth et al. 2012). Particular activities were some- Thus open-air sites complement subsistence patterns gleaned times conducted repeatedly in the same area within a site, Figure 2. Values of IL, ILty, and percentages of retouched blades in various MP sites. Reprinted from Hovers (2009, fig. 8.4) with permission from Oxford University Press. S344 Current Anthropology Volume 54, Supplement 8, December 2013 suggesting that the specific functional assignments of areas The Beginning of the Levantine MP: Gains, Losses, and were retained over time. Variations on Themes The functions of sites within their respective settlement The onset of the MP in Eurasia and of the MSA in Africa is systems changed through time. Taphonomy, body part and characterized by the disappearance of LP bifaces from the age distribution of ungulate remains, and frequency changes archaeological record,7 which appears as a rupture from earlier within the pertinent lithic assemblages suggest that the roles industries, concurrent with the proliferation of Levallois flak- of sites shifted between hunting to habitation or butchering ing as a system for flake production. The origins of Levallois locations (Boe¨da, Griggo, and Noe¨l-Soriano 2001; Hauck flaking concepts in the Levant are traced to the late Acheulian, 2011; Meignen et al. 2006; Speth and Tchernov 2007). Faunal with two pathways identified for this change: recycling hand- data have also been used to infer occupation duration/inten- axes into (centripetal) Levallois cores for preferential flakes, sity. At EMP Hayonim Cave (ca. 200,000 yr ago), the large and flake removals from the center of a core surface but sizes of slow-growing tortoises that were harvested by the without rigid hierarchical relationship between core surfaces hominins implied infrequent and ephemeral occupations (Sti- and platforms (e.g., DeBono and Goren-Inbar 2001; Malin- ner, Munro, and Surovell 2000). In contrast, throughout the sky-Buller, Grosman, and Marder 2011).8 However, in the late late MP sequence of Kebara Cave (60,000–48,000 yr ago), the LP Acheulo-Yabrudian technocomplex (400,000–220,000 yr site’s occupants gradually shifted to exploitation of smaller- ago) that immediately precedes the MP, evidence for Levallois bodied animals and younger individuals of those species that flaking (e.g., in Adlun, Tabun, Masloukh, Qesem Caves, Ya- offered lower meat or fat returns. This has been interpreted brud I Rockshelter) is nonexistent (Gopher et al. 2005) or as hunting at ever-increasing distances from the site to com- scanty at best (Copeland 1983; Garrod and Bate 1937:79–89; pensate for locally diminishing returns resulting from over- Jelinek et al. 1973; Skinner 1970; E. Hovers, personal obser- hunting (Speth 2004; Speth and Clark 2006). Hunting pres- vations). This is unlike the long “transitional” phases in Africa sure throughout the Upper Pleistocene may have had a or Europe at more or less the same time. long-term negative effect on the abundance of large ungulates The systematic production of elongated blanks is well doc- (Davis, Rabinovich, and Goren-Inbar 1988). umented in the Acheulo-Yabrudian, yet these flaking tech- An emphasis on “simple” (bidirectional or unidirectional) nologies (Shimelmitz, Barkai, and Gopher 2011) do not seem core reduction modes (Kuhn 1995; Parry and Kelly 1987; to continue into the EMP. The increased control over cores Wallace and Shea 2006) increases gradually during the later (and thus over the shapes of the products), which is a tech- part of the MP. This may correspond to either reduction in nological advantage of the Levallois and Laminar systems, led mobility levels or an increase in site use intensity. However, to a range of recurrent flake morphologies that are the hall- changes in site function(s) are not necessarily reflected in all mark of the Levantine Mousterian throughout its course. Sys- features of the occupation. At Qafzeh Cave, hearths, burials, tematic production of pointed elongated blanks as desired and micromammals occur only in the early layers, yet changes products, obtained through such enhanced lithic technolog- in lithic technology do not correspond to this dichotomy. ical control (sometimes with further modification by re- Shifts in retouched tool-type frequencies crosscut the dis- touch), is a novel component of EMP flaking systems and tinction between the lower and upper sections, and the pro- appears to be a local innovation (see below). portions of the large ungulates remain similar through time Shea, Davis, and Brown (2001) concluded from experi- (Bar-Yosef and Vandermeersch 1993; Hovers 2009; Rabino- mental work that the early elongated points were more suited vich and Tchernov 1995). for use as knives than as tips of heavy thrusting weapons, and their emergence in the Levantine MP record may represent the inception of hafting techniques. Importantly, elongated points co-occur in the earliest MP assemblages with short, What Is “The Levantine MP Variability”? subtriangular Levallois points (Ashkenazi 2005; Copeland 1975; Meignen 1998, 2011; Monigal 2002; Nishiaki 1989; Because of the intrinsic properties of the prehistoric record Weinstein-Evron et al. 2003; Wojtzak 2011). The two mor- (Goring-Morris, Hovers, and Belfer-Cohen 2009) but also photypes undergo a frequency shift over the course of the because of taphonomic and research biases, the hominin evo- lutionary record is constructed from unequal distributions of 7. MP bifaces in Europe are a late phenomenon, different technolog- data points in time and space. Characterizations of Levantine ically from the Acheulian ones (e.g., Soressi 2005, and references therein). MP lithic technology sometimes selectively focus on isolated Therefore, they should not be considered as a continuous trait but rather technological aspects rather than whole-assemblage proper- as technological innovations within the European MP. ties. Still, the Levantine MP can be characterized as a discrete 8. Similar phenomena during early MIS 8 in Europe (Moncel et al. archaeological phenomenon. Behavioral features that contin- 2011; Tuffreau 1995; White, Ashton, and Scott 2011) are identified as immediate precursors of Levallois methods, whereas in eastern Africa ued from earlier periods survived throughout the record, oth- they are regarded as fully Levallois, yet they occur in a mosaic pattern ers that were lost early on were not regained, and a few in- with ESA bifacial elements (McBrearty and Tryon 2006; Tryon, McBrearty, novations held on throughout the period. and Texier 2005). Hovers and Belfer-Cohen Variability and Complexity S345

MP, with the frequencies of the long points declining abruptly of this population (Hovers, Vandermeersch, and Bar-Yosef, from ca. 170,000 yr ago, while short (unretouched or lightly 1997; Hovers et al. 2003).10 retouched) Levallois points become more numerous (though rarely the major component) through time (Hovers 2009:216, Changes within the Levantine MP Record table 8.3; but see Hauck 2011). A now-classical model of linear development of Levantine It is doubtful that the rise of broad-based points is related MP lithic industries based on the sequence of Tabun Cave to changing extractive strategies or to their greater efficacy as (Bar-Yosef 1998; Copeland 1975; but see Ronen 1979; table hunting weapon components. By current archaeozoological B1 in CAϩ online supplement B) emphasizes the presence evidence, species compositions remained similar from the or absence of a few selected technological traits (e.g., Laminar 9 Late LP throughout the MP. This is especially true for the products, specific modes of Levallois flaking or of Levallois Mediterranean ecological zone because of its higher sustain- morphotypes) as characteristics that differentiate three chron- ability in the face of climate variations (Belmaker and Hovers ocultural phases of the Levantine MP. However, as the di- 2011; Hovers 2009 and references therein). Hence, changes versity of MP flaking systems (i.e., the number of flaking in faunal resource structure need not have driven the change systems in the technological repertoire) had already become in point frequencies (Shea 2006). established in the EMP and persisted throughout the period Habitual fire making and fire control are the main behav- (Goren-Inbar and Belfer-Cohen 1998), assemblages portray ioral features that continued uninterrupted from the late LP shifts in the frequencies of technological variants rather than into and through the MP, with a range of variation in size any single dominant variant or morphotype. This is a sig- and shape of the fire structures encountered from the EMP nificant distinction, because an “assemblage” is a basic ana- onward (e.g., Weiner, Goldberg, and Bar-Yosef 2002; Yeshu- lytical unit that nonetheless averages events of group-level run, Bar-Oz, and Weinstein-Evron 2007). In contrast, ochre decision making. Following this, interassemblage variations (and the evidence of its heating to change its color) as well and how they are patterned chronologically and geographi- as marine shells occur briefly, during MIS 5e-b, only in Skhul cally offer interesting insights as regards the behaviors they and Qafzeh. Another MP innovation is hafting with adhesives. might reflect. While its earliest known occurrence is in late Middle Pleis- Interassemblage variability is identified along two main tocene Europe (Mazza et al. 2006), it appeared in the Levant axes. Along the temporal axis, a major change occurred be- only during MIS 4 and involved different components from tween the EMP and the later MP in relative frequencies of those used either in Europe (where birch tar was used in technological variants, with more obvious raw material and artifact curation strategies observed in the earlier MP. Geo- addition to bitumen; Caˆrciumaru et al. 2012; Gru¨nberg 2002; graphic variations are less conspicuous in the EMP.11 In con- Pawlik and Thissen 2011) or in Africa (ochre; Lombard 2007; trast, interassemblage variability in the later MP patterns Rots and Van Peer 2006; Rots, Van Peer, and Vermeersch 2011; largely geographically (the Mediterranean vs. the semiarid and Wadley 2005), suggesting that it may be another local in- arid zones) even though the strategies of raw material curation novation, again involving the use of fire as an engineering in both zones emphasize the knapping of maintainable, ver- tool. Because of the functional advantages of hafting with satile tools (Bleed 1986). Within the Mediterranean zone, later adhesive, Boe¨da et al. (2008) ascribe the gap in bitumen use MP interassemblage variability decreases with geographic dis- at Umm el Tlel to changes in territorial behavior, implicitly tance; that is, assemblages from sites that are geographically assuming its continuous use. closer to one another (or occur at the same locality through- The evolution of mortuary behavior in the Levantine MP out its stratigraphy) are more similar than those that are differs from either Africa or Europe, where, respectively, bone farther away.12 With the exception of Tabun, assemblages in modifications and polishes unrelated to cannibalism (Bodo, a given sequence tend to be more similar to one another Herto) and body caching and cannibalism (Sierra de Ata- (Hovers 1997). puerca) appear already in the Middle Pleistocene. In Europe this behavior developed into interment in the Upper Pleis- 10. The claim is stronger in Qafzeh given the more accurate excavation tocene (Hovers and Belfer-Cohen 2013; Pettitt 2010). In the techniques applied during fieldwork. The Skhul evidence is not as per- Levantine MP, burial appears without recognized precursors suasive. Note that ochre, shells, or incised items do not occur in the during MIS 5–3. The stratigraphic associations of “nonutili- burials yet are associated with them stratigraphically and are absent from contexts without burials. tarian” objects (see above) strictly with burials of modern 11. Although this may be partly related to the lower resolution of the humans provides contextual support for the suggestion that EMP record. these traits may constitute a novel symbolic/behavioral suite 12. This is probably the reason why the temporal Tabun model fails to encompass the full range of variability of the Levantine Mousterian (Goren-Inbar and Belfer-Cohen 1998; Hovers 1998b; Meignen 2011). The 9. Large-game species usually reflect local availability (Bar-Yosef 2004), model holds better for assemblages within the Mediterranean region though there is a possibility of choice-related bias (e.g., Marder et al. (Hauck 2011; Hovers 1997, 2009) but is less successful in chronological 2011). attribution of assemblages outside the Mediterranean zone. S346 Current Anthropology Volume 54, Supplement 8, December 2013

Changes in Settlement Patterns On the basis of paleoecological reconstructions, the distri- bution and structure of resources did not shift significantly throughout the MP, and human groups would tend to be small and highly mobile. The change in curation behavior through time therefore suggests a shift in associated organi- zational strategies—specifically, the geographic scale of group movements. It occurs in tandem with changes in the markers of occupation intensity within the Mediterranean zone: den- sities of the lithic assemblages (fig. 3) and faunal remains per time unit as well as investment in site features seem to increase from ca. 130,000 yr ago, setting EMP occurrences apart from later ones.13 The archaeological record suggests that at least from that time, larger populations were concentrated within the Mediterranean zone, ecologically the most attractive part of the Levant. Such changes in modes of occupation and the geographic patterning of interassemblage technological variability are Figure 3. Lithic densities/accumulation time, in four MP sites. consistent with a scenario of increased group territoriality Data for Hayonim and Kebara, Bar-Yosef (1998); for Qafzeh, within the Mediterranean zone. MP groups that converged Hovers (1997); for Amud, E. Hovers, personal observation. A into the Mediterranean zone would exploit more efficiently color version of this figure is available in the online edition of the seasonal availability of resources within their contracted Current Anthropology. territories.14 Repeated burials in specific locations may be a result of this contraction process, perhaps denoting the emer- later MP are not tantamount to an overall increase in pop- gence of some degree of territoriality and corporateness (Hov- ulation size. If anything, settlement patterns in the very late ers 2001, 2009). Ecological constraints (see above) may have MP and in the UP show more ephemeral than intensive oc- kept group size at low numbers, but sites within a group’s cupations (Goring-Morris, Hovers, and Belfer-Cohen 2009; territory would be visited more frequently, leading to a more Rabinovich 2003; Speth and Tchernov 2007). There are no rapid and extensive accumulation of archaeological remains indications for social, ritual, and cultural innovations that and to recurring spatial relationships between domestic fea- might be associated with robust demographic networks of tures within the sites (Hovers 1997, 2001; Meignen et al. high-density (or growing) populations (e.g., Hovers and Bel- 2006). Outside of the Mediterranean zone, groups probably fer-Cohen 2006; Jochim 1983; Powell, Shennan, and Thomas still exploited larger territories (Boe¨da et al. 2008; Hauck 2009; Shennan 2001). To the contrary, indications for such 2011). cultural intensification (e.g., shell beads or pigment use) in- The changes in settlement patterns between the EMP and creased gradually throughout the UP from sporadic occur- rences in the earliest phases (e.g., Bar-Yosef Mayer 2005; Kuhn 13. Misliya Cave, with a large hearth (Yeshurun, Bar-Oz, and Wein- stein-Evron 2007) and high lithic and faunal densities (Y. Zaidner, per- et al. 2001), while burials occur only later in the UP. Some sonal communication), is exceptional in the EMP. One possible expla- conceptual approaches to the volumetric management of nation is that it was the locale for social interactions of groups that cores were retained from the MP to the UP, and minute yet otherwise used sites ephemerally throughout their large exploitation ter- substantial changes in core-shaping techniques (e.g., abrasion) ritories. Initial observations suggest an increase in artifact and faunal as a way to facilitate hafting occurred only in later stages of densities at Nesher-Ramleh at ca. 170,000 yr ago (Zaidner et al., forth- coming). the UP (Belfer-Cohen and Goring-Morris 2007, 2009; Belfer- 14. An area of 120,000 km2 in the Mediterranean zone (roughly the Cohen and Hovers 2010; see Meignen and Bar-Yosef 2002). size of the area in which MP sites are known) could carry between 8,640 Given these observations, a parsimonious explanation of and 14,992 persons (see Hovers 2009:206 for discussion; Shea 2004:170– the trends in land-use patterns within the later MP is that 173). Note that the estimates here are based on values for extant groups they depict a historical occurrence rather than a long-term in temperate forests, where primary production is lower than in the Mediterranean forests; also, effects of technological extractive aids was evolutionary trajectory of gradual population growth during not factored into the equations. Thus carrying capacity may have been the MP. The posited human density in the Mediterranean higher during MP times. Based on these numbers alone, a 25-person zone is attributed to the effects of changing landscapes and band of hunter-gatherers could survive in the Mediterranean zone in an environmental constraints (e.g., high sea levels, high stands 2 annual territory as small as 350–400 km (realistically, it would have been of Lake Lisan in the Dead Sea Rift that reduced the area of slightly larger); a viable population of 500 individuals (Wobst 1974, 1976) would require a territory of at least 800–1,000 km2. In terms of carrying habitable land, greater aridity outside of the Mediterranean capacity, viable groups could survive in small territories as postulated zone), which may have resulted in an influx of people from here. the arid areas. Possibly this is also related to dispersal events Hovers and Belfer-Cohen Variability and Complexity S347 into the Levant during environmental windows of opportu- Microevolutionary Processes: Cultural Transmission nity in the later MP (Bar-Yosef 2000; Drake et al. 2011; Lahr An alternative hypothesis for explaining the variability in the and Foley 1998; Vaks et al. 2007). material culture record of the Levantine MP invokes micro- evolutionary mechanisms of cultural transmission (e.g., Bet- Discussion tinger, Boyd, and Richerson 2009). The discernible change between the EMP and later MP reflects reorganization of land- From the broadscale, low-resolution cultural evolutionary use patterns. Interassemblage variability observed in the later perspective, the Eurasian MP record has been perceived as a MP stems from flexible combinations of provisioning and period of cultural stasis over some 200,000 yr without cu- curation strategies along ecological gradients. We have sug- mulative changes that led to cultural evolution. This has been gested that these changes are related to demographic shifts. attractively conceptualized by “rugged fitness landscapes” Indeed, loss and retention of cultural diversity are associ- (Boyd and Richerson 1996; Dobzhansky 1951), whereby sig- ated with demographic properties; for example, loss of di- nificant, costly changes that enhance fitness only occurred versity is greater, and will occur faster, in small groups. Some when physical and social conditions on the adaptive landscape demographic events (local extinctions, demic diffusion) act were disrupted dramatically (e.g., the shift to sedentism; Bel- (respectively) as direct causes of eradication/reduction of cul- fer-Cohen and Bar-Yosef 2000). This broad perspective, how- tural traits (Henrich 2004; Mesoudi 2011b; Premo and Kuhn 15 ever, does not tell us much about the dynamics that created 2010). Otherwise, changes in cultural diversity may accu- and preserved such stasis. If we are to address questions about mulate because of random drift (Mesoudi 2011a; Neiman alternative pathways to a known end result of an evolutionary 1995) or socially mediated (“biased”) cultural transmission process, a higher-resolution record of the historical details is by individuals, which leads to group-level changes (Mesoudi required. Otherwise explanatory hypotheses become tauto- 2011a; Richerson and Boyd 2005). The latter takes on variable logical. forms of “conformist” (frequency-dependent) and “prestige” (agent-dependent) biases. Both transmission mechanisms re- quire that social institutions of learning (e.g., socially rec- Potential Causes of Variability ognized learning environments where individuals can copy prevalent variants or select mentors from whom they choose The Levantine MP constitutes a distinct entity within the MP to learn) be in place (Henrich and Boyd 1998; Sterelny 2012). world (Bar-Yosef 2006). This overview has underlined the Over time, both mechanisms lead to loss of diversity at properties that distinguish it. The repertoire of material cul- variable paces depending on the strength of model selection ture behaviors defining the Levantine MP falls within the by individual decision makers or the number of mentors range known from Europe and Africa, yet the abruptness of (Boyd and Richerson 1985; Mesoudi 2011a, 2011b; Mesoudi both loss and emergence of the defining lithic characteristics and Lycett 2009). A population size threshold of N p ∼500 differs from the gradual mosaic-like processes observed in the differentiates between marginal and substantial loss of cultural late Middle Pleistocene of Europe and Africa. Some nonlithic complexity (Vaesen 2012:e40989).16 On the other hand, gains behaviors—cave occupation, the use and control of fire, and in technological diversity (i.e., incorporation of inventions as a prime-adult, large-game focused hunting—continued over society-wide innovations; Renfrew 1978) are not dependent from the LP. The LP/MP turnover in lithic practices was fol- only on population size. Even if a new behavior is beneficial lowed by 200,000-yr-long variations on a restricted repertoire and population size is above 250 individuals, it will not spread with few and relatively short-lived innovations. and cultural diversity will not increase if levels of intercon- A detailed terrestrial paleoclimatic record of the Upper nectedness (e.g., the number of linkages between individuals Pleistocene Levant enables an understanding of the effects of in networks of different mating behaviors) are low (Vaesen climate variations on a regional scale, which is more relevant 2012). Social learning can occur in kin-based learning envi- than globally recognized climate patterns for understanding ronments (e.g., extended families) with a restricted trans- the behavior of prehistoric foragers. Given the mild amplitude mission network or within broader social contexts that may of Levantine climate changes, this leads to the insight that promote the spread of behaviors selected for learning because aspects of material culture variability, even if they are envi- of their economic payoff or social rewards (Vaesen 2012; ronment related (e.g., patterns of raw material use, tool func- Young 2012). Even if the same behavior is transmitted within tions), do not respond directly to climatic shifts. Similarly, predictions that lithic variability or organizational strategies 15. Vital information may be lost when an influential or knowledge- will be clearly dichotomized according to the two hominin able individual passes away without having passed on the information species present in the Levantine MP are questionable both (Whallon, Lovis, and Hitchcock 2011). 16. Under certain conditions of the mathematical models, this thresh- theoretically and empirically (Hovers 2006, 2009; Lieberman old may be as high as 1,290 individuals (Vaesen 2012 and references and Bar-Yosef 2005). therein). S348 Current Anthropology Volume 54, Supplement 8, December 2013 each context, the end result in terms of cultural diversification At present there is no concrete evidence that Neanderthals and accruing novel behaviors will differ significantly. and moderns overlapped chronologically in the Levant. This Given the idea that demographic aspects can spur or slow has led to scenarios of competitive exclusion, recursion, or down technological change (e.g., Henrich 2004, 2010; Powell, alternate extinctions due to climate (Shea 2003, 2006, 2008). Shennan, and Thomas 2009; Shennan and Steele 2009), we However, the absence of evidence for temporal overlap may find it useful to consider the possibility of understanding the be an artifact of the small samples of identifiable human Levantine MP stasis from this microevolutionary perspective. remains that archaeologists have at their disposal (Hovers Right now, this notion can be explored only informally, given 2006). If all dated MP assemblages are considered, the tem- the incompatibility of the evolutionary timescale of the MP poral distribution is less discontinuous (fig. 4), and some record and the temporal scale of decision-making and social degree of continuity (and coexistence) cannot be rejected transmission processes (e.g., Bettinger, Boyd, and Richerson (Hovers 2006, 2009). Paleogenetic analyses (albeit notoriously 2009; Jordan 2007) as well as the incipient stage of the relevant transient) have implications that are potentially consistent formal modeling. with this hypothesis (Green et al. 2010; though see Eriksson and Manica 2012). Recognizing Cultural Transmission Processes The persistent occurrence of the same repertoire of flaking in the Levantine MP technologies throughout the MP is noteworthy. Long-term occurrence of any single flaking system need not imply con- To address processes of biased cultural transmission, two is- tinuity of information transmission (and thus demographic) sues need to be tackled. The first is whether there existed systems. However, the consistent use of the same technological environments of social transmission, and the second is repertoire is less likely to represent repeated independent rein- whether demographic continuity is a feasible scenario in the ventions because complex knapping processes do necessitate Levantine MP. teaching (or, at least, direct observation).17 Accordingly, we 1. Although their archaeological signatures may be the most assume some degree of demographic continuity in the Med- versatile and most difficult to decipher, habitation sites were iterranean zone during the later MP. Still, we cannot—and likely the locations of significant intragroup social interactions do not—argue that it prevailed at all times regionwide. (e.g., Rolland 2004). The use and control over fire generated 2. The archaeological record is spotty, and discontinuity a domestic and social context for knowledge transmission. In scenarios are the null hypothesis in many studies of cultural addition to the cooperation implied by hunting, transport of evolution, rendering drift, biological extinctions, dispersals, high-utility body parts into caves and their butchery away and collapses of demographic systems as the main mecha- from hunting locations suggest that delayed consumption and nisms implicated in shaping the pace and trajectory of Pa- food sharing were part of the subsistence-cum-social activities leolithic cultural evolution (e.g., Boyd and Richerson 1985, that took place within habitation sites. The evidence of do- 2009; Henrich 2004; Mesoudi 2011a:8; Premo and Kuhn mestic space allocation and differentiation, raw material con- 2010). Notably, the Levantine MP archaeological record is centration and redistribution, and delayed returns in food characterized by retention rather than gain or loss. The suite sharing, which occur variably in MP open-air and cave sites, of nonutilitarian innovations in Skhul and Qafzeh during MIS suggests that the social relationship of Levantine MP hominins 5, which disappeared altogether by MIS 4 time, represents an (moderns and Neanderthals) extended beyond immediate exception. Such traits appear slightly earlier and less sporad- economic returns and was formally constructed. Such a social ically in Africa (although there, too, they are not continuous organization would provide opportunities for biased trans- throughout the whole MSA time span; Barham 2002; Cain mission of cultural traits, for example, highly specific ways of 2006; Clark 2011; d’Errico, Garcı´a Moreno, and Rifkin 2011; stone tool making or a social perception of a site’s space (e.g., Marean et al. 2007; Texier et al. 2010; although see Marshack David and Kramer 2001). Appearance of mortuary practices 1981 and Roebroeks et al. 2012 for early occurrences in Eu- as a local phenomenon in the later Levantine MP may be rope). These traits may have been introduced into the Lev- significant in view of suggestions that the human practice of antine behavioral repertoire by dispersing populations of burial evidences “new forms of social organization” (Binford modern humans from Africa. Yet if there was indeed a degree 1968). This contrasts with the Acheulo-Yabrudian popula- of demographic continuity in the Levant, it may be difficult tions, whose social behaviors around fireplaces may have had different contexts for resource redistribution and lower levels to decide whether these temporally constrained phenomena of intergroup cooperation than did MP groups (Stiner, Go- represented demic or cultural diffusion processes (Collard et pher, and Barkai 2011). Notably, the Acheulo-Yabrudian pat- al. 2010). The loss of these features may have entailed physical tern itself may represent a shift from the social structure of extinction but could alternatively be a reflection of collapse earlier Acheulian groups, hypothesized to have operated as 17. Modern-day knappers have indicated that mastering any form of large, loosely, and pragmatically united corporate groups Levallois knapping is training, skill, and cognitively expensive compared based on dyadic relationships between mothers and their off- with earlier or later formal flaking systems (M. Eren, G. Sharon, and J. spring (Marx 2004). Shea, personal communication). Hovers and Belfer-Cohen Variability and Complexity S349

Figure 4. Distribution of dated MP assemblages. Darker vertical bars indicate stratigraphic horizons with remains of anatomically modern humans; lighter vertical bars indicate stratigraphic horizons with Neanderthal remains. For Tabun, both the thermolu- minescence (TL) and electron spin resonance (ESR) date ranges are shown. The question mark and the arrow reflect the controversy about the stratigraphic context of the Tabun C1 fossil and its corrected placement in layer B rather than C (base graphic, courtesy Ofer Bar-Yosef). A color version of this figure is available in the online edition of Current Anthropology. of the social networks of small groups (Hovers and Belfer- iterranean zone, was not a period of accelerated rates of major Cohen 2006; Hovers et al. 2003), where even beneficial traits innovations (the persistence of mortuary behavior being a may not survive (Vaesen 2012). notable exception). In this scenario, such groups had to keep Conversely, there are surprisingly few indications, especially occasional physical contact to maintain viable populations for a region with documented population inflows, for diver- (e.g., a site such as Misliya Cave could serve as an “aggregation sification through diffusion (cultural or demic) from neigh- locality” where otherwise widely dispersed EMP hominins boring areas. Not a single piece that could be related to the could have interacted occasionally). The minimal number of African MSA or European MP lithic systems is published from members in such networks is ∼500 (Wobst 1974, 1976). This Levantine MP archaeological contexts despite the presence of overlaps with the suggested critical population size threshold highly diagnostic morphotypes and practices in these areas beyond which major losses of cultural diversity occur when (e.g., Aterian or bifacial points, Quina flaking, Micoquian biased transmission takes place (Vaesen 2012).18 bifaces). That elongated EMP points represented “a moment of contact” with African populations (Brooks et al. 2006:249) Concluding Comments is questionable on chronological and technological grounds. The Levantine MP emerges as a period when small human The Levantine case study, as perceived in this paper, indicates groups had a robust enough social transmission environment that environmental conditions influence human niche con- but were not interconnected sufficiently to generate techno- struction (Laland and O’Brien 2010) but in ways more in- logical innovations (Henrich 2010; Young 2012) leading to tricate and subtle than has previously been suggested. For cultural evolutionary change that is observed archaeologically. example, it has been suggested recently that habitual use of The spatial parameters of the settlement systems identified in fire emerged in Europe only post-LP (Roeberoeks and Villa the Levantine MP (e.g., small, local clusters) could have been 2011) and that European Neanderthals were unable to control conducive to creating short-term diversity that was nonethe- fire (Sandgathe et al. 2011). It was also suggested that in less at high risk of being lost because of biased transmission before becoming habitual innovations (conformity bias es- 18. This scenario can be tested archaeologically because it predicts that given similar ecological and economic conditions, assemblages in different pecially weakens the relationship between diversity and in- sites will differ in the fine nuances of lithic (or other, if preserved) novations; Kandler and Laland 2009). This explains why the technologies (e.g., Hovers 1997, 2009; Meignen 1995; and see MacKay later MP, when groups were more densely packed in the Med- 2011). Research to that effect is currently under way. S350 Current Anthropology Volume 54, Supplement 8, December 2013

Europe, Neanderthal social bonding was dyadic (as in the Palaeolithic levels of Hayonim Cave (Galilee, Israel). Journal of Archaeo- logical Science 30:461–480. Levantine Acheulian) while exploiting very large territories Albert, Rosa-Maria, Francesco Berna, and Paul Goldberg. 2012. Insights on (Kuhn and Stiner 2006; Stiner 2013). If those claims are val- Neanderthal fire use at Kebara Cave (Israel) through high resolution study idated, they may suggest that MP hominins in Europe as of prehistoric combustion features: evidence from phytoliths and thin sec- tions. Quaternary International 247:278–293. opposed to those in the Levant constructed different envi- Albert, Rosa-Maria, Ofer Lavi, Lara Estroff, Steve Weiner, Alexander Tsatskin, ronments of social interaction as well as microevolutionary Avraham Ronen, and Simcha Lev-Yadun. 1999. Mode of occupation of mechanisms of cultural transmission. Tabun Cave, Mt Carmel, Israel, during the Mousterian period: a study of the sediments and phytoliths. Journal of Archaeological Science 26:1249– That said it is important to bear in mind that the geographic 1260. scale of the Levant differs significantly from those of Africa Almogi-Labin, Ahuva, Miryam Bar-Matthews, and Avner Ayalon. 2004. Cli- or Europe. Still, if we are to compare demographic patterns mate variability in the Levant and northeast Africa during the Late Qua- ternary based on marine and land records. In Human paleoecology in the and cultural trajectories of the Levant with those larger Levantine corridor. Naama Goren-Inbar and John D. Speth, eds. Pp. 117– regions, the geographic “clumping” that typifies cultural 134. Oxford: Oxbow. transmission of small-scale social units requires detailed anal- Almogi-Labin, Ahuva, Miryam Bar-Matthews, Dan Shriki, Elina Kolosovsky, Martine Paterne, Bettina Schilman, Avner Ayalon, et al. 2009. Climatic yses on comparable geographic scales (e.g., Bocquet-Appel variability during the last 90 ka of the southern and northern Levantine and Tuffreau 2009; de la Torre 2013; Wurz 2013). Basin as evident from marine records and speleothems. Quaternary Science Humble gains in technological diversity and slow changes Reviews 28:2882–2896. Alperson-Afil, Nira, and Erella Hovers. 2005. Differential use of space at the in the pace of cultural accumulation characterize also the Neandertal site of Amud Cave, Israel. Eurasian Prehistory 3:3–22. Levantine UP. Despite the diversification of lithic technologies Andrefsky, William. 1994. The geological occurrence of lithic material and (Goring-Morris and Belfer-Cohen 2003), it was not until well stone tool production strategies. Geoarchaeology 9:375–391. Arensburg, Baruch, and Anna Belfer-Cohen. 1998. Neandertals and moderns: into the UP that beneficial technological innovations (e.g., re-thinking the Levantine Middle Paleolithic hominids. In Neandertals and microlithization and highly composite tools: Belfer-Cohen modern humans in Western Asia. Takeru Akazawa, Kenichi Aoki, and Ofer and Goring-Morris 2002; the use of shell beads as information Bar-Yosef, eds. Pp. 311–322. New York: Plenum. Ashkenazi, Hila. 2005. Standardization of blade production in the Levantine transmission systems: Kuhn et al. 2001) spread, indicating a MP. MA dissertation, Hebrew University of Jerusalem. possible change in the levels of interconnectedness of social Avery, D. Margaret, David Halkett, Jayson Orton, Teresa Steel, Madelon Tus- groups (as modeled by Kandler and Laland 2009; Young enius, and Richard Klein. 2008. The Ysterfontein 1 Middle Stone Age rock shelter and the evolution of coastal foraging. South African Archaeological 2012). If a cultural and demographic “Upper Paleolithic Rev- Society Goodwin Series 10:66–89. olution” occurred, it happened during, rather than at the Bamforth, Douglas B. 1986. Technological efficiency and tool curation. Amer- beginning, of the UP. The pace of diversification and inno- ican Antiquity 51:38–50. Bamforth, Douglas B., and Peter Bleed. 1997. Technology, flaked stone, and vativeness remained relatively slow till the close of the Pleis- risk. In Rediscovering Darwin: evolutionary theory and archaeological expla- tocene. Thus most of the UP resembles the “state of affairs” nation. C. Michael Barton and Geoffrey A. Clark, eds. Pp. 109–139. Ar- observed in the local MP during which Levantine hominins chaeological Papers of the American Anthropological Association, no. 7. Arlington, VA: American Anthropological Association. occupied suboptimal social fitness peaks, getting by with Barham, Lawrence S. 2002. Systemic pigment use in the Middle Pleistocene “good enough.” It is this particular slow pace of cultural of South-Central Africa. Current Anthropology 43:181–189. change that characterizes the Levantine prehistoric record of Barkai, Ran, and Avi Gopher. 2009. Changing the face of the earth: human behavior at Sede Ilan, an extensive Lower-Middle Paleolithic quarry site in the late Middle and Upper Pleistocene and that opens up Israel. In Lithic materials and Palaeolithic societies. Brian Adams and Brooke interesting research questions about the mechanisms of cul- S. Blades, eds. Pp. 174–185. New York: Blackwell. tural transformation and stability and the historical and evo- Bar-Matthews, Miryam, Avner Ayalon, Mabs Gilmour, Alan Matthews, and Chris J. Hawkesworth. 2003. Sea-land oxygen isotopic relationships from lutionary questions underlying them. planktonic froaminifera and speleothems in the eastern Mediterranean re- gion and their implication for paleorainfall during interglacial intervals. Geochimica et Cosmochimica Acta 67:3181–3199. Bar-Yosef, Ofer. 1998. The chronology of the Middle Paleolithic of the Levant. In Neandertals and modern humans in western Asia. Takeru Akazawa, Ken- Acknowledgments ichi Aoki, and Ofer Bar-Yosef, eds. Pp. 39–56. New York: Plenum. ———. 2000. The Middle and Early Upper Paleolithic in southwest Asia and Steve Kuhn, John Speth, Christian Tryon, Yossi Zaidner, and neighboring regions. In The geography of Neandertals and modern humans two anonymous reviewers provided insightful critical com- in Europe and the greater Mediterranean. Ofer Bar-Yosef and David Pilbeam, ments on earlier drafts of this paper. eds. Pp. 107–156. Cambridge, MA: Peabody Museum of Archaeology and Ethnography. ———. 2004. Eat what is there: hunting and gathering in the world of Ne- anderthals and their neighbours. International Journal of Osteoarchaeology References Cited 14:333–342. Ahmad, G. S. 2009. The Middle Paleolithic stone tool assemblage from Ar ———. 2006. Between observations and models: an eclectic view of Middle Rasfa: reconstructing Late Pleistocene human behavior in the Jordan Rift Paleolithic archaeology. In Transitions before the transition: evolution and Valley. MA thesis, State University of New York, Stony Brook. stability in the Middle Paleolithic and Middle Stone Age. Erella Hovers and Aiello, Leslie C. 2003. Neanderthal thermo-regulation and the glacial climate. Steven L. Kuhn, eds. Pp. 305–325. New York: Springer. In Neanderthals and modern humans in Europe during the last glaciation. Bar-Yosef, Ofer, and Richard H. Meadow. 1995. The origins of agriculture in Tjeerd H. van Andel and William Davies, eds. Pp. 147–156. Cambridge: the Near East. In Last hunters, first farmers. T. Douglas Price and A.-B. McDonald Institute for Archaeological Research. Gebauer, eds. Pp. 39–94. Santa Fe, NM: School of American Research. Albert, Rosa-Maria, Ofer Bar-Yosef, Liliane Meignen, and Steve Weiner. 2003. Bar-Yosef, Ofer, and Bernard Vandermeersch. 1993. Modern humans in the Quantitative phytolith study of hearths from the Natufian and Middle Levant. Scientific American 268:94–99. Hovers and Belfer-Cohen Variability and Complexity S351

Bar-Yosef, Ofer, Bernard Vandermeersch, Baruch Arensburg, Anna Belfer- of Neanderthals to macroclimatic variations (240,000–40,000 BP). Human Cohen, Paul Goldberg, Henri Laville, Liliane Meignen, et al. 1992. The Biology 81:287–307. excavations in Kebara Cave, Mt. Carmel. Current Anthropology 33:497– Boe¨da, Eric. 1993. Le de´bitage discoı¨deetlede´bitage Levallois re´current cen- 550. tripe`te. Bulletin de la Societe´Pre´historique Franc¸aise 90:392–404. Bar-Yosef, Ofer, and Philip Van Peer. 2009. The chaıˆne ope´ratoire approach in Boe¨da, Eric, Ste´phanie Bonilauri, Jacque Connan, Dan Jarvie, Norbert Mercier, Middle Paleolithic archaeology. Current Anthropology 50:103–131. Mark Tobey, He´le`ne Valladas, Heba al Sakhel, and Sultan Muhesen. 2008. Bar-Yosef Mayer, Daniella E. 2005. The exploitation of shells as beads in the Middle Paleolithic bitumen use at Umm el Tlel around 70,000 BP. Antiquity Palaeolithic and Neolithic of the Levant. Pale´orient 31:176–185. 82:853–861. Bar-Yosef Mayer, Daniella E., Bernard Vandermeersch, and Ofer Bar-Yosef. Boe¨da, Eric, Jacques Connan, and Sultan Muhesen. 1998. Bitumen as hafting 2009. Shells and ochre in Middle Paleolithic Qafzeh Cave, Israel: indications material on Middle Paleolithic artifacts from the el-Kowm Basin, Syria. In for modern behavior. Journal of Human Evolution 56:307–314. Neandertals and modern humans in western Asia. Takeru Akazawa, Kenich Belfer-Cohen, Anna, and Ofer Bar-Yosef. 2000. Early sedentism in the Near Aoki, and Ofer Bar-Yosef, eds. Pp. 181–204. New York: Plenum. East: a bumpy ride to village life. In Life in Neolithic farming communities: Boe¨da, Eric, Jean-Michel Geneste, Christophe Griggo, Norbert Mercier, Sultan social organization, identity, and differentiation. Ian Kuijt, ed. Pp. 19–38. Muhesen, J. L. Reyss, A. Taha, and He´le`ne Valladas. 1999. A Levallois point New York: Kluwer Academic/Plenum. embedded in the vertebra of a wild ass (Equus africanus): hafting, projectiles Belfer-Cohen, Anna, and Naama Goren-Inbar. 1994. Cognition and com- and Mousterian hunting weapons. Antiquity 73:394–402. munication in the Levantine Lower Palaeolithic. World Archaeology 26:144– Boe¨da, Eric, Jean-Michel Geneste, and Liliane Meignen. 1990. Identification 157. de chaıˆne ope´ratoires lithiques du pale´olithique ancient et moyen. Pale´o 2: Belfer-Cohen, Anna, and A. Nigel Goring-Morris. 2002. Why microliths? mi- 43–80. crolithization in the Levant. In Thinking small: global perspectives on mi- Boe¨da, Eric, Christophe Griggo, and Sandrine Noe¨l-Soriano. 2001. Diffe´rents crolithization. Robert G. Elston and Steven L. Kuhn, eds. Pp. 57–68. Ar- modes d’occupation du site d’Umm el Tlel au cours du pale´olithique moyen chaeological Papers of the American Anthropological Association, no. 12. (El Kowm, Syrie centrale). Pale´orient 27:13–28. Santa Fe, NM: American Anthropological Association. Bonilauri, Ste´phanie, Eric Boe¨da, Christophe Griggo, Heba Al-Sakhel, and ———. 2007. From the beginning: Levantine Upper Palaeolithic cultural Sultan Muhesen. 2007. Un e´clat de silex mouste´rien coince dans un bassin change and continuity. In Rethinking the human revolution: new behavioural d’autruche (Struthio camelus) a Umm el Tlel (Syrie centrale). Pale´orient 33: and biological perspectives on the origin and dispersal of modern humans. 39–46. Paul Mellars, Katie Boyle, Ofer Bar-Yosef, and Chris Stringer, eds. Pp. 199– Bousman, C. Britt. 2005. Coping with risk: Later Stone Age technological 205. Cambridge: McDonald Institute for Archaeological Research. strategies at Blydefontein Rock Shelter, South Africa. Journal of Anthro- ———. 2009. The shift from the Middle Palaeolithic to the Upper Palaeolithic: pological Archaeology 24:193–226. Levantine perspectives. In The Mediterranean from 50,000 to 25,000 BP: Boyd, Robert, and Peter J. Richerson. 1985. Culture and the evolutionary turning points and new directions. Marta Camps and Carolyn C. Szmidt, process. Chicago: University of Chicago Press. eds. Pp. 89–100. Oxford: Oxbow. ———. 1996. Why culture is common, but cultural evolution is rare. Pro- Belfer-Cohen, Anna, and Erella Hovers. 1992. In the eye of the beholder: ceedings of the British Academy 88:77–93. Mousterian and Natufian burials in the Levant. Current Anthropology 33: ———. 2009. Voting with your feet: payoff biased migration and the 463–471. evolution of group beneficial behavior. Journal of Theoretical Biology ———. 2010. Modernity, enhanced working memory, and the Middle to 257:331–339. Upper Paleolithic record in the Levant. Current Anthropology 51(suppl. 1): Brooks, Alison, John E. Yellen, Lisa Nevell, and Gideon Hartman. 2006. Pro- S167–S175. jectile technologies of the African MSA: implications for modern humans Belmaker, Miriam, and Erella Hovers. 2011. Ecological change and the ex- origins. In Transitions before the transition: evolution and stability in the tinction of the Levantine Neanderthals: implications from a diachronic Middle Paleolithic and Middle Stone Age. Erella Hovers and Steven L. Kuhn, study of micromammals from Amud Cave, Israel. Quaternary Science Re- eds. Pp. 233–255. New York: Springer. views 30:3196–3209. Brown, Kyle S., Curtis W. Marean, Andy I. R. Herries, Zenobia Jacobs, Chantal Bettinger, Robert L., Robert Boyd, and Peter J. Richerson. 2009. Cultural Tribolo, David Braun, David L. Roberts, Michael C. Meyer, and Jocelyn innovations and demographic change. Human Biology 81:211–235. Bernatchez. 2009. Fire as an engineering tool of early modern humans. Beyries, Silvie. 1988. Functional variability of lithic sets in the Middle Pa- Science 325:859–862. laeolithic. In Upper Pleistocene prehistory of western Euroasia. Harold L. Cain, Chester R. 2006. Implications of the marked artifacts of the Middle Dibble and Anta Montet-White, eds. Pp. 213–225. University Museum Stone Age in Africa. Current Anthropology 47:675–681. Monograph 54. Philadelphia: University Museum, University of Pennsyl- Caˆrciumaru, Marin, Rodica-Mariana Ion, Elenna-Christina Nit¸u, and Radu vania. S¸tefa˘nescu. 2012. New evidence of adhesive as hafting material on Middle Binford, Lewis R. 1977. Forty-seven trips: a case study in the character of and Upper Palaeolithic artefacts from Gura Cheii-Raˆs¸nov Cave (Romania). archaeological site formation process. In Stone tools as cultural markers.R. Journal of Archaeological Science 39:1942–1950. V. S. Wright, ed. Pp. 24–36. Canberra: Australian Institute of Aboriginal Churchill, Steven E. 2006. Bioenergetic perspectives on Neanderthal ther- Studies. moregulatory and activity budgets. In Neanderthals revisited: new ap- ———. 1979. Organization and formation processes: looking at curated tech- proaches and perspectives. Katrina Harvati and Terry Harrison, eds. Pp. nologies. Journal of Anthropological Research 35:255–273. 113–133. Vertebrate Paleobiology and Paloeanthroplogy Series. Dor- ———. 1989. Isolating the transition to cultural adaptations: an organiza- drecht: Springer. tional approach. In The emergence of modern humans: biocultural adaptations Clark, Jamie L. 2011. The evolution of human culture during the Later Pleis- in the Late Pleistocene. Erik Trinkaus, ed. Pp. 18–41. Cambridge: Cambridge tocene: using fauna to test models on the emergence and nature of University Press. “modern” human behavior. Journal of Anthropological Archaeology 30:273– ———. 2001. Constructing frames of reference: an analytical method for ar- 291. chaeological theory building using hunter-gatherer and environmental data Collard, Mark, Briggs Buchanan, Marcus J. Hamilton, and Michael J. O’Brien. sets. Berkeley: University of California Press. 2010. Spatiotemporal dynamics of the Clovis-Folsom transition. Journal of Binford, Sally R. 1968. Early Upper Pleistocene adaptations in the Levant. Archaeological Science 37:2513–2519. American Anthropologist 70:707–717. Conard, Nicholas, M. Masri, K. Bretzkei, H. A. Naipieralai, B. Weltel, and Bisson, Michael, April Nowell, M. Poupart, C. Cordova, T. J. Pokines, and C. Andrew W. Kandel. 2010. The 2008 excavation at the Middle Paleolithic Ames. Forthcoming. WE-2: the Middle Paleolithic of the Wadi Enoqqiya site of Wadi Mushkuna Rockshelter, Damascus Province, Syria. In Docu- revisited. In Jordan’s prehistory: past and future revisited. Gary Rollefson and menting the Annual Excavation Reports concerning the archaeological activities Bill Finlayson, eds. Amman: Department of Antiquities of the Hashemite in Syria. Special issue, Chronique arche´ologique en Syrie 6:13–22. Kingdom of Jordan. Copeland, Lorraine. 1975. The Middle and Upper Palaeolithic in Lebanon Bleed, Peter. 1986. The optimal design of hunting weapons: maintainability and Syria in the light of recent research. In Problems in prehistory: North or reliability. American Antiquity 51:737–747. Africa and the Levant. Fred Wendorf and Angela Close, eds. Pp. 317–350. Bocquet-Appel, Jean-Pierre, and Alan Tuffreau. 2009. Technological responses Dallas, TX: Southern Methodist University Press. S352 Current Anthropology Volume 54, Supplement 8, December 2013

———. 1983. Levallois/non-Levallois determinations in the Early Levantine Finlayson, Geraldine, R. Barton, and Chris Stringer. 2001. The Gibraltar Ne- Mousterian: problems and questions for 1983. Pale´orient 9:15–27. anderthals and their extinction. In Les pre´miers hommes modernes de la Cordain, Loren, Janette B. Miller, S. Boyd Eaton, Neil Mann, Susanne H. pe´ninsule Ibe´rique. Joao Zilha˜o, T. Aubry, and A. Carvalho, eds. Pp. 117– Holt, and John D. Speth. 2000. Plant-animal subsistence ratios and mac- 122. Trabalhos de Arqueologia 17. Lisbon: Instituto Portugues de Arqueo- ronutrient energy estimations in worldwide hunter-gatherer diets. American logia. Journal of Clinical Nutrition 71:682–692. Friedman, Erich, Naama Goren-Inbar, Amnon Rosenfeld, Ofer Marder, and Corning, Peter A. 2002. The re-emergence of “emergence”: a venerable concept Felix Burian. 1994–1995. Hafting during Mousterian times: further evi- in search of a theory. Complexity 7:18–30. dence. Journal of the Israel Prehistoric Society 26:8–31. Corte´s-Sa´nchez, Miguel, Arturo Morales-Mun˜iz, Marı´a D. Simo´n-Vallejo, Frumkin, Amos, Ofer Bar-Yosef, and Henry P. Schwarcz. 2011. Possible pa- Marı´a C. Lozano-Francisco, Jose´ L. Vera-Pela´ez, Clive Finlayson, Joaquı´n leohydrologic and paleoclimatic effects on hominin migration and occu- Rodrı´gues-Vidal, et al. 2011. Earliest known use of marine resources by pation of the Levantine Middle Paleolithic. Journal of Human Evolution 60: Neanderthals. PLoS ONE 6:e24026. 437–451. David, Nick, and Carole Kramer. 2001. Ethnoarchaeology in action. Cambridge: Garrod, Dorothy A. E., and Dorothea M. A. Bate. 1937. The Stone Age of Cambridge University Press. Mount Carmel, vol. 1. Oxford: Clarendon. Davis, Simon J. M., Rivka Rabinovich, and Naama Goren-Inbar. 1988. Qua- Geneste, Jean-Michel. 1985. Analyse lithique d’industries mouste´riennes du ternary extinctions and population increase in western Asia: the animal Pe´rigord: une approche technologique des comportements des groupes remains from Biq’at Quneitra. Pale´orient 14:95–105. humains au pale´olithique moyen. The`se Sc., Universite´ de Bordeaux 1. DeBono, Hila, and Naama Goren-Inbar. 2001. Note on a link between Acheu- Gilead, Isaac. 1980. A Middle Palaeolithic open-air site near Tell Far’ah, west- lian handaxes and the Levallois method. Journal of the Israel Prehistoric ern Negev: preliminary report. Israel Exploration Journal 30:52–62. Society 31:9–24. ———. 1988. Le site mouste´rien de Fara II (Neguev septentrional, Israel) et Delage, Christophe, ed. 2007. Chert availability and prehistoric exploitation in le remontage de son industrie. L’Anthropologie 92:797–807. the Near East. BAR International Series 1615. Oxford: Archaeopress. Gilead, Isaac, and Caroline Grigson. 1984. Far’ah II: a Middle Palaeolithic de la Torre, Ignacio, Jorge Martı´nez-Moreno, and Rafael Mora. 2013. Change open-air site in the Northern Negev, Israel. Proceedings of the Prehistoric and stasis in the Iberian Middle Paleolithic: considerations on the signifi- Society 50:71–91. cance of Mousterian technological variability. Current Anthropology Godfrey-Smith, Dorothy I., and Shimon Ilani. 2004. Past thermal history of 54(suppl. 8):S320–S336. goethite and hematite fragments from Qafzeh Cave deduced from thermal Demidenko, Y. E., and V. I. Usik. 2004. Into the mind of the maker: refitting activation characteristics of the 110 centigrade TL peak of enclosed quartz study and technological reconstructions. In Neandertals in the Levant: be- grains. Revue d’Arche´ome´trie 28:185–190. havioral organization and the beginnings of human modernity. D. O. Henry, Goldberg, Paul, and Ofer Bar-Yosef. 1998. Site formation processes at Keabara ed. Pp. 107–155. London: Continuum. and Hayonim caves and their significance in Levantine prehistoric caves. d’Errico, Francesco, Renata Garcı´a Moreno, and Riaan F. Rifkin. 2011. Tech- In Neandertals and modern humans in western Asia. Takeru Akazawa, Ken- nological, elemental and colorimetric analysis of an engraved ochre frag- ichi Aoki, and Ofer Bar-Yosef, eds. Pp. 107–126. New York: Plenum. ment from the Middle Stone Age levels of Klasies River Cave 1, South Gopher, Avi, Ran Barkai, Ron Shimelmitz, Hamoudi Khalaily, Christina Le- Africa. Journal of Archaeological Science 39:942–952. morini, Israel Hershkovitz, and Mary C. Stiner. 2005. Qesem Cave: an Dobzhansky, Theodosius G. 1951. Genetics and human origins. New York: Amudian site in central Israel. Journal of the Israel Prehistoric Society 35: Columbia University Press. 69–92. Drake, Nick A., Roger M. Blench, Simon J. Armitage, Charlie S. Bristow, and Gordon, David. 1993. Mousterian tool selection, reduction, and discard at Kevin H. White. 2011. Ancient watercourses and biogeography of the Sahara Ghar, Israel. Journal of Field Archaeology 20:205–218. explain the peopling of the desert. Proceedings of the National Academy of Goren-Inbar, Naama. 1988. Too small to be true? reevaluation of cores Sciences of the USA 108:458–462. on flakes in Levantine Mousterian assemblages. Lithic Technology 17: Druck, Dotan. 2004. The exploitation of flint resources throughout the se- 37–44. quence of prehistoric cultures in Nahal Mea’rot, Mt. Carmel. MA thesis, ———. 1990. The lithic assemblages. In Quneitra: a Mousterian site on the University of Haifa. Golan Heights. Qedem Naama Goren-Inbar, ed. Pp. 61–167. Jerusalem: Dyson-Hudson, Rada, and Eric Alden Smith. 1978. Human territoriality: an Institute of Archaeology. ecological reassessment. American Anthropologist 80:21–41. ———. 1994. The Lower Palaeolithic of Israel. In The archaeology of society Ehrlich, Paul, and Marcus W. Feldman. 2003. Genes and cultures: what creates in the Holy Land. Thomas. E. Levy, ed. Pp. 93–109. London: Leicester our behavioral phenome? Current Anthropology 44:87–107. University Press. Ekshtain, Ravid, Moshe Inbar, Ianir Milevski, Micka Ullman, and Omry Bar- ———. 2011. Behavioral and cultural origins of Neanderthals: a Levantine izlai. 2012. Giv’at Rabi East: a new Middle Paleolithic knapping site in the perspective. Continuity and discontinuity in the peopling of Europe: one hun- Lower Galilee, Israel. Pale´orient 37:107–122. dred fifty years of Neanderthal study. Silvana Condemi and Gerd-Christian Ekshtain, Ravid, Ariel Malinsky-Buller, Shimon Ilani, Irena Segal, and Weninger, eds. Pp. 89–100. Dordrecht: Springer. Erella Hovers. 2013. Raw material exploitation around the Middle Pa- Goren-Inbar, Naama, and Anna Belfer-Cohen. 1998. The technological abil- leolithic site of ’Ein Qashish. Quaternary International, forthcoming, ities of the Levantine Mousterians: cultural and mental capacities. In Nean- doi:10.1016/j.quaint.3013.07.025. dertals and modern humans in western Asia. Takeru Akazawa, Kenichi Aoki, El Zaatari, Sireen, Frederick E. Grine, Peter S. Ungar, and Jean-Jaque Hublin. and Ofer Bar-Yosef, eds. Pp. 205–222. New York: Plenum. 2011. Ecogeographic variation in Neandertal dietary habits: evidence from Goring-Morris, Adrian N., and Anna Belfer-Cohen, eds. 2003. More than meets occlusal molar microwear texture analysis. Journal of Human Evolution 61: the eye: studies on Upper Palaeolithic diversity in the Near East. Oxford: 411–424. Oxbow. Enzel, Yehouda, Rivka Amit, Uri Dayan, Onn Crouvi, Ron Kahana, Baruch Goring-Morris, A. Nigel, Erella Hovers, and Anna Belfer-Cohen. 2009. The Ziv, and David Sharon. 2008. The climatic and physiographic controls of dynamics of Pleistocene and early Holocene settlement patterns and human the eastern Mediterranean over the late Pleistocene climates in the southern adaptations in the Levant: an overview. In Transitions in prehistory: essays Levant and its neighboring deserts. Global and Planetary Change 60:165– in honor of Ofer Bar-Yosef. John J. Shea and Daniel E. Lieberman, eds. Pp. 192. 185–252. Oxford: Oxbow. Eren, Metin I., Manuel Domı´nguez-Rodrigo, Steven L. Kuhn, Dan S. Adler, Green, Richard E., Johannes Krause, Adrian W. Briggs, Tolmislav Maricic, Ian Le, and Ofer Bar-Yosef. 2005. Defining and measuring reduction in Udo Stenzel, Martin Kircher, Nick Patterson, et al. 2010. A draft sequence unifacial stone tools. Journal of Archaeological Science 32:1190–1201. of the Neandertal genome. Science 328:710–722. Eren, Metin I., and Stephen J. Lycett. 2012. Why Levallois? a morphometric Greenfield, Patricia M. 1991. Language, tools, and brain: the ontogeny and comparison of experimental “preferential” Levallois flakes versus debitage phylogeny of hierarchically organized sequential behavior. Behavioral and flakes. PLoS ONE 7:e29273. Brain Sciences 14:531–595. Eriksson, Anders, and Andrea Manica. 2012. Effect of ancient population Griggo, Christophe. 1998. La faune mouste´rienne du site d’umm el Tlel (Syrie): structure on the degree of polymorphism shared between modern human e´tude pre´liminarie: Fouilles 1991–1993. Cahiers el l’Euphrates 8:11–25. populations and ancient hominins. Proceedings of the National Academy of Gru¨nberg, J.-M. 2002. Middle Palaeolithic birch-bark pitch. Antiquity 76:15– Sciences of the USA 109:13956–13960. 16. Hovers and Belfer-Cohen Variability and Complexity S353

Hallin, Kristin A., Margaret J. Schoeninger, and Henry P. Schwarcz. 2012. Middle Stone Age. Erella Hovers and Steven L. Kuhn, eds. Pp. 295–304. Paleoclimate during Neandertal and anatomically modern human occu- New York: Springer. pation at Amud and Qafzeh, Israel: the stable isotope data. Journal of Human ———. 2013. Insights into early mortuary practices of Homo.InThe Oxford Evolution 62:59–73. handbook of the archaeology of death and burial. Sarah Tarlow and Liv Hauck, T. C. 2011. Mousterian technology and settlement dynamics in the Nilsson-Stutz, eds. Oxford: Oxford University Press. site of Hummal (Syria). Journal of Human Evolution 61:519–537. Hovers, Erella, Shimon Ilani, Ofer Bar-Yosef, and Bernard Vandermeersch. Hauck, Thomas C., Jacques Connan, Armelle Charrie´-Duhaut, Jean-Marie Le 2003. An early case of color symbolism: ochre use by early modern humans Tensorerd, and Heba al Sakhel. 2013. Molecular evidence of bitumen in in Qafzeh Cave. Current Anthropology 44:491–522. the Mousterian lithic assemblage of Hummal (Central Syria). Journal of Hovers, Erella, William H. Kimbel, and Yoel Rak. 2000. Amud 7: still, a burial: Archaeological Science 40(8):3252–3262. http://dx.doi.org/10.1016/j.jas.2013 response to Gargett. Journal of Human Evolution 39:253–260. .03.022. Hovers, Erella, Ariel Malinsky-Buller, Mae Goder-Goldberger, and Ravid Eksh- Hayden, Brian. 1981. Subsistence and ecological adaptations of modern tain. 2011. Capturing a moment: identifying short-lived activity locations hunter-gatherers. In Omnivorous primates: gathering and hunting in human in Amud Cave, Israel. In The Lower and Middle Palaeolithic in the Middle evolution. Robert S. O. Harding and Geza Teleki, eds. Pp. 344–421. New East and neighbouring regions: Basel symposium, mai 8–10 2008. Jean-Marie York: Columbia University Press. Le Tensorer, Rheto Jagher, and Marcel Otte, eds. Pp. 101–114. Etudes et Henrich, Joseph. 2004. Demography and cultural evolution: how adaptive recherches arche´ologiques de l’Universite´de Lie`ge 126. Lie`ge: University of cultural processes can produce maladaptive losses: the Tasmania case. Amer- Lie`ge Press. ican Antiquity 69:197–214. Hovers, Erella, Yoel Rak, Ron Lavi, and William H. Kimbel. 1995. Hominid ———. 2010. The evolution of innovation-enhancing institutions. Innovations remains from Amud Cave in the context of the Levantine Middle Paleolithic. in cultural systems. Michael J. O’Brien and Stephen J. Shennan, eds. Pp. Pale´orient 21:47–61. 99–120. Cambridge MA, London England. Cambridge: MIT Press. Hovers, Erella, and Adi Raveh. 2000. The use of a multivariate display tech- Henrich, Joseph, and Robert Boyd. 1998. The evolution of conformist trans- nique in the analysis of inter-assemblage lithic variability: a case study from mission and the emergence of between-group differences. Evolution and Qafzeh Cave, Israel. Journal of Archaeological Science 27:1023–1038. Human Behavior 19:215–241. Hovers, Erella, Bernard Vandermeersch, and Ofer Bar-Yosef. 1997. A Middle Henry, Amanda G., Alison S. Brooks, and Dolores R. Piperno. 2011. Micro- Palaeolithic engraved artefact from Qafzeh Cave, Israel. Rock Art Research fossils in calculus demonstrate consumption of plants and cooked foods in 14:79–87. Neanderthal diets (Shanidar III, Iraq; Spy I and II, Belgium). Proceedings Hutchings, W. Karl. 2011. Measuring use-related fracture velocity in lithic of the National Academy of Sciences of the USA, doi:10.1073/pnas armatures to identify spears, javelins, darts, and arrows. Journal of Archae- .1016868108. ological Science 38:1737–1746. Henry, Donald O. 1995a. The influence of mobility levels on Levallois point Inizan, Marie-Louise, Helene Roche, and Jacques Tixier. 1992. Technology production, Late Levantine Mousterian, southern Jordan. In The definition of knapping stone: pre´histoire de la Pierre Taille´e 3. Meudon, France: and interpretation of Levallois technology. Harold L. Dibble and Ofer Bar- CREP. Yosef, eds. Pp. 185–200. Madison, WI: Prehistory Press. Jelinek, Arthur. 1982. The Middle Paleolithic in the southern Levant, with ———. 1995b. The Middle Paleolithic sites. In Prehistoric cultural ecology and comments on the appearance of modern Homo sapiens.InThe transition diversity: insights from southern Jordan. Donald O. Henry, ed. Pp. 49–84. from Lower to Middle Palaeolithic and the origin of modern man. Avraham New York: Plenum. Ronen, ed. Pp. 57–104. BAR International Series 151. Oxford: BAR. Henry, D. O., H. J. Hietala, A. M. Rosen, Y. E. Demidenko, V. I. Usik, Jelinek, Arthur J., William Farrand, George Haas, Aharon Horowitz, and Paul and T. L. Armagan. 2004. Human behavioral organization in the Middle Goldberg. 1973. New excavations at the Tabun Cave, Mount Carmel, Israel, Paleolithic: were Neanderthals different? American Anthropologist 106: 17–31. 1967–1972: a preliminary report. Pale´orient 1:151–183. Hietala, H. J. 2003. Site structure and material culture in space on the Tor Jochim, Michael B. 1983. Palaeolithic cave art in ecological perspective. In Faraj living floors. In Neandertals in the Levant: behavioral organization and hunter-gatherer economy in prehistory: a European perspective. Geoff Bailey, the beginnings of human modernity. Donald O. Henry, ed. Pp. 198–233. ed. Pp. 212–219. Cambridge: Cambridge University Press. London: Continuum. Johnson, Gregory A. 1978. Information sources and the development of Hockett, Bryan. 2012. The consequences of Middle Paleolithic diets on preg- decision-making organization. In Social archaeology: beyond subsistence and nant Neanderthal women. Quaternary International 264:78–82. dating. Charles L. Redman, M. J. Berman, E. V. Curtin, W. J. Longhorne Hovers, Erella. 1997. Variability of lithic assemblages and settlement patterns Jr., N. M. Versaggi, and C. Wansen, eds. Pp. 87–112. New York: Academic in the Levantine Middle Paleolithic: implications for the development of Press. human behavior. PhD dissertation, Hebrew University of Jerusalem. Jordan, P. 2007. Continuity and change in different domains of culture: an ———. 1998a. Comment on “Neandertal and early modern human behav- emerging approach to understanding diversity in technological traditions. ioral variability: a regional-scale approach to lithic evidence for hunting in In The model-based archaeology of socionatural ecosystems. T. A. Kohler and the Levantine Mousterian” by J. J. Shea. Current Anthropology 39(suppl.): S. E. van der Leeuw, eds. Pp. 13–39. Santa Fe, NM: School for Advanced S65–S66. Research. ———. 1998b. The lithic assemblages of Amud Cave: implications for the Kandler, Anne, and Kevin N. Laland. 2009. An investigation of the relationship end of the Mousterian in the Levant. In Neandertals and modern humans between innovation and cultural diversity. Theoretical Population Biology in western Asia. Takeru Akazawa, Kenichi Aoki, and Ofer Bar-Yosef, eds. 76:59–67. Pp. 143–163. New York: Plenum. Karkanas, Panagiotis, Ruth Shahack-Gross, Avner Ayalon, Miryam Bar- ———. 2001. Territorial behavior in the Middle Paleolithic of the Southern Matthews, Ran Barkai, Amos Frumkin, Avi Gopher, and Mary C. Stiner. Levant. In Settlement dynamics of the Middle Paleolithic and Middle Stone 2007. Evidence for habitual use of fire at the end of the Lower Paleolithic: Age . Nicholas Conard, ed. Pp. 123–152. Tu¨bingen Publications in Prehis- site-formation processes at Qesem Cave, Israel. Journal of Human Evolution tory. Tu¨bingen: Kerns. 53:197–212. ———. 2006. Neandertals and modern humans in the Middle Paleolithic of Kelly, Robert L. 1983. Hunter-gatherer mobility strategies. Journal of Anthro- the Levant: what kind of interaction? In When Neandertals and moderns pological Research 39:277–306. met. Nicholas Conard, ed. Pp. 65–86. Tu¨bingen: Kerns. ———. 1995. The foraging spectrum: diversity in hunter-gatherer lifeways. ———. 2007. The many faces of cores-on-flakes: a perspective from the Washington, DC: Smithsonian Institution. Levantine Mousterian. In Cores or tools? alternative approaches to stone tool Klein, Richard G., and K. Cruz-Uribe. 1996. Exploitation of large bovids and analysis. Shannon P. McPherron, ed. Pp. 42–74. Cambridge: Cambridge seals at Middle and Later Stone Age sites in South Africa. Journal of Human Scholars Press. Evolution 31:315–334. ———. 2009. The lithic assemblages of Qafzeh Cave. New York: Oxford Uni- Kuhn, Steven L. 1993. Mousterian technology as adaptive response: a case versity Press. study. In Hunting and animal exploitation in the late Palaeolithic and Hovers, Erella, and Anna Belfer-Cohen. 2006. “Now you see it, now you Mesolithic of Europe. Gail H. Peterkin, Harvey M. Bricker, and Paul don’t”: modern human behavior in the Middle Paleolithic. In Transitions Mellars, eds. Pp. 20–25. Washington, DC: American Anthropological before the transition: evolution and stability in the Middle Paleolithic and Association. S354 Current Anthropology Volume 54, Supplement 8, December 2013

———. 1994. A formal approach to the design and assembly of transported and paleoenvironments in the central Negev, Israel. Anthony E. Marks, ed. toolkits. American Antiquity 59:426–442. Pp. 131–158. Dallas, TX: Southern Methodist University Press. ———. 1995. Mousterian lithic technology: an ecological perspective. Princeton, Marshack, Alexander. 1981. On Paleolithic ochre and the early uses of color NJ: Princeton University Press. and symbol. Current Anthropology 22:188–191. ———. 2013. Roots of the Middle Paleolithic in Eurasia. Current Anthropology ———. 1996. A Middle Paleolithic symbolic composition from the Golan 54(suppl. 8):S255–S268. Heights: the earliest known depictive image. Current Anthropology 37:357– Kuhn, Steven L., and Mary C. Stiner. 1998. Middle Paleolithic “creativity”: 365. reflections on an oxymoron? In Creativity in human evolution and prehistory. Marx, Emmanuel. 2004. Bands and other corporate societies in Acheulian Steven Mithen, ed. Pp. 143–164. London: Routledge. culture. In Human paleoecology in the Levantine corridor. Naama Goren- ———. 2006. What’s a mother to do? the division of labor among Ne- Inbar and John D. Speth, eds. Pp. 89–104. Oxford: Oxbow. anderthals and modern humans in Eurasia. Current Anthropology 47: Matsutani, A. 1973. Microscopic study on the “amorphous silica” in sediments 953–980. from the Douara Cave. In The Paleolithic site of Doura Cave in Syria, part Kuhn, Steven L., Mary C. Stiner, David S. Reese, and Erskin Gu¨lec¸. 2001. I. Hisashi Suzuki and Fuyuji Takai, eds. Pp. 17–21. Tokyo: University of Ornaments of the earliest Upper Paleolithic: new insights from the Tokyo Press. Levant. Proceedings of the National Academy of Sciences of the USA 98: ———. 1987. Plant remains from the 1984 excavations at Douara Cave. In 7641–7646. Paleolithic site of Douara Cave and paleogeography of Palmyra Basin, part Lahr, Martha M., and Robert Foley. 1998. Towards a theory of modern human IV: 1984 excavations. Takeru Akazawa and Y. Sakaguchi, eds. Pp. 117–122. origins: geography, demography, and diversity in recent human evolution. Tokyo: University of Tokyo Press. Yearbook of Physical Anthropology 41:137–176. Mazza, Paul P. A., Fabio Martini, Benedeto Sala, Maurizio Magi, Maria P. Laland, Kevin, and Michael O’Brien. 2010. Niche construction theory and Colombini, Gianna Giachi, Francesco Landucci, Christina Lemorini, Fran- archaeology. Journal of Archaeological Method and Theory 17:303–322. cesco Modugno, and Erika Ribechini. 2006. A new Palaeolithic discovery: Langbroek, Marco. 2012. Trees and ladders: a critique of the theory of human tar-hafted stone tools in a European Mid-Pleistocene bone-bearing bed. cognitive and behavioural evolution in Palaeolithic archaeology. Quaternary Journal of Archaeological Science 33:1310–1318. International 270:4–14. McBrearty, Sally, and Christian A. Tryon. 2006. From Acheulean to Middle Lev, Ephraim, Mordechai E. Kislev, and Ofer Bar-Yosef. 2005. Mousterian Stone Age in the Kapthurin Formation, Kenya. In Transitions before the vegetal food in Kebara Cave, Mt. Carmel. Journal of Archaeological Science transition: evolution and stability in the Middle Paleolithic and Middle Stone 32:475–484. Age. Erella Hovers and Steven L. Kuhn, eds. Pp. 257–277. New York: Lieberman, Daniel E., and Ofer Bar-Yosef. 2005. Apples and oranges: mor- Springer. phological versus behavioral transitions in the Pleistocene. In Interpreting Meignen, Liliane. 1995. Levallois lithic production systems in the Middle the past: essays in human, primate, and mammal evolution in honor of David Palaeolithic of the Near East: the case of the unidirectional method. In The Pilbeam. Daniel E. Lieberman, Richard J. Smith, and Jay Kelley, eds. Pp. definition and interpretation of Levallois technology. Harold L. Dibble and Ofer Bar-Yosef, eds. Pp. 361–380. Madison, WI: Prehistory Press. 275–296. Boston: Bril Academic. ———. 1998. A preliminary report on Hayonim cave lithic assemblages in Lombard, Marlize. 2007. The gripping nature of ochre: the association of the context of the Near Eastern Middle Palaeolithic. In Neandertals and ochre with Howiesons Poort adhesives and Later Stone Age mastics from modern humans in western Asia. Takeru Akazawa, Kenichi Aoki, and Ofer South Africa. Journal of Human Evolution 53:406–419. Bar-Yosef, eds. Pp. 165–180. New York: Plenum. MacDonald, Katharine, Will Roebroeks, and Alexander Verpoort. 2009. An ———. 2011. The contribution of Hayonim Cave assemblage to the under- energetics perspective on the Neandertal record. In The evolution of hominin standing of the so-called Early Levantine Mousterian. In The Lower and diets: integrating approaches to the study of palaeolithic subsistence. Jean- Middle Palaeolithic in the Middle East and neighbouring regions: Basel sym- Jacque Hublin and Michael P. Richards, eds. Pp. 211–220. Vertebrate Pale- posium, mai 8–10 2008. Jean-Marie Le Tensorer, Rheto Jagher, and Marcel obiology and Paleoanthropology. Dordrecht: Springer. Otte, eds. Pp. 85–100. Etudes et recherches arche´ologiques de l’Universite´ Mackay, Alex. 2011. Potentially stylistic differences between backed artefacts de Lie`ge 126. Lie`ge: University of Lie`ge Press. from two nearby sites occupied 60,000 years before present in South Africa. Meignen, Liliane, and Ofer Bar-Yosef. 1992. Middle Paleolithic variability in Journal of Anthropological Archaeology 30:235–245. Kebara Cave, Israel. In The evolution and dispersal of modern humans in Madella, Marco, Martin K. Jones, Paul Goldberg, Yuval Goren, and Erella Asia. Takeru Akazawa, Kenichi Aoki, and Tatsuku Kimura, eds. Pp. 129– Hovers. 2002. The exploitation of plant resources by Neanderthals in Amud 148. Tokyo: Hakusen-Sha. Cave (Israel): the evidence from phytolith studies. Journal of Archaeological ———. 2002. The lithic industries of the Middle and Upper Paleolithic of Science 29:703–719. the Levant: continuity or break? Archaeology, Ethnology and Anthropology Malinsky-Buller, Ariel, Leore Grosman, and Ofer Marder. 2011. A case of of Eurasia 3:12–21. techno-typological lithic variability and continuity in the late Lower Pa- Meignen, Liliane, Ofer Bar-Yosef, John D. Speth, and Mary C. Stiner. 2006. laeolithic. Before Farming 2011/1(art. 3):1–32. Middle Paleolithic settlement patterns in the Levant. In Transitions before March, Ramiro, Alexandre Lucquin, Delphine Joly, Juan Ferreri, and Mo- the transition: evolution and stability in the Middle Paleolithic and Middle hamad Muhieddine. 2012. Processes of formation and alteration of ar- Stone Age. Erella Hovers and Steven L. Kuhn, eds. Pp. 149–169. New York: chaeological fire structures: complexity viewed in the light of experimental Springer. approaches. Journal of Archaeological Method and Theory, doi:10.1007 Meignen, Liliane, Paul Goldberg, and Ofer Bar-Yosef. 2007. The hearths at /s10816-012-9134-7. Kebara Cave and their role in site formation processes. In Kebara Cave, Marder, Ofer, Reuven Yeshurun, Ronit Lupu, Guy Bar-Oz, Miriam Bel- Mt. Carmel, Israel: the Middle and Upper Paleolithic archaeology, part I.Ofer maker, Naomi Porat, Hagai Ron, and Amos Frumkin. 2011. Mammal Bar-Yosef and Liliane Meignen, eds. Pp. 91–122. Cambridge, MA: American remains at Rantis Cave, Israel, and Middle-Late Pleistocene human School of Prehistoric Research, Peabody Museum of Anthropology and subsistence and ecology in the southern Levant. Journal of Quaternary Ethnology, Harvard University. Science 26:769–780. Mesoudi, Alex. 2011a. Cultural evolution: how Darwinian theory can explain Marean, Curtis W., Miryam Bar-Matthews, J. Bernatchez, Erich Fisher, Paul human culture and synthesize the social sciences. Chicago: University of Chi- Goldberg, Andy I. R. Herries, Zenobia Jacobs, et al. 2007. Early human use cago Press. of marine resources and pigment in South Africa during the Middle Pleis- ———. 2011b. Variable cultural acquisition costs constrain cumulative cul- tocene. Nature 449:905–908. tural evolution. PLoS ONE 6:e18239. Marks, Anthony E. 1988. The curation of stone tools during the Upper Pleis- Mesoudi, Alex, and Stephen J. Lycett. 2009. Random copying, frequency- tocene: a view from the central Negev, Israel. In Upper Pleistocene prehistory dependent copying and culture change. Evolution and Human Behavior 30: of western Eurasia. Harold L. Dibble and Anta Montet-White, eds. Pp. 275– 41–48. 285. Philadelphia: University Museum, University of Pennsylvania. Metcalfe, D., and K. R. Barlow. 1992. A model for exploring the optimal trade- ———. 2008. Into Arabia, perhaps, but if so, from where? Proceedings of the off between field processing and transport. American Anthropologist 94:340– Seminar for Arabian Studies 38:15–24. 356. Marks, Anthony E., and David A. Freidel. 1977. Prehistoric settlement patterns Moncel, Marie-He´le´n, Anne-Marie Moigne, Youssef Sam, and Jean Combier. in the Avdat/Aqev area. In The Avdat/Aqev area, part 2, vol. 2 of Prehistory 2011. The emergence of Neanderthal technical behavior: new evidence from Hovers and Belfer-Cohen Variability and Complexity S355

Orgnac 3 (Level 1, MIS 8), southeastern France. Current Anthropology 52: Rabinovich, Rivka, and Eitan Tchernov. 1995. Chronological, paleoecological 37–75. and taphonomical aspects of the Middle Paleolithic site of Qafzeh, Israel. Monigal, Katherine. 2002. The Levantine leptolithic: blade production from In Archaeozoology of the Near East: proceedings of the second symposium of the Lower Paleolithic to the dawn of the Upper Paleolithic. PhD dissertation, the archaeozoology of southwestern Asia and adjacent areas. H. Buitenhuis Southern Methodist University, Dallas, TX. and Hans-Peter Uerpmann, eds. Pp. 5–44. Leiden: Backhuys. Moore, James A. 1981. The effects of information networks in hunter-gatherer Renfrew, Colin. 1978. The anatomy of innovation. In Approaches to social societies. In Hunter-gatherer foraging strategies: ethnographic and archaeo- archaeology. Colin Renfrew, ed. Pp. 390–418. Cambridge, MA: Harvard logical case studies. Bruce Winterhalder and Eric A. Smith, eds. Pp. 194– University Press. 217. Prehistoric Archeology and Ecology. Chicago: University of Chicago Reynolds, Robert G. 1978. On modeling the evolution of hunter-gatherer Press. decision-making systems. Geographical Analysis 10:31–46. Mourre, Vincent, Paola Villa, and Christopher S. Henshilwood. 2010. Early Richerson, Peter J., and Robert Boyd. 2005. Not by genes alone: how culture use of pressure flaking on lithic artifacts at Blombos Cave, South Africa. transformed human evolution. Chicago: University of Chicago Press. Science 330:659–662. Roebroeks, Wil, Mark J. Sier, Trine K. Nielsen, Dimitri De Loecker, Josep M. Munday, Frederick. C. 1976. Intersite variability in the Mousterian of the Pare´sb, Charles E. S. Arps, and Herman J. Mu¨cher. 2012. Use of red ochre Central Negev. In The Avdat/Aqev area, part 1, vol. 1 of Prehistory and by early Neandertals. Proceedings of the National Academy of Sciences of the paleoenvironments in the central Negev, Israel. Anthony E. Marks, ed. Pp. USA 109:1889–1894. 113–140. Dallas, TX: Southern Methodist University Press. Roebroeks, Wil, and Paola Villa. 2011. On the earliest evidence for habitual Neiman, Frederick D. 1995. Stylistic variation in evolutionary perspective: use of fire in Europe. Proceedings of the National Academy of Sciences of the inferences from decorative diversity and interassemblage distance in Illinois USA 108:5209–5214. woodland ceramic assemblages. American Antiquity 60:7–36. Rohling, Eelco J., Katharine M. Grant, Andrew P. Roberts, and Juan-Cruz Nelson, Margaret C. 1991. The study of technological organization. In Ar- Larrasoan˜a. 2013. Paleoclimate variability in the Mediterranean and Red chaeological method and theory, vol. 3. Michael B. Schiffer, ed. Pp. 57–100. Sea regions during the last 500,000 years: implications for hominin migra- Tucson: University of Arizona Press. tions. Current Anthropology 54(suppl. 8):S183–S201. Nilsson, Liv. 1998. Dynamic cadavers: a “field-anthropological” analysis of the Rolland, Nicholas. 2004. Was the emergence of home bases and domestic fire Skateholm II burials. Lund Archaeological Review 4:5–17. a punctuated event? a review of the Middle Pleistocene record in Eurasia. Nishiaki, Yoshihiro. 1989. Early blade industries in the Levant: the placement Asian Perspectives 43:248–280. of Douara IV industry in the context of the Levantine Early Middle Pa- Ronen, Avraham. 1979. Paleolithic industries. In The Quaternary of Israel. leolithic. Pale´orient 15:215–229. Aharon Horowitz, ed. Pp. 296–307. New York: Academic Press. Nishiaki, Yoshihiro, and Lorraine Copeland. 1992. Keoue Cave, northern Leb- Rossano, Matt J. 2010. Making friends, making tools, and making symbols. anon, and its place in the context of the Levantine Mousterian. In The Current Anthropology 51(suppl.):S89–S98. evolution and dispersal of modern humans in Asia. Takeru Akazawa, Kenichi Rots, Veerle, and Philip Van Peer. 2006. Early evidence of complexity in lithic Aoki, and Tatsuku Kimura, eds. Pp. 107–127. Tokyo: Hakusen-sha. economy: core-axe production, hafting and use at Late Middle Pleistocene Nowell, April, Francesco d’Errico, and Erella Hovers. 2001. The origin of site 8-B-11, Sai Island (Sudan). Journal of Archaeological Science 33:360– symbolism in the Near East. Abstracts of the 66th annual meeting, April 18- 371. 22, 2001, New Orleans, Louisiana. Washington, DC: Society for American Rots, Veerle, Philip Van Peer, and Pierre M. Vermeersch. 2011. Aspects of Archaeology. tool production, use, and hafting in Palaeolithic assemblages from Northeast Odell, George H. 1996. Economizing behavior and the concept of curation. Africa. Journal of Human Evolution 60:637–664. In Stone tools: theoretical insights into human prehistory. George H. Odell, Salomon, He´le`ne, C. Vignaud, Y. Coquinot, L. Beck, Chris Stringer, D. Strivay, eds. Pp. 51–80. New York: Plenum. and Frencesco D’Errico. 2012. Selection and heating of colouring materials Oron, Maya, and Naama Goren-Inbar. 2013. Mousterian intra-site spatial in the Mousterian level of es-Skhul (c. 100,000 years BP, Mt. Carmel, Israel. patterning at Quneitra, Golan Heights. Quaternary International. http:// Archaeometry 54:698–722. dx.doi.org/10.1016/j.quaint.2013.04.013. Sandgathe, Dennis M., Harold L. Dibble, Paul Goldberg, Shannon P. Mc- O’Shea, John. 1981. Coping with scarcity: exchange and social storage. In Pherron, Alain Turq, Laura Niven, and Jamie Hodgkins. 2011. On the role Economic archaeology: towards an integration of economic and social ap- of fire in Neandertal adaptations in western Europe: evidence from Pech proaches. A. Sheridan and Geoff Bailey, eds. Pp. 167–183. BAR International de l’Aze´and Roc de Marsal, France. Paleoanthropology 2011:216–242. Series 96. Oxford: BAR. Schick, Tamar, and Moshe Stekelis. 1977. Mousterian assemblages in Kebara Oswalt, Wendel H. 1976. An anthropological analysis of food-getting technology. Cave, Israel. In Moshe Stekelis memorial volume: Eretz-Israe, vol. 13. Baruch New York: Wiley. Arensburg and Ofer Bar-Yosef, eds. Pp. 97–149. Jerusalem: Israel Explo- Parry, William J., and Robert L. Kelly. 1987. Expedient core technology and ration Society. sedentism. In The organization of core technology. J. K. Johnson and Tobi Shahack-Gross, Ruth, Avner Ayalon, Paul Goldberg, Yuval Goren, Boaz Ofek, A. Morrow, eds. Pp. 285–309. Boulder, CO: Westview. Rivka Rabinovich, and Erella Hovers. 2008. Formation processes of ce- Pawlik, Alfred F., and J. P. Thissen. 2011. Hafted armatures and multi-com- mented features in karstic cave sites revealed using stable oxygen and carbon ponent tool design at the Micoquian site of Inden-Altdorf, Germany. Journal isotopic analyses: a case study at Middle Paleolithic Amud Cave, Israel. of Archaeological Science 38:1699–1708. Geoarchaeology 23:43–62. Pettitt, Paul. 2010. The Palaeolithic origins of human burial. London: Routledge. Sharon, Gonen, Leore Grosman, Hannah Fluck, Yoel Melamed, Yoel Rak, Plisson, Hugue, and Silvie Beyries. 1998. Pointes ou outils triangulaires? done´es Rivka Rabinovich, and Maya Oron. 2010. The first two excavation seasons fonctionnelles dans le mouste´rian levantin. Pale´orient 24:5–16, 23–24. at NMO: a Mousterian site at the bank of the Jordan river. Eurasian Pre- Powell, Adam, Stephen Shennan, and Mark G. Thomas. 2009. Late Pleistocene history 7:129–151. demography and the appearance of modern human behavior. Science 324: Sharon, Gonen, and Maya Oron. 2013. The lithic tool arsenal of a Mousterian 1298–1301. hunter. Quaternary International, forthcoming, doi:10.1016/j.quaint.2013 Premo, Lukas S., and Steven L. Kuhn. 2010. Modeling effects of local ex- .10.024. tinctions on culture change and diversity in the Paleolithic. PLoS ONE 5: Shea, John J. 1988. Spear points from the Middle Paleolithic of the Levant. e15582, doi:10.1671/journal.pone.0015582. Journal of Field Archaeology 15:441–450. Pulliam, H. Ronald, and Christopher Dunford. 1980. Programmed to ———. 1991. The behavioral significance of Levantine Mousterian variability. learn: an essay on the evolution of culture. New York: Columbia Uni- PhD dissertation, Harvard University. versity Press. ———. 1998. Neandertal and early modern human behavioral variability: a Rabinovich, Rivka. 2003. The Levantine Upper Palaeolithic faunal record. In regional scale approach to lithic evidence for hunting in the Levantine More than meets the eye: studies on Upper Palaeolithic diversity in the Near Mousterian. Current Anthropology 39(suppl.):S45–S78. East. A. Nigel Goring-Morris and Anna Belfer-Cohen, eds. Pp. 33–48. Ox- ———. 2003. Neandertals, competition, and the origin of modern human ford: Oxbow. behavior in the Levant. Evolutionary Anthropology 12:173–187. Rabinovich, Rivka, and Erella Hovers. 2004. Faunal assemblages from Amud ———. 2004. The Middle Paleolithic of the East Mediterranean Levant. Jour- Cave: preliminary results and interpretations. International Journal of Os- nal of World Prehistory 17:313–394. teoarchaeology 14:287–306. ———. 2006. The origins of lithic projectile point technology: evidence from S356 Current Anthropology Volume 54, Supplement 8, December 2013

Africa, the Levant, and Europe. Journal of Archaeological Science 33:823– of marine mammals in Gibraltar. Proceedings of the National Academy of 846. Sciences of the USA 105:14319–14324. ———. 2008. Transitions or turnovers? climatically-forced extinctions of Texier, Pierre-Jean, Guillaume Porraz, John Parkington, Jean-Phillipe Rigaud, Homo sapiens and Neanderthals in the East Mediterranean Levant. Qua- Cedric Poggenpoel, Christopher Miller, Chantal Tribolo, et al. 2010. A How- ternary Science Reviews 27:2253–2270. iesons Poort tradition of engraving ostrich eggshell containers dated to Shea, John J., Zachary Davis, and Kyle Brown. 2001. Experimental tests of 60,000 years ago at Diepkloof Rock Shelter, South Africa. Proceedings of the Middle Paleolithic spear points using a calibrated crossbow. Journal of Ar- National Academy of Sciences of the USA 107:6180–6185. chaeological Science 28:807–816. Thie´baut, Celine, E´ milie Claud, Vincent Mourre, Maria-Gema Chacon, Guil- Shennan, Stephen. 2001. Demography and cultural innovation: a model and laume Asselin, Michel Brenet, and Benoıˆt Paravel. 2010. The recycling and its implications for the emergence of modern human culture. Cambridge reuse of cores and bifaces during the Middle Paleolithic in western Europe. Archaeological Journal 11:5–16. Palethnologie 2, 2010. Shennan, Stephen, and James Steele. 2009. Introduction: demography and Thieme, Hartmut. 1997. Lower Palaeolithic hunting spears from Germany. cultural macroevolution. Human Biology 81:105–119. Nature 385:807–810. Shimelmitz, Ron, Ran Barkai, and Avi Gopher. 2011. Systematic blade pro- Tillier, Anne-Marie. 1990. Une controverse de´passee´: l’existence de pra- duction at late Lower Paleolithic (400–200 kyr) Qesem Cave, Israel. Journal tiques fune´raires au pale´olithique moyen. Les Nouvelles de l’Arche´ologie of Human Evolution 61:458–479. 40:22–24. Shott, Michael. 1989. On tool-class use lives and the formation of archaeo- Tryon, Christian A., and J. Tyler Faith. 2013. Variability in the Middle logical assemblages. American Antiquity 54:9–30. Stone Age of eastern Africa. Current Anthropology 54(suppl. 8):S234– Skinner, J. H. 1970. El Masloukh: a Yabrudian site in Lebanon. Bulletin du S254. Muse´e de Beyrouth 23:143–172. Tryon, Christian A., Sally McBrearty, and Pierre-Jean Texier. 2005. Le- Soressi, Marie. 2005. Late Mousterian lithic technology: its implications for vallois lithic technology from the Kapthurin Formation, Kenya: Acheu- the pace of the emergence of behavioural modernity and the relationship lian origin and Middle Stone Age diversity. African Archaeological Re- between behavioural modernity and biological modernity. In From tools to view 22:199–229. symbols. Lucinda Backwell and Francesco d’Errico, eds. Pp. 389–417. Jo- Tuffreau, Aalin. 1995. The variability of Levallois technology in northern hannesburg: Witwatersrand University Press. France and neighbouring areas. In The definition and interpretation of Le- Speth, John D. 2004. Hunting pressure, subsistence intensification, and de- vallois technology. Harold L. Dibble and Ofer Bar-Yosef, eds. Pp. 413–427. mographic change in the Levantine Late Middle Paleolithic. In Human Madison, WI: Prehistory Press. paleoecology in the Levantine corridor. Naama Goren-Inbar and John D. Ugan, Andrew, Jason Bright, and Alan Rogers. 2003. When is technology Speth, eds. Pp. 149–166. Oxford: Oxbow. worth the trouble? Journal of Archaeological Science 30:1315–1329. ———. 2006. Housekeeping, Neanderthal-style: hearth placement and Vaesen, Krist. 2012. Cumulative cultural evolution and demography. PLoS midden formation in Kebara Cave (Israel). In Transitions before the ONE 7:e40989. transition: evolution and stability in the Middle Paleolithic and Middle Vaks, Anton, Miryam Bar-Matthews, Avner Ayalon, Alan Matthews, Lud- Stone Age. Erella Hovers and Steven L. Kuhn, eds. Pp. 171–188. New wig Halicz, and Amos Frumkin. 2007. Desert speleothems reveal cli- York: Springer. matic window for African exodus of early modern humans. Geology ———. 2010. The paleoanthropology and archaeology of big-game hunting: 35:831–834. protein, fat, or politics? New York: Springer. Vanhaeren, Marian, Francesco d’Errico, Chris Stringer, Sarah L. James, Jon- ———. 2012. Middle Palaeolithic subsistence in the Near East: zooarchae- athan A. Todd, and Henk K. Mienis. 2006. Middle Paleolithic shell beads ological perspectives: past, present, and future. Before Farming 2012(2), art. in Israel and Algeria. Science 312:1785–1788. 1. Van Peer, Philip. 1992. The Levallois reduction strategy. Monographs in World Speth, John D., and Jamie Clark. 2006. Hunting and overhunting in the Archaeology 13. Madison, WI: Prehistory Press. Levantine Late Middle Palaeolithic. Before Farming 2006(3), art. 1. Villa, Paola, Paolo Boscato, Filomena Ranaldo, and Annamaria Ronchitelli. Speth, John D., Liliane Meignen, Ofer Bar-Yosef, and Paul Goldberg. 2012. 2009. Stone tools for the hunt: points with impact scars from a Middle Spatial organization of Middle Paleolithic occupation X in Kebara Cave Paleolithic site in southern Italy. Journal of Archaeological Science 36:850– (Israel): concentrations of animal bones. Quaternary International 247:85– 859. 102. Wadley, Lyn. 2005. Putting ochre to the test: replication studies of adhesives Speth, John D., and Eitan Tchernov. 2007. The Middle Paleolithic occupations that may have been used for hafting tools in the Middle Stone Age. Journal at Kebara Cave: a faunal perspective. In The Middle and Upper Paleolithic of Human Evolution 49:587–601. archaeology of Kebara Cave, part I. Ofer Bar-Yosef and Liliane Meignen, eds. Wadley, Lyn, Tamaryin Hodgskiss, and Michael Grant. 2009. Implications for Pp. 165–260. Cambridge, MA: Peabody Museum of Archaeology and Eth- complex cognition from the hafting of tools with compound adhesives in nology, Harvard University. the Middle Stone Age, South Africa. Proceedings of the National Academy Steegmann, A. Theodore, Jr., Frank J. Cerny, and Trenton W. Holliday. 2002. of Sciences of the USA 106:9590–9594. Neandertal cold adaptation: physiological and energetic factors. American Wallace, Ian J., and John J. Shea. 2006. Mobility patterns and core technologies Journal of Human Biology 14:566–583. in the Middle Paleolithic of the Levant. Journal of Archaeological Science 33: Sterelny, Kim. 2012. The evolved apprentice: how evolution made humans 1293–1309. unique. Cambridge, MA: MIT Press. Walter, Phillipe. 2003. Characte´risation des traces rouges et noires sur les Stiner, Mary C. 1994. Honor among thieves: a zooarchaeological study of Nean- coquillage perfore´s de Qafzeh. In E´changes et diffusion dans la pre´histoire dertal ecology. Princeton, NJ: Princeton University Press. me´diterrane´enne. Bernard Vandermeerch, ed. Pp. 122. Paris: E´ ditions du ———. 2005. The faunas of Hayonim Cave, Israel: a 200,000-year record of Comite´ des travaux historiques et scientifiques. paleolithic diet, demography, and society. American School of Prehistoric Weiner, Steve, Paul Goldberg, and Ofer Bar-Yosef. 2002. Three-dimensional Research Bulletin 48. Cambridge, MA: Peabody Museum of Archaeology distribution of minerals in the sediments of Hayonim Cave, Israel: diage- and Ethnology, Harvard University. netic processes and archaeological implications. Journal of Archaeological Stiner, Mary C. 2013. An unshakable Middle Paleolithic? trends versus con- Science 29:1289–1308. servatism in the predatory niche and their social ramifications. Current Weinstein-Evron, Mina, Guy Bar-Oz, Yossi Zaidner, Alexander Tsatskin, Dotan Anthropology 54(suppl. 8):S288–S304. Druck, Naomi Porat, and Israel Hershkovitz. 2003. Introducing Misliya Stiner, Mary C., Avi Gopher, and Ran Barkai. 2011. Hearth-side socioeco- Cave, Mount Carmel, Israel: a new continuous Lower/Middle Paleolithic nomics, hunting and paleoecology during the late Lower Paleolithic at sequence in the Levant. Eurasian Prehistory 1:31–55. Qesem Cave, Israel. Journal of Human Evolution 60:213–233. Whallon, Robert, William Lovis, and Robert Hitchcock, eds. 2011. Information Stiner, Mary C., Natalie D. Munro, and Todd A. Surovell. 2000. The tortoise and its role in hunter-gatherer bands. Los Angeles: UCLA Cotsen Institute and the hare: small-game use, the broad-spectrum revolution, and Paleo- for Archaeology. lithic demography. Current Anthropology 41:39–73. White, Mark, Nick Ashton, and B. Scott. 2011. The emergence, diversity and Stringer, Chris B., J. C. Finlayson, R. N. E. Barton, Yolanda Ferna´ndez-Jalvo, significance of Mode 3 (prepared core) technologies. Developments in Qua- I. Ca´ceres, R. C. Sabin, E. J. Rhodes, et al. 2008. Neanderthal exploitation ternary Science 14:53–65. Hovers and Belfer-Cohen Variability and Complexity S357

Wobst, H. Martin. 1974. Boundary conditions for Paleolithic social systems: Yeshurun, Reuven, Guy Bar-Oz, and Mina Weinstein-Evron. 2007. Mod- a simulation approach. American Antiquity 39:147–178. ern hunting behavior in the Early Middle Paleolithic: faunal remains ———. 1976. Locational relationships in Paleolithic society. Journal of Human from Misliya Cave, Mount Carmel, Israel. Journal of Human Evolution Evolution 5:49–58. 53:656–677. Wojtzak, Dorothea. 2011. Hummal (central Syria) and it eponymous industry. Young, H. Peyton. 2012. The dynamics of social innovation. Proceedings of In The Lower and Middle Palaeolithic in the Middle East and neighbouring the National Academy of Sciences of the USA 108:21285–21291. regions: Basel symposium, mai 8–10 2008. Jean-Marie Le Tensorer, Rheto Zaidner, Yossi, Amow Frumkin, Nqomi Porat, Alexander Tsatskin, Reuven Jagher, and Marcel Otte, eds. Pp. 289–307. Etudes et recherches arche´o- Yeshurun, and Lior Weissbrod. Forthcoming. A series of Mousterian oc- logiques de l’Universite´deLie`ge 126. Lie`ge: University of Lie`ge Press. cupation in a new type of site: the Nesher Ramla karst depression, Israel. Wurz, Sarah. 2013. Technological trends in the Middle Stone Age of South Journal of Human Evolution. Africa between MIS 7 and MIS 3. Current Anthropology 54(suppl. 8):S305– Zilha˜o, Joa˜o, Diego E. Angelucci, Ernestina Badal-Garcı´a, Francesco d’Errico, S319. Flore´al Daniel, Laure Dayet, Katerina Douka, et al. 2010. Symbolic use of Wynn, Thomas, and Frederick L. Coolidge. 2012. How to think like a Nean- marine shells and mineral pigments by Iberian Neandertals. Proceedings of dertal. Oxford: Oxford University Press. the National Academy of Sciences of the USA 107:1338–1342. S358 Current Anthropology Volume 54, Supplement 8, December 2013

Paleolithic Cultures in China Uniqueness and Divergence

by Xing Gao

This paper presents an overview of the Chinese Paleolithic industries between 300 ka and 40 ka, a time span now termed the “later Early Paleolithic” (LEP) in the Chinese chronological scheme. It describes the unique features of LEP remains in China compared with contemporaneous materials in Africa and western Eurasia as well as the internal diversity and complexity of these Chinese Paleolithic assemblages. Basic features of LEP remains in China include the persistent and conservative pebble-tool and simple flake-tool traditions, the use of poor-quality local raw materials, tool fabrication on pebbles and direct use of unretouched flakes, opportunistic flaking, simple and casual modification, and the lack of obvious temporal trends. The diversity and complexity of Chinese Paleolithic cultures as they are expressed in terms of the major difference between southern China’s pebble-tool tradition and northern China’s simple flake-tool tradition are also assessed. Based on such generalizations and analyses, a com- prehensive behavioral model is proposed to explain the unique features of LEP cultures in China and the alternative pathway of human evolution and adaptation in China during that period of time.

Introduction considered to be of marginal importance. Indeed, a global perspective on human origins cannot be properly understood without a detailed consideration of the largest continent Recent research and discussions concerning Pleistocene hu- (Dennell 2009). While studies on African Middle Stone Age man technological development and adaptive strategies have and western Eurasian Middle Paleolithic industries may pro- largely concentrated on archaeological materials during the duce insights about the emergence of modern human be- Middle Late Stone Age and the Middle Upper Paleolithic havior, technology, and the relationship/possible interaction transitions in Africa and western Eurasia, where scenarios of between the intruding early modern human groups and the early modern human origins, dispersal, and their replacement Neanderthals, research on contemporary East Asian Paleo- of the Neanderthals are believed to have taken place. In this lithic remains may provide us a comparative data set of cul- wave of heated discussions and debates, China and East Asia keep almost silent. While the Out of Africa theory enjoys tural and behavioral variability for ancient human groups overwhelming support, the Continuity with Hybridization living in different environmental and ecological zones, and theory looks odd and outmoded, and the cultural remains of this may encourage us to look at the third lineage of human Homo erectus and archaic Homo sapiens seem irrelevant and evolution in late Middle Pleistocene. The unique feature and negligible, for they might be outside the lineage leading di- “conservative” progression of eastern Asian Paleolithic in- rectly to living humans. Once again, as with the influential dustries in general and their obvious distinction from the West Movius Line hypothesis that prevailed in the mid-twentieth seems to support the scenario of continuous evolution and century (Movius 1944, 1948), Paleolithic cultures in China development of local populations, which may present a major are found deep in a “backwater.” challenge to the Out of Africa or Total Replacement hypoth- Human evolution toward complexity and modernity might esis (Gao et al. 2010). have taken different pathways in different regions. In this In order to provide relevant information for this workshop, regard, Asia has received far less attention than Africa and titled “Alternative Pathways to Complexity: Evolutionary Tra- Europe in the search for human origins, but it is no longer jectories in the Middle Paleolithic and Middle Stone Age,” descriptions of the Chinese Paleolithic remains and discus- sions of related questions of this paper will focus mainly on Xing Gao is a Research Member of the Laboratory of Vertebrate the time period between 300 ka and 40 ka, a cultural stage Evolution and Human Origins at the Institute of Vertebrate Paleontology and Paleoanthropology of the Chinese Academy of previously described as the later part of the Lower Paleolithic Sciences (P.O. Box 643, Beijing 100044, China [[email protected]]). and the whole Middle Paleolithic, now reclassified as the later This paper was submitted 3 VII 13, accepted 13 VIII 13, and Early Paleolithic (LEP) by me and my colleagues (Gao 2000; electronically published 14 XI 13. Gao and Norton 2002).

᭧ 2013 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved. 0011-3204/2013/54S8-0014$10.00. DOI: 10.1086/673502 Gao Paleolithic Cultures in China S359

Research Tradition and Conception art, and/or symbolism, indicators of modern human behavior concerning the Chinese Early Paleolithic (ca. 35–30 ka; Gao and Norton 2002).

The practice of Paleolithic research in China still to a certain Movius’s Partition of Two Paleolithic Traditions and extent differs from that of the West, including terminology, Its Influence the classification-typology system, and the way data are pre- Today, most archaeologists agree that the Early Paleolithic sented and interpreted. Therefore, a brief introduction to the assemblages of China and East Asia differ in a number of research tradition and its conception may help Western schol- fundamental ways from Lower and Middle Paleolithic assem- ars understand the content of this paper and research accom- blages of Africa and western Eurasia. For decades, Movius’s plishments in this field in China. partition of two cultural traditions and his hypothetical in- terpretations of it have dominated discussions. Movius proposed two technological traditions in the Lower A Shift from the Middle Paleolithic to the LEP in Chinese Paleolithic: one is the Acheulean handaxe tradition of Africa Paleolithic Research and western Eurasia, characterized by handaxes and other large bifaces. The other is the chopper-chopping tool tradition of East Asia, represented by simple core tools made of pebbles Paleolithic archaeology in China was an adopted enterprise (Movius 1948). His explanation for this difference was that from the West in the 1920s and 1930s, and the three-stage (1) East Asia is a marginal region of human biological and Paleolithic cultural model was carried along with the Western cultural evolution that somehow broke away from the main- scientists who came to China to initiate this field and train stream of human development and maintained the technology local scholars. Accordingly, the Chinese method copied the of the earliest phase of human culture in an isolated context, Western model that was based on artifactual material indig- and (2) the quality of raw material in East Asia was so poor enous to western Eurasia, and the use of similar develop- that it would not permit the ancient population there to make mental stages implies that the cultural evolutionary trajectory better implements. in China was similar to Africa and western Eurasia. However, Even though Movius focused mainly on the Lower Pa- this practice ran into problems when it became obvious that leolithic or the earlier period of the Chinese Early Paleolithic in fact few similarities appear to exist between the Chinese in the new scheme, his conceptualization has profound in- materials and those of the West (Gao and Olsen 1997; Ikawa- fluence on the study of other periods, and Paleolithic research Smith 1978; Movius 1944) and that the Western “index” has somehow been placed under its shadow since then. For stone-tool types of different cultural stages are quite scarce half a century, Chinese scholars have voiced some of the in China and East Asia. Consequently, the derived Paleolithic strongest opposition to the Movius Line. They criticize Mov- cultural development periodization in China began to take a ius’s conception of “chopper-chopping tool” in the East, ar- different approach. Two criteria have been utilized for defin- guing that such artifacts are not typical and dominant in the ing a distinct Middle Paleolithic in China: (1) age of site (i.e., East Asian Paleolithic complexes, that the Chinese and East all archaeological materials dating from the late Middle–Early Asian Paleolithic assemblages are not simple and homoge- Upper Pleistocene—ca. 200–40 ka—were considered Middle neous, and that a certain degree of cultural variability and Paleolithic); and (2) association with archaic Homo sapiens innovation are evident in the archaeological record. Chinese remains. researchers are also eager to demonstrate that some Western Gradually, as the weakness of defining cultural stages based cultural elements, such as Acheulean tool kits and the Lev- on chronometric information and association with certain alloisian technological products, are also presented in the East kinds of human fossils became obvious, researchers began to and that therefore there are no fundamental differences be- agree that such a practice must be based exclusively on the tween the East and West and that East Asia was never a archaeological record (Gao 1999). An analysis of four stone- “cultural backwater” (Huang 1989a, 1989b, 1993; Huang, tool criteria (raw material procurement, core reduction, re- Hou, and Gao 2009). However, not everybody agrees with touch, and typology) to determine whether a distinct Middle such a counterview. Some Western scholars still believe that Paleolithic existed in the Chinese record indicates that very Movius’s “basic characterization of the major characteristics little or very gradual change occurred in lithic technology and of early stage technologies in eastern Asia still holds up” typology between the Lower and Middle Paleolithic. Accord- (Schick 1994:579), and some Chinese researchers also made ingly, there is little reason to retain the three-stage model of similar statements that Movius’s basic observation and con- cultural sequence. Instead, a two-stage progression is pro- clusion were still applicable to the Chinese Paleolithic ma- posed consisting of the Early and the Late Paleolithic. The terials before the Late Paleolithic (Lin 1994, 1996). In other transition between these two cultural periods occurred with words, attempts to support or invalidate the Movius Line the development of more refined lithic techniques (e.g., blade hypothesis have become central to much Paleolithic research and microblade technology) and the presence of ornaments, in China. S360 Current Anthropology Volume 54, Supplement 8, December 2013

The Controversial Hypothesis of Two Parallel Paleolithic 22) and “the number of well-documented Lower Paleolithic Traditions in North China sites remains very small, and the credibility of some of the industries must be questioned because of limited investigation An example of the Chinese archaeologists’ attempts to de- or the lack of the firm chronometric data. Because of these nounce the Movius theory and to demonstrate the diversity problems, it is unreasonable to expect very sophisticated treat- of Paleolithic cultures in the region is the proposition of two ment of specific areal, temporal, and cultural topics. Students parallel Paleolithic traditions in North China. This notion will find much of the literature very limited from theoretical was first put forth in 1972 by Jia and colleagues (Jia, Gai, and and methodological perspectives and rather vague when com- You 1972) and was further developed later (Jia and Huang pared with that of better-studied areas” (Yi and Clark 1983: 1985). Basically, it states that there are two parallel lithic 181). traditions in North China. One is the Kehe-Dingcun Series, Today, Chinese Paleolithic research and Chinese archae- characterized by large chopper-chopping tools and triangular ology as a whole still maintain an ambiguous relationship points; the other is the Zhoukoudian Locality 1-Shiyu Series, with Western practice and theory. Several factors could be characterized by small flake tools such as scrapers and burins. responsible for this. (1) Language: most of the site reports The two “traditions” were postulated as extending in parallel and research papers by the Chinese researchers are published from the Early Paleolithic all the way into the Late Paleolithic in Chinese journals, and the language barrier prevents Chinese and even the Neolithic and developing into two different and foreign scholars from sharing information and exchang- agricultural patterns. This hypothesis had been influential and ing ideas freely. (2) Different research priorities: a large part made significant impact on Paleolithic research in China dur- of archaeological activities in China are undertaken as salvage ing the last three decades of the twentieth century, and some projects, and rescue of the artifacts or cultural relics and basic researchers are still working in this domain up to the present. classification and description rather than detailed analysis and Such a notion is quite troublesome. The sites of the two theoretical explanations are the principal tasks of the field- “traditions” are distributed basically in the same region, and worker. (3) The profound impact of traditional epigraphy on it is difficult to imagine that two distinct cultural traditions modern archaeological practice helps maintain a long-lasting can exist side by side in the same area for about a million and persistent classification and description tendency, and the years. Studies have revealed that most of the sites of the large- political situation, especially the adoption of Marxism and tool tradition, including the key site Dingcun, are in fact Maoism as notional ideology by the New China, strengthened dominated by small flake tools (Zhang 1993). The most se- the tradition. (4) Believing in the philosophy that scientific rious problem with the “large-tool tradition” is taphonomic: reasoning should be very cautious and that one has to ac- almost all the localities assigned to the “large-tool tradition” cumulate enough data before reaching meaningful conclu- were fluvial sites exhibiting traces of disturbance and sec- sions, most Chinese scholars are reluctant or lack confidence ondary deposition, and many of these large stone tools were to touch theoretical issues, and if they have to do so, they selectively collected from the ground surface. Therefore, these would like to take an inductive approach rather than being collections could not represent a complete “assemblage” or deductive. (5) A strict and somewhat exclusive government “tool kit” and certainly not a “cultural tradition.” Another policy regarding foreigners’ involvement in archaeological re- drawback of this hypothesis is that it may well overlook or search in China, especially fieldwork, makes it difficult for minimize variations within the specific lithic assemblages as- foreign scholars to get firsthand information and experience cribed to each of these so-called traditions and, at the same and to establish long-term research programs there. We time, potentially underestimate the similarities among in- should also draw attention to the fact that compared with dustries of different “traditions.” In short, the diversity and the large size of the country and the rich archaeological ma- complexity of the lithic industries in China cannot be sum- terials, the Paleolithic research community in China is very marized simply by two unilinear traditions based solely on small, and the number of well-trained professional researchers typological and morphological analyses. familiar with global methodological and theoretical ap- proaches is inadequate. In addition, many Chinese researchers Communication Obstacles and the Distinct Paleolithic are accustomed to doing research in their own territory and Research Tradition in China seldom go beyond its border, even within China. I would argue that the difficulties and differences in lan- China has rich collections of Paleolithic remains. Many for- guage and research practices are superficial. The more fun- eign researchers are eager to access the collections and to damental reason that the Chinese record plays such a limited establish fruitful contact with colleagues there. However, they role in discussion of evolutionary trends within the Middle often feel frustrated; typical comments are “prehistorians out- and Late Pleistocene is that the archaeological evidence is still side of China have found it difficult to obtain good infor- difficult to reconcile with what is known from Europe and mation about the results of this surge in archaeological in- Africa. Chinese and eastern Asian Paleolithic materials really quiry, much less to synthesize a comprehensive understanding are different from those of the West in many ways; their of Paleolithic trends in eastern Asia” (Schick and Dong 1993: unique types and morphological and developmental features Gao Paleolithic Cultures in China S361 make Western typology and terminology difficult to apply, Homo sapiens, numerous mammalian fossils, some stone ar- and many Western scholars do not fully understand them. tifacts, and ash and burned items were unearthed from several They expect to find similar cultural remains and familiar re- cultural horizons. These cultural remains were generally dated search results, but they are often disappointed. We have failed to 140–250 ka, and even older ages were suggested (Wei 2009; to work out a typological-descriptive system and research Zhang et al. 2007). A total of 64 lithic artifacts have been norms that can be effectively applied to archaeological ma- reported and analyzed, including simple cores, flakes, and terials from both the West and the East. retouched pieces. Direct hammer percussion was used as the principal method of core reduction; core preparation was The Chinese LEP Archaeological Record: seldom applied. The tools were fabricated coarsely, and the Uniqueness versus Divergence artifacts vary greatly in size and morphology. Side scrapers are the dominant tools, and most of them are small; chopper- Literally hundreds of archaeological sites estimated to belong chopping tools take the second position, and some of them to the 300–40 ka time span have been reported in China (fig. are very large. 1). The exact figure is hard to calculate, for some sites were assigned a wide chronological range, and it is difficult to say Jinniushan. Jinniushan is a cave site complex located near whether they belong to this stage; and in some regions, many Yingkou City, Liaoning Province, Northeast China. It was localities were identified and numbered, and it is difficult to discovered in 1974 and excavated in the 1970s and 1980s (Lu¨ determine whether they belong to one site or discrete sites. 2004). The excavations resulted in the discovery of a partial I will describe a few key sites to provide some basic infor- skeleton of archaic Homo sapiens, numerous mammalian fos- mation. sils, and stone artifacts. A few hearths covered by rocks be- lieved to be evidence of fire preservation, along with burnt Key Sites materials, were also unearthed. Several chronometric tests Miaohoushan. Miaohoushan is a cave site situated near have been applied to the human fossil and artifact-bearing Benxi City, Liaoning Province, in Northeast China (Zhang horizons, and ages of 230–300 Ka, 228 Ka, and 187 Ka were 1989). It was discovered in 1978 and excavated in 1978, 1979, reported (Chen, Yang, and Wu 1994). Only a part of the and 1980. A few human fossil fragments, probably of archaic unearthed materials has been analyzed and published. Ac-

Figure 1. Key localities among major LEP sites in China. 1, Jinniushan; 2, Miaohoushan; 3, Xujiayao; 4, Zhoukoudian Locality 15; 5, Dingcun; 6, Dali; 7, Longyadong; 8, Lingjing; 9, Jigongshan; 10, Jingshuiwan; 11, Guanyindong; 12, Dadong. S362 Current Anthropology Volume 54, Supplement 8, December 2013 cording to the available information, the lithic assemblage Dingcun. The Dingcun site complex is located in Shanxi includes simple cores, flakes, scrapers, points, and burin. Core Province, North China, and was originally discovered in 1953. reduction was mainly carried out through direct hammer per- The site is made up of a dozen separate open-air localities. cussion without core preparation, and the bipolar technique Several excavations have been carried out at the site; a parietal was also used. The tool kit is dominated by side scrapers, and skull and some isolated teeth of archaic Homo sapiens and a retouch on them is simple and casual. Most artifacts are small large quantity of vertebrate fossils and lithic artifacts were and vary in size and morphology. Some chipped-bone tools collected. U-series dates and lithostratigraphic and bio- were also believed to be produced and used. stratigraphic reconstructions indicated an age range of 260– 107 ka for the cultural remains (Norton, Gao, and Feng 2009). Zhoukoudian Locality 15. Zhoukoudian Locality 15 is part For a long time, the rich lithic collection from the site has of the limestone cave complex of the Zhoukoudian site, lo- been the center of discussion and debate in Paleolithic re- cated 50 km southwest of Beijing. The locality was discovered search in China. Jia nominated Dingcun as the representative in 1932 and excavated from 1935 to 1937. A large quantity site of the “large chopper-chopping tool and triangular point of vertebrate fossils and lithic artifacts was unearthed. Faunal tradition” of North China, for most of the artifacts were assemblage indicates a late Middle to early Upper Pleistocene considered to be large ones, and some of them exhibited age, and limited uranium series dates estimated an age range distinctive typological and technological features. Huang put between 140 and 110 ka for the cultural horizon (Gao 2000). forward the notion that many large pointed tools in the as- The rich lithic assemblage is composed of more than 10,000 semblage were handaxes; therefore, Dingcun was the center stone artifacts, including hammerstones, cores, flakes, re- of a handaxe zone along the Fen and Wei rivers in North touched pieces, and chunks, with the latter predominating. China (Huang 1989a). However, close examination of the Hammer percussion was the principal flaking strategy, but collections reveals that the assemblages are dominated by bipolar flaking was also employed frequently, which made the small flake tools, mainly side scrapers and points, not large assemblage unique in the Chinese LEP and a clear successor pebble tools, and there are no real handaxes from the site of the earlier “Peking Man” industry known from Zhoukou- (Gao 2011). Still, the Dingcun industry shows some special dian Locality 1. In addition to the simple cores, regular discoid characteristics within the flake-tool techno-complex in China cores and heavily reduced polyhedral cores were also present, in that core reduction, even though still through simple hard- testifying to a sophisticated fashion of core reduction. The hammer percussion and possible anvil technique, seems to retouched tools are mostly side scrapers; other tool types in- be more sophisticated, and many large and regular flakes and clude chopper-chopping tools, backed knives, points, awls, tools were produced. Some large well-made triangular picks notches, and burins (fig. 2B). The tools are mostly fabricated made on flakes with unique morphology as well as chopper- on flakes and are small in size, but the few pieces of backed chopping tools and spheroids were also present (fig. 2D). A knives or cleavers are large and distinctive. Most of the ar- major reason for the uniqueness of the Dingcun industry tifacts were fabricated from locally available quartz, a raw might be the exploitation of high-quality dark hornfels avail- material source characterized by high abundance and low able in nearby river beds. workability. An analysis of raw material utilization reveals that some simple but practical and efficient strategies were adopted Xujiayao. Xujiayao is a fluvial-lacustrine open-air site lo- to make use of these raw materials: different materials were cated in the western margin of the Nihewan Basin in Hebei procured and consumed differently (table 1); the site was Province, North China (Jia, Wei, and Li 1979). It was exca- provisioned with abundant potential tool-making materials; vated three times in the late 1970s and more in recent years. numerous flakes were detached, but only a portion was se- Some fragmental archaic Homo sapiens fossils and an array lected for modification and utilization (Gao 2003). of vertebrate fossils and Paleolithic materials were recovered. U-series, paleomagnetism, and optically stimulated lumines- Dali. Dali is an open-air site discovered in 1978 in Shanxi cence (OSL) dating have been applied to the site, and dates Province, North China, and is best known for the presence of 125–104 ka, 117 ka, and 69 ka have been obtained, re- of an archaic Homo sapiens skull (Wu 2009), but some faunal spectively (Chen et al. 1982; Liu, Su, and Jin 1992; Nagatomo remains and 564 pieces of stone artifacts were also collected et al. 2009). A recent study on 1,765 lithic artifacts unearthed from sandy deposits. U-series and electron spin produced an in 1977 indicates that the assemblage includes cores, flakes, age range between 380 and 140 ka (Chen, Yuan, and Gao retouched pieces, chunks, and debris (Ma, Pei, and Gao 2011). 1984; Yin et al. 2001). Core reduction at the site was found Raw materials were mainly quartzite and quartz pebbles se- to be conducted through hammer percussion and the bipolar lected from nearby river beds, simple hammer percussion was method. Most of lithic artifacts are very small side scrapers, used for core reduction, and a certain number of discoid and points, burins, and drills produced from quartzite, flint, and polyhedral cores were left behind. About half of the retouched quartz materials. Retouch on these pieces is very simple and pieces are small side scrapers made on flake blanks, and sphe- casual, and some of them were heavily worn and difficult to roids constitute more than 27% of the assemblage, which is study typologically and technologically. very distinctive. Other tool types include point, notch, den- Figure 2. Line drawings of stone artifacts from some key LEP sites in North China. A, Longyadong; B, Zhoukoudian Locality 15; C, Xujiayao; D, Dingcun. S364 Current Anthropology Volume 54, Supplement 8, December 2013

Table 1. Raw material frequencies for artifacts by class from Zhoukoudian Locality 15

Quartz Igneous Crystal Flint Sandstone Quartzite Class N % N % N % N % N % N Core 126 1.8 2 1 1 Flake 393 5.7 113 1.6 15 .2 9 .1 Bipolar 86 1.3 1 Hammer 5 .1 2 Chunk 4,730 68.9 32 .5 66 1.0 1 Tool 1,198 17.4 54 .8 14 .2 12 .2 4 .1 1 ticulate, chopper-chopping tools, burin, and drill tools (fig. finds (Wang 2005). Paleomagnetism and OSL dating suggest 2C). Some bone tools were also recognized in previous studies. that ancient humans stayed in the area off and on in the time Because of the high density of equid and rhinoceros bones span of 800–140 ka (Lu et al. 2007). These sites have attracted and artifacts, particularly stone spheroids and bone tools, Xu- a great deal of attention in China because of the discovery of jiayao was interpreted as a horse kill site. a certain number of Acheulean-like handaxes, cleavers, and picks along with other large pebble tools and small flake tools Lingjing. Lingjing is an open-air site located near Xuchang (fig. 2A). Raw materials used for stone-tool manufacture are City, Henan Province, central China. The site was discovered overwhelmingly quartzite pebbles. Such a tool kit is quite in the mid-1960s and excavated recently. A broken skullcap unique in central-southern China’s pebble-tool zone in which of possible archaic Homo sapiens was unearthed in association simple and large chopper-chopping tools dominated the in- with a large quantity of mammalian fossil fragments and dustries throughout the Paleolithic. Recent excavation and thousands of stone tools. OSL dates and biostratigraphic data chronometric dating suggest that some of the Acheulean-style estimate the age of 110–80 Ka for the Paleolithic horizon. artifacts might come from the Upper Pleistocene horizon, The lithic assemblage is dominated by small side scrapers, which may present challenges to the study of Paleolithic hu- points, and drills made of vein quartz flakes, but some heavy- man adaptation, migration, interaction, or convergent cul- duty chopper-chopping tools and picks made of quartzite tural development. blank are also present (Li 2007). Core reduction and tool manufacture are found to be simple and casual. Some mod- Jigongshan. The Jigongshan site is situated in the Jingzhou ified bone tools, mostly pointed ones, have been identified district of Hubei Province along the Yangtze River. It was and analyzed (fig. 3), and use-wear analysis results suggest discovered in 1984 and excavated in 1992. Numerous lithic that some bone tools were used for drilling, penetrating, and materials were unearthed from two cultural horizons at the scraping animal substances and that some might have been site. The lower horizon was estimated to be of the early Upper hafted during use (Li and Shen 2010). Pleistocene and the upper horizon of the late Upper Pleis- A study of mortality profiles of the large herbivores from tocene. Lithic assemblages from the two horizons were dom- the site suggests that the accumulation of rich mammalian inated by simple cores, flakes, and chunks. Most of the tools bone fragments is the result of human hunting and butch- from the lower horizon were made of pebbles, and heavy- ering. Aurochs (Bos primigenius) and horse (Equus caballus) duty tools such as picks and chopper-chopping tools make are the major prey species, and the age structures of these up the large majority of the assemblage (fig. 4D). Tools from animals can be best described as the “prime-dominated pat- the upper horizon were mostly made on flakes, and most of tern.” This study confirms the well-established notions at them are small irregular scrapers. According to the site report many Middle and Upper Paleolithic sites across Eurasia and (Liu and Wang 2001), the “living floor” with a circular struc- Africa that Middle Stone Age/Middle Paleolithic foragers were ture composed of pebbles and artifacts was identified from fully effective in hunting large prey species, particularly au- the lower cultural horizon. rochs and horse, which might indicate that the hunting be- Sites in Sanxia (Three Gorges Region). More than 20 Pa- haviors and subsistence strategies were not significantly dif- leolithic sites have been discovered and excavated in the Three ferent between Middle Paleolithic and Upper Paleolithic Gorges Region (Sanxia in Chinese) in Chongqing Municipal humans in East Asia and hence suggest the early emergence City, central China, in the past two decades, and some sites of modern human behaviors in this area (Zhang et al. 2009). have been dated to the time span of 140–70 ka, such as Sites in the Luonan Basin. Since 1995, more than 300 sites Jingshuiwan (Gao and Pei 2010; Pei et al. 2006). Lithic ar- have been found in the Luonan Basin of southern Shanxi tifacts from these sites are typical of the pebble-tool industries Province, central China. Among them, only Longyadong is a that prevailed in southern and central China during the entire cave site, and the others are open-air sites. Tens of thousands Pleistocene. Assemblages are dominated overwhelmingly by of stone artifacts have been collected from different river ter- large chopper-chopping tools and picks made of pebbles. Only races along the Luonan River, and most of them are surface a small portion of the tools were fabricated on flake blanks. Gao Paleolithic Cultures in China S365

Figure 3. Possible bone tools from Lingjing.

Raw materials exploited are mostly quartzite and quartzite points, drills, burins, and chopper-chopping tools; most of sandstone. Core reduction was carried out using direct ham- them are small flake tools (fig. 4B). The retouched pieces mer percussion and a kind of special “throw and collision exhibit a simple and irregular mode of modification; some method.” Only simple cores and flakes were produced. Re- of them possess more than one cutting edge, and the edges touch of tools was simple and casual (fig. 4A). are usually thick and steep. The Guanyindong site was initially estimated to be of Middle Pleistocene age, and the cultural Dadong (Grand Cave). The Dadong or Grand Cave site is remains were believed to be comparable with those of the located in Guizhou Province, South China, and was discov- Peking Man site at Zhoukoudian. However, later chrono- ered and excavated in 1990s. More than 2,000 artifacts have metric dating placed the lower horizon at 50–140 ka and 180– been unearthed (Huang, Hou, and Si 1997), and the age of 240 ka and the upper horizon to be younger than 40 ka (Shen human occupation of the site has been estimated to be 260– and Jin 1992). 142 ka (Wang et al. 2003). Raw materials exploited are mostly small flint nodules. Only direct hard-hammer percussion was employed for core reduction, and a few Levallois-like flakes Principal Features of the Chinese LEP Paleolithic Assemblages were reported. The assemblage is dominated by small flake It is obvious that Early Paleolithic industries in China have tools, including side scrapers, drills, notches, denticulates, and their unique features compared with contemporary cultural end scrapers (fig. 4C). A few pieces of rhinoceros teeth were remains in Africa and western Eurasia. The most distinctive identified as modified into scraping tools. cultural characteristics of the Chinese LEP Paleolithic assem- Guanyindong. The Guanyindong cave site is also located in blages can be summarized as follows. Guizhou Province, South China, and was discovered and ex- 1. Slow or conservative development process in that Mode cavated in the 1960s (Li and Wen 1986). More than 3,000 I technology and assemblages prevailed for all the LEP. Most stone artifacts and numerous animal fossils were unearthed of the assemblages consist of simple cores, irregular flakes, from two depositional units. Stone-tool raw materials ex- side scrapers, chopper-chopping tools, points, picks, and so ploited are locally available flints. Direct hard-hammer per- forth. It is true that Acheulean-like tool kits, including hand- cussion was believed to be used for core reduction and re- axes, cleavers, and picks were reported from some localities touch, and a few Levallois-like flakes were identified. A rich of the Luonan Basin and the Dingcun site. Some Acheulean- variety of tool types were recognized from the assemblage, like assemblages from the Luonan Basin were estimated to be including side scrapers, end scrapers, notches, denticulates, of the late Middle Pleistocene or even the upper Pleistocene. Figure 4. Line drawings of stone artifacts from some key LEP sites in South China. A, Jingshuiwan; B, Guanyindong; C, Dadong; D, Jigongshan. Gao Paleolithic Cultures in China S367

However, most of these artifacts are surface finds, and the area, and it is more appropriate to think about regional find- age of such assemblages needs to be further analyzed. Fur- ings according to paleoecological conditions rather than treat- thermore, handaxes found in this region are mostly pointed ing the vast territory as a whole. and thick, and no trace of soft-hammer retouch and thinning The complexity of Early Paleolithic industries in China and (typical of contemporaneous industries in Europe, the Near their regional variability have been realized in the past three East, and Africa) can be observed from the samples. No real decades. Zhang Senshui’s observation of “two main Chinese handaxes or cleavers were collected from the Dingcun site, Paleolithic industries with numerous local cultural variants” and no technological mark of soft-hammer flaking and thin- is a good example (Zhang 1999:198). He proposed that two ning can be identified from the artifacts. Before 40 ka, no principal industries could be recognized from the Chinese Mousterian-style assemblages or blade technology can be Paleolithic remains, that is, the small flake tool industry in identified in the Chinese Paleolithic industries. North China and the large pebble-tool industry in South 2. Only local raw materials were exploited for tool making, China. He called them the northern main industry (NMI) and they are mostly poor-quality quartz, quartzite, sandstone, and the southern main industry (SMI), respectively. Within and silicified limestone. At some sites chert and flint were these two cultural zones, divided by the Huai River in the used, but they are usually of poor quality and in small nodules. east and the Qinling Mountains in the west, he further rec- No evidence of quarrying and long-distance transportation ognized numerous local cultural variants. Major features of of high-quality raw material has been detected so far. NMI were summarized as including the domination of small 3. A variety of flaking methods used for core reduction is flake tools, mainly various kinds of side scrapers, points, awls, evident in these industries, including direct hard-hammer burins; large pebble tools, such as chopper-chopping tools, percussion, bipolar, and block-on-block techniques. Cores picks, and spheroids, were of secondary importance; direct were rarely prepared, and no application of real Levalloisian hammer percussion, the bipolar technique, and the block-on- technique was recognized. Simple cores with a few detach- block method were used for flaking; and the industries ex- flake scars and polyhedral ones are the most numerous, and hibited slow and conservative development during the Early discoidal cores appear in some assemblages. Flakes are usually Paleolithic but showed acceleration of technological devel- small, irregular, and vary in size and morphology. Evidence opment and innovation as well as the emergence of new tech- for “predetermination” of flake shapes by core preparation is niques and tool types (such as blade and microblade tools lacking. No trace of soft-hammer flaking has been recognized and technology) in the Late Paleolithic. Meanwhile, charac- for this period of time. teristics of SMI include the domination of large pebble tools, 4. Tools are mostly simple, irregular, and casually modified, mainly chopper-chopping tools, picks, and handaxes; rarely and some are difficult to classify into discrete types. In the present and poorly fabricated scrapers and points; hammer- north—that is, to the north of the Qinling Mountains and percussion and block-on-block methods used for core re- the Huai River, a natural boundary that normally divides duction; fewer clear tool types and coarser retouch compared China into two ecological zones—industries were mostly with NMI; and a much stronger conservative developmental dominated by small flake tools such as side scrapers, points, trend even all the way to the early Neolithic. On the basis of and drills supplemented by chopper-chopping tools, picks, the partition of the north versus the south cultural zones and and spheroids, while in the south, industries were dominated the formation of many regional cultural variants, Zhang by large pebble tools, especially chopper-chopping tools and pointed out possible factors, such as environmental differ- picks. Generally speaking, the degree of tool standardization ences and human migrations and interactions, but was short is pretty low, and flakes were frequently utilized without fur- of detailed analysis. ther modification. There are certainly exceptions. Some large Simple stone-tool technology and assemblage and slow de- and well-made digging-cutting tools were collected from a velopment of the Chinese LEP should not lead to the con- few localities in the Dingcun site complex and the Luonan clusion that they are stagnant. In fact, changes and devel- Basin, such as triangular picks, handaxes, and cleavers, and opment in these industries are still evident through time. some small flake tools, such as side scrapers and points, un- Zhang Senshui (1989) summarizes the major developmental earthed from sites such as Zhoukoudian Location 15 and trends of the Chinese Lower Paleolithic as follows. (1) More Guanyindong, are found to be fabricated delicately and skill- and more high-quality raw materials were exploited, especially fully. It might mean that when raw material was suitable and silicified limestone and flint. (2) Direct hard-hammer per- necessity arose, human groups living at these sites were ca- cussion technique underwent a discernible process of mat- pable of producing regular, efficient and curated tools. uration while block-on-block techniques diminished in im- Any attempt to summarize the basic features of the Chinese portance. (3) Morphologically regular flakes and tools made LEP as a whole is inevitably oversimplified. While the fact on them increased in number. (4) More flakes were used as that Paleolithic industries in China and East Asia are different tool blanks, and even in the pebble-tool zone of southern from those of the West has been recognized, regional diversity China, flake tools increased through time. (5) more tool types and internal complexity are also evident in these industries. were added to the assemblages, and the discrepancy between China covers a vast and geographically/ecologically diverse tool classes became clearer. (6) Chopper-chopping tools be- S368 Current Anthropology Volume 54, Supplement 8, December 2013 came less common, and points and drills in accordance in- isolation, restrictions of raw materials, and ecological/envi- creased and became more regularized. (7) Through time, ronmental conditions, and some theories are short of sup- more delicately retouched tools increased in number. (8) Re- porting evidence. The formation of a lithic industry or of a touch changed from multidirectional to mainly unidirectional certain cultural tradition should be a complex process that on the dorsal surface. Such technological and morphological might involve many influencing factors. Maybe it is time to change or development is not as obvious and dramatic as work out a model that is more comprehensive and takes an what happened to the Early and Middle Stone Age or Lower integrative approach to consider both environmental effects and Middle Paleolithic in the West. However, they provide and human behavior and adaptive strategies. I call it a “com- us the opportunity to think about and reconstruct different prehensive behavioral model.” The model may offer the fol- or alternative trajectories of human evolution during the re- lowing observations and explanations on the stable devel- mote past. opment and unique features of Early Paleolithic industries in China. Discussion Stable Environments and Continuity of Human Evolution For a long time, discussions of technological similarities and differences between Paleolithic tool traditions have been fre- During the LEP, China was under the control of monsoon quently based on assumptions regarding what biological af- climates. Studies of Loess depositional sequence and faunal finities may indicate about cultural ties and vice versa. Such assemblages suggest that even if climatic fluctuations occurred assumptions are not always borne out in the real world when periodically, environmental conditions were relatively stable we realize that biologically closely related populations some- in the region, and most of the area was suitable for human times exhibit fundamental differences in their social structure habitation (Liu 2009). Rich and continuous archaeological and even language, and culture and technology can be trans- records indicate that human evolution in the region was stable mitted quickly between geographically distant groups (Schick and uninterrupted and without large-scale population re- 1994). Therefore, archaeologists must pursue alternative hy- placement. Strong and stable cultural traditions were formed potheses to explain cultural variability. during this process, and occasional outside intruders were Other than the Movius Line theory, many hypotheses have assimilated into the mainstream populations and lost their been proposed to interpret the unique features of Paleolithic cultural identities. industries in China and East Asia. A few researchers have suggested that the main tools used by Paleolithic humans for Low-Intensity Resource Exploitation and High Mobility adaptation in the region were those made from bamboo and Most of the LEP sites in China are seasonal, short-time oc- wood materials, and the simple stone tools were actually used cupied ones, and artifacts collected from them are mostly to make such vegetal tools. Therefore, stone artifacts are not simple and share basic features of technology, typology, and the right indicator of human technological development and morphology, which may indicate that human groups living adaptive strategy in the Pleistocene (Pope 1989). Some pro- in the region had a simple and “easy” hunting-gathering life- posed that the lack of high-quality raw material in East Asia style. They kept the exploitation of natural resources at a was the major obstacle for ancient populations living in the rather low intensity and seldom felt the pressure of innovating region to develop more sophisticated lithic technology and lithic technology to procure difficult resources. In keeping make stone tools as good as those of Africa and western such a lifeway, they moved frequently to other places to find Eurasia (Schick 1994). Some even suggested that when early new food resources, and therefore they left identical artifacts human groups migrated into East Asia, they first encountered and other materials at many sites in certain regions. tropical and subtropical ecological conditions in the southern part of the territory. In such environments, foods were ob- High Flexibility in Tool Technology and Adaptation tained mainly through gathering plant fruits and roots rather than hunting game; large hunting and butchering tools were The lack of high-quality stone raw materials and suitable quar- useless, and simple pebble and flake tools took the dominant rying sites forced Early Paleolithic humans living in the area role. Such a shift of survival conditions and adaptive strategies to make best use of poor-quality and locally available raw brought about fundamental changes to stone-tool technology materials. In dealing with such materials with great variability and the composition of tool kits, and the watershed in Pa- in lithology and morphology, these people learned to be highly leolithic industries between the East and the West began to flexible and use simple but suitable and effective ways to appear (Watanabe 1985). produce tool blanks and make stone tools. For instance, peo- The above hypotheses all offered some explanations on the ple living at Zhoukoudian Locality 15 relied heavily on the distinctive features of Paleolithic remains in China and East bipolar technique to make use of quartz nodules that were Asia and cultural differences between the East and the West abundant in the nearby river bed. People living at the Dingcun in most of the Pleistocene. However, such discussions are site applied the hammer-percussion method skillfully to dark often confined to some isolated factors, such as geographic hornfels, a relatively high-quality local raw material, to detach Gao Paleolithic Cultures in China S369 large and regular flakes and to produce large and sharp cutting Dennell, Robin. 2009. The Paleolithic settlement of Asia. New York: Cambridge University Press. and digging tools. Still, people in the Sanxia region invented Gao, Xing. 1999. A discussion of the “Chinese Middle Paleolithic.” Acta An- the unique “throw and collision method” to exploit highly thropologica Sinica 18(1):1–16. polished and rounded river pebbles. Such flexibility might ———. 2000. Interpretations of typological variability within Paleolithic re- mains from Zhoukoudian Locality 15, China. PhD dissertation, University have enforced ancient human survival capability and helped of Arizona, Tucson. the Paleolithic traditions to be strong, stable, and full of vi- ———. 2003. A study of the lithic assemblage from Zhoukoudian Locality tality. If the Bamboo Tool hypothesis can be verified, it will 15. Acta Anthropologica Sinica 21(suppl.):31–52. ———. 2011. The nature of Paleolithic handaxes from China and its impli- be a good example of the flexibility and intelligence of ancient cations for Lower Paleolithic cultural variation. In Handaxes in the Imjin humans living in the region. Basin: diversity and variability in the East Asian Paleolithic. Seonbok Yi, ed. Pp. 193–217. Seoul: Seoul National University Press. Gao, Xing, and Christopher J. Norton. 2002. A critique of the Chinese “Middle Conclusions Paleolithic.” Antiquity 76:397–412. Gao, Xing, and John W. Olsen. 1997. Similarity and variation within the Lower It is obvious that the LEP industries in China and East Asia Paleolithic: East Asia, western Europe and Africa compared. In Evidence for are different from those of contemporary Paleolithic remains evolution. T. Yongsheng, ed. Pp. 63–76. Beijing: Ocean Press. Gao, Xing, and Pei ShuWen. 2010. Footprints of ancient humans in the Three in Africa and western Eurasia in many ways. However, it is Gorges region. Chengdu: Sichuan Publishing. also clear that the Lower Paleolithic world should not be Gao, Xing, Zhang XiaoLing, Yang DongYa, Shen Chen, and Wu XinZhi. 2010. simply divided into two different cultural/technological Revisiting the origin of modern humans in China and its implications for global human evolution. Science China (Earth Sciences) 40(9):1287–1300. traditions based solely on stone-tool technological and ty- Huang, WeiWen. 1989a. Bifaces in China. Human Evolution 4(1):87–92. pological comparisons. The variation in Paleolithic industries ———. 1989b. The early Paleolithic of China. Quaternary Research 28:237– between the West and the East is undeniable, but it should 242. ———. 1993. Typology of Lower Paleolithic heavy-duty implements from be understood within a broad framework of universal cultural East and Southeast Asia: comments on the Movius’s typological system. diversity. It should be realized that while ancient hominids Acta Anthropologica Sinica 12(4):297–304. in different parts of the world shared some basic lithic tech- Huang, WeiWen, Hou YaMei, and Gao LiHong. 2009. “Western elements” in the Chinese Paleolithic as viewed in a framework of early human cultural nologies and produced and utilized similar stone tools (such evolution. Acta Anthropologica Sinica 28(1):16–25. as core/flake tools), each group was unique in its methods of Huang, WeWen, Hou YaMei, and Si XinQiang. 1997. Stone industry from survival and adaptation because of ecological context and raw Panxian Dadong, a cave-site of southwestern China. Acta Anthropologica Sinica 16(3):171–192. material availability and quality. What is more important is Ikawa-Smith, F., ed. 1978. Early Paleolithic in South and East Asia. The Hague: to look beyond lithic technological and typological variability Mouton. and find the factors and dynamics behind such cultural dif- Jia, LanPo, Gai Pei, and You YuZhu. 1972. Excavation report on the Shiyu site in Shanxi. Acta Archaeologica Sinica (1):39–58. ferences and reconstruct different pathways of human evo- Jia, LanPo, Wei, Qi, and Li, ChaoRong. 1979. Report on the excavation of lution toward complexity and modernity. the Hsuchiayao Man site in 1976. Vertebrata PalAsiatica 17:277–293. [In Chinese.] Jia, LanPo, and WeiWen Huang. 1985. On the recognition of China’s Pa- leolithic cultural traditions. In Paleoanthropology and Paleolithic archaeology in the People’s Republic of China. Wu RuKang and John W. Olsen, eds. Pp. Acknowledgments 259–265. New York: Academic Press. Li, YanXian, and Wen BenHeng. 1986. Guanyindong: a Lower Paleolithic site I would like to thank Steve Kuhn and Erella Hovers for their in Qianxi county, Guizhou Province. Beijing: Cultural Relic Press. invitation to the symposium “Alternative Pathways to Com- Li, ZhanYang. 2007. A preliminary study on the stone artifacts of Lingjing site excavated in 2005. Acta Anthropologica Sinica 26(2):138–154. plexity: Evolutionary Trajectories in the Middle Paleolithic Li, ZhanYang, and Shen Chen. 2010. Use-wear analysis confirms the use of and Middle Stone Age” and their suggestions for and help Palaeolithic bone tools by the Lingjing Xuchang early human. Chinese Sci- with the revision of this paper. I also thank Leslie Aiello and ence Bulletin 55:2282–2289. Lin, ShengLong. 1994. Restudy of nine hand-axe specimens and the appli- Laurie Obbink for their organization of the symposium and cability of Movius’ theory. Acta Anthropologica Sinica 13(3):189–208. their assistance. The research that led to the writing of this ———. 1996. A comparative study on the Paleolithic technological modes paper was supported by the Chinese Academy of Sciences between China and the West. Acta Anthropologica Sinica 15(1):1–20. Liu, C., P. Su, and Z. Jin. 1992. Discovery of Blake Episode in the Xujiayao Strategic Priority Research Program (XDA05130202) and the Paleolithic site, Shanxi, China. Scientia Geologica Sinica 1:87–95. Ministry of Science and Technology of China Groundwork Liu, DeYin, and Wang YouPing. 2001. A preliminary report on the excavation Project (2007FY110200). of Jigongshan site. Acta Anthropologie Sinica 20(2):102–114. Liu, DongSheng. 2009. Loess and arid environment. Hefei, China: Anhui Sci- entific and Technological Press. Lu, HuaYu, Zhang HongYan, Wang SheJiang, Zhao CunFa, and Zhao Jun. References Cited 2007. A preliminary survey on loess deposit in eastern Qinling Mountain Chen, Tiemei, Yang Quan, and Wu En. 1994. Antiquity of Homo sapiens in in central China and its implication for estimating age of the Pleistocene China. Nature 368:55–56. lithic artifacts. Quaternary Sciences 27(4):550–567. Chen, Tiemei, Yuan SiXun, and Gao ShiJun. 1984. The study on uranium- Lu¨Zune. 2004. Paleolithic archaeology in Liaoning Province. In Chinese ar- series dating of fossil bones as an absolute age sequence for the main chaeological research in the 20th century (Paleolithic volume). Lu¨ Zune, ed. Paleolithic sites of North China. Acta Anthropologica Sinica 3(2):259–269. Pp. 194–219. Beijing: Science Press. [In Chinese.] Chen, Tiemei, Yuan SiXun, Gao ShiJun, Wang LiangXun, and Zhao GuiYing. Ma, Ning, Pei ShuWen, and Gao Xing. 2011. A preliminary study on the 1982. U-series dating on the mammalian fossils from the Xujiayao site. stone artifacts excavated from Locality 74093 of the Xijiayao Site in 1977. Acta Anthropologica Sinica 1(1):91–95. Acta Anthropologica Sinica 30(3):275–288. S370 Current Anthropology Volume 54, Supplement 8, December 2013

Movius, H. L. 1944. Early man and Pleistocene stratigraphy in southern and Rink, Xin-quiang Si, and Wei-wen Huang. 2003. Stratigraphic and paleoen- eastern Asia. Papers of the Peabody Museum of American Archaeology and vironmental studies at the Dadong Cave, Panxian. Acta Anthropologica Sin- Ethnology, vol. 19, no. 2. Cambridge, MA: Peabody Museum. ica 22(2):131–138. ———. 1948. The Lower Paleolithic cultures of southern and eastern Asia. Watanabe H. 1985. The chopper-chopping tool complex of eastern Asia: an Transactions of the American Philosophical Society 38(4):329–420. ethnoarchaeological-ecological reexamination. Journal of Anthropological Nagatomo, Tsuneto, Yorinao Shitaoka, Hisae Namioka, Masatoshi Sagawa, Archaeology 4:1–18. and Wei Qi. 2009. OSL dating of the strata at Paleolithic sites in the Nihewan Wei HaiBo. 2009. Continued research on the Miaohoushan site, Liaoning Basin, China. Acta Anthropologica Sinica 28(3):276–284. Province. Acta Anthropologica Sinica 28(2):154–161. Norton, Christopher J., Gao Xing, and Feng XingWu. 2009. The East Asian Wu, XinZhi. 2009. A metrical study of the Dali cranium. Acta Anthropologica Middle Paleolithic reexamined. In Sourcebook of Paleolithic transition.Marta Sinica 28(3):217–236. Camps and Parth Chauhan, eds. Pp. 245–254. Dordrecht: Springer. Yi, S., and G. A. Clark. 1983. Observations on the Lower Paleolithic of north- Pei, ShuWen, Zhang JiaFu, Gao Xing, Zhou LiPing, Feng XingWu, and Chen east Asia. Current Anthropology 24(2):181–202. FuYou. 2006. Optical dating of the Jingshuiwan Paleolithic site of Three Yin, GongMing, Sun Yingjie, Ye YuGuang, and Liu Wu. 2001. The ESR age Gorges, China. Chinese Science Bulletin 51(11):1334–1342. of shells from the bed of Dali with human fossil. Acta Anthropologica Sinica Pope, Geoffrey G. 1989. Bamboo and human evolution. Natural History 20:34–38. 89(10):48–57. Zhang Li, Shen Guanjun, Fu Renyi, et al. 2007. Preliminary U-series dating Schick, Kathy D. 1994. The Movius line reconsidered: perspectives on the of hominid locality Miaohoushan in Benxi, Liaoning Province. Southeast earlier Paleolithic of eastern Asia. In Integrative paths to the past. Robert S. Culture 2007(3):54–57. Corruccini and Russell L. Ciochon, eds. Pp. 569–596. Englewood Cliffs, Zhang, SenShui. 1989. The Lower Paleolithic cultures in North China. In NJ: Prentice Hall. Ancient humans in China. Wu Rukang, Wu Xinzhi, and Zhang Senshui, Schick, Kathy D., and Z. Dong. 1993. Early Paleolithic of China and eastern eds. Pp. 97–158. Beijing: Science Press. Asia. Evolutionary Anthropology 2(1):22–35. ———. 1993. A study on the stone artifacts from Loc. 54:100 of the Dingcun Shen, GuanJun, and Jin LinHong. 1992. U-series dating of speleothem samples site. Acta Anthropologica Sinica 12(3):195–213. from Guanyindong Cave at Qianxi County, Guizhou Province. Acta An- ———. 1999. On the important advancement of the Paleolithic archaeology thropologica Sinica 11(1):93–100. in China since 1949. Acta Anthropologica Sinica 18(3):193–214. Wang, Shejiang. 2005. Perspectives on hominid behaviour and settlement pat- Zhang, ShuangQuan, Li ZhanYang, Zhang Ye, and Gao Xing. 2009. Mortality terns: a study of the Lower Paleolithic sites in the Luonan Basin, China. Oxford: profiles of the large herbivores from the Lingjing, Xuchang man site, Henan British Archaeological Reports. Province, and the early emergence of the modern human behaviors in East Wang, Wei, Liu Jun, Hou YaMei, Lynne A. Shepartz, S. Miller-Antonio, W. J. Asia. Chinese Science Bulletin 54:3857–3863. Current Anthropology Volume 54, Supplement 8, December 2013 S371

Identifying Mechanisms behind Middle Paleolithic and Middle Stone Age Cultural Trajectories

by Francesco d’Errico and William E. Banks

A critical analysis of the debate that has surrounded the emergence of “modern behavior” during the last two decades and new ways to study material culture and human-environment relationships allow us to design a novel approach with which we can understand the mechanisms that have led human populations to develop the variety of cultures that we recognize today. We propose a methodological framework that moves away from narrative explanations for the origin of “behavioral modernity” and instead focuses on the interplay between cultural adaptation and environmental change. We argue that by applying this approach to the many different instances of cultural change as well as stasis that characterized the last 300 kyr of human societies we may identify the mechanisms that have led us to become what we are and, if any, the underlying trends that guided this process.

Introduction chaeological records over the past 300 kyr (Marean et al. 2007; McBrearty and Brooks 2000). The idea that the emergence of cultures such as ours was abrupt nevertheless remained in Twenty years ago, the path followed by humans to attain play, and some viewed innovations occurring in southern “behavioral modernity” appeared evident to most archaeol- Africa ca. 70 ka as the factor that allowed cognitively modern ogists and paleoanthropologists. Best exemplified by the pub- AMHs to qualitatively change their adaptive abilities and rap- lication of The Human Revolution monograph (Mellars and idly expand out of that continent ca. 60 ka (Mellars 2006). Stringer 1989), this path was short, abrupt, exclusively as- In parallel, other researchers proposed that “modern” cog- sociated with anatomically modern humans (AMHs), and best nition was associated with various members of our lineage, reflected in the European Upper Paleolithic archaeological not just AMHs, and that social and demographic factors, record. However, apart from a possible neurological switch arguably triggered by climate change, could explain the asyn- (Klein 2000, 2009), no solid cause was proposed that could chronous emergence, disappearance, and reemergence of key explain why this should have happened where it did and in cultural innovations among both African Middle Stone Age such an instantaneous way. Subsequently, the discovery that and Eurasian Middle Paleolithic populations (Conard 2008; AMHs originated in Africa (Henn et al. 2011; Trinkaus 2005; d’Errico 2003; d’Errico and Stringer 2011; Hovers and Belfer- Weaver and Roseman 2008) along with a growing body of Cohen 2006; Langley, Clarkson, and Ulm 2008; Nowell 2010; archaeological evidence supporting the emergence of key cul- Zilha˜o 2001, 2007). The partisans of this model relied on tural innovations in that continent before the purported Eu- cultural innovations found in the Neanderthal archaeological ropean “revolution” ca. 40 ka gave rise to a different explan- record (burials, use of pigments, complex lithic and hafting atory model: “modern behavior” must have developed technologies, and personal ornamentation at the end of the gradually in Africa as a consequence of the origin of our Neanderthal evolutionary trajectory) to counter the idea that species there and would have been expressed by a process of behavioral modernity is unique to our species. The recent gradual accretion of innovations observed in the African ar- finding that significant interbreeding occurred between Ne- anderthals, Denisovan, and modern populations originating Francesco d’Errico is Director of Research and William E. Banks in Africa (Green et al. 2010; Meyer et al. 2012; Reich et al. is Research Associate at the University of Bordeaux, Centre National 2010) is used to support such a scenario because it blurs de la Recherche Scientifique-Unite´ Mixte de Recherche de la previously accepted taxonomic boundaries between Upper Pre´histoire a` l’Actuel: Culture, Environnement, Anthropologie, Equipe Pre´histoire, Pale´oenvironnement, Patrimoine (Baˆtiment Pleistocene hominins. In parallel with these scenarios, which B18, Avenue des Faculte´s, 33405 Talence, France [f.derrico@pacea are in one way or another anchored in the archaeological and .u-bordeaux1.fr]). This paper was submitted 3 VII 13, accepted 1 genetic records, some researchers have proposed that the evo- VIII 13, and electronically published 8 XI 13. lution of inherent components of present-day modern cog-

᭧ 2013 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved. 0011-3204/2013/54S8-0015$10.00. DOI: 10.1086/673388 S372 Current Anthropology Volume 54, Supplement 8, December 2013 nition may have played a key role in reaching the cognitive mography is important but that it is only one factor in the capacity expressed in our species and the ensuing “complex- spread and maintenance of innovations, not the prime mover ification” of material culture. Altruism (Bowles 2009), en- nor the unique explanatory mechanism behind their adop- hanced memory (Wynn and Coolidge 2010), complex lan- tion. guage (Dunbar 2003), increased capacity for social learning In parallel with the proposition of these various explanatory (Mesoudi, Whiten, and Laland 2006; Richerson, Boyd, and models and triggers, a number of researchers have called into Bettinger 2009; Tomasello 1994), creation of adapted learning question whether behavioral modernity is the appropriate environments (Sterelny 2011), hierarchical mental construc- concept with which to identify when we became as we are. tions (Gibson 2007), and acquisition of syntactical language One such criticism has been directed at the reliance on ma- (Bickerton 1990) have each been proposed as the prime mover terial culture trait lists to recognize the crossing of the thresh- that allowed AMHs to cross the threshold of behavioral mo- old to modernity. An early example of the use of a trait list dernity. It has been argued, though, that the capacity for is by Mellars (1989), who used disjointed elements of the “modernity” resulting from a speciation event is a necessary Upper Paleolithic archaeological record to establish a sup- but not sufficient condition for the development of modern posed cognitive threshold between Neanderthal and AMH, cultural traits. In this vein, a number of scholars have explored suggesting that the latter had crossed the Rubicon to behav- the roles played by population pressure (Compton 2011) as ioral modernity by the beginning of the Upper Paleolithic. well as demography and cultural transmission (Henrich 2004; However, Deacon (1990) cautioned against the use of a Eu- Powell, Shennan, and Thomas 2009; Shennan 2001) in the rocentric yardstick for measuring cognitive modernity by spread and maintenance of cultural innovations. By applying pointing out that there were indications that these criteria modeling techniques, these latter authors argue that popu- were not pertinent in other regions of the world, where this lation size, density, rates of reproduction, and networks of process appeared to have followed a different tempo and information exchange determine whether or not cultural in- mode. After the publication of the more comprehensive trait novations can be diffused, maintained, and in some cases lost. list proposed by McBrearty and Brooks (2000), d’Errico The interest of their results is that they account for such events (2003) noted that instead of being based on universal features without invoking speciation as a prime mover and that their found in all present-day human cultures, this list was a mix- expectation better fits the archaeological record than models ture of traits that the authors had recognized in the Middle predicting abrupt or incremental changes in behavior. How- Stone Age and the Upper Paleolithic. This naturally led them ever, it has been argued that while demography may be key to recognize behavioral modernity in the records on which to the diffusion and maintenance of innovative behaviors, it their list was based and create the misleading impression of does not necessarily favor their emergence and acceptance. a gradual accretion of these traits in Africa. In his view these Each society has its own way of regulating the acceptance of trait lists cannot necessarily be applied to human populations deviations from a behavioral norm, which produces different that lived in dramatically different environments and that had postures toward innovation even when potentially advanta- different cultural trajectories. While proposing that behavioral geous (Bar-Yosef and Belfer-Cohen 2011). The spread of in- modernity played a useful role in highlighting the inconsis- novations is also dependent on a society’s ability to create tencies of the “human revolution” model and showing that settings of high-fidelity learning: even with sufficient popu- more gradualistic patterns of cultural change existed outside lation size, if such settings are absent, innovations may not of Europe, d’Errico (2003) argued that behavioral modernity be maintained (Sterelny 2011). Another shortcoming, as was no longer a useful operational concept and that one pointed out by d’Errico and Stringer (2011), is that Powell, should instead explore the emergence of cultural innovation Shennan, and Thomas (2009) signal climate change as a ge- in each region of the world irrespective of the taxonomic neric factor behind the increase or fragmentation of popu- affiliation of the population in question. lations in their demographic-based “mechanism,” but they Zilha˜o (2011) points out that the real problem with be- offer no means of testing this, especially when considering havioral modernity is that it is based on the notion that dif- that specific climate changes had very different effects in var- ferent species of Homo were each characterized by a unique ious regions of the world. Implicit in their (Powell, Shennan, set of behaviors and that modern behavior is unique to AMHs. and Thomas 2009) contention that favors demography is the He argues that attempts to define “modern human behavior” assumption that in order to produce the spread and main- in opposition to “Neanderthal behavior” have consistently tenance of innovations, this demographic-based mechanism shown that some modern humans were “behaviorally Ne- is only relevant to AMH populations. It has been noticed that anderthal” and some Neanderthals were “behaviorally mod- contrary to their assumption, such a mechanism is equally ern” (Zilha˜o 2006). In his view, there is no such thing as applicable to archaic hominins and that differences between Neanderthal behavior because the archaeological record in cultural trajectories of Neanderthals and AMHs may have both Europe and the Near East demonstrates that Neanderthal been dependent on group size and rates of cultural exchange adaptations span the entire range of ethnographically docu- rather than on built-in differences in cognition (d’Errico and mented settlement-subsistence strategies. Nowell (2010) also Henshilwood 2011). These criticisms serve to show that de- highlights the concept of modernity’s ambiguous status due d’Errico and Banks Mechanisms behind Cultural Trajectories S373 to its routine association with the development of AMH so- hominins perceived by definition as a behaviorally different cieties, which is at odds with the presence of many instances biological species and therefore irrelevant in this debate. of modernity in Neanderthal populations. Comments on Shea’s paper (2011), which was published Henshilwood and Marean (2003) argue that behavioral mo- in Current Anthropology, welcome his proposed shift toward dernity is too loose a concept and that we should rely on the reconstruction of different cultural adaptations in specific clear evidence for symbolically mediated actions as the best environmental situations but point out that his search for way to recognize societies that have acquired “fully symbolic behavioral variability does not go very far. Comments by sapiens behavior.” However, when applying this concept to Nicholas J. Conard and Rick Potts in particular highlight the the archaeological record of southern Central Africa, Barham fact that behavioral variability, as Shea describes it, is as vague (2007) highlights how difficult it may be to identify the emer- as behavioral modernity because he does not propose with gence of such a capacity in records that hint at the presence the concept any clear operational tools with which one can of modernity but do not have the full suite of material culture evaluate cultural adaptations within their respective environ- that supports its “formal” recognition. This problem may be mental contexts. The tool he used, Clark’s system of tech- due to our lack of heuristic tools for recognizing fully symbolic nological modes, appears to them as highly inappropriate behavior when expressed in ways other than our preconceived because it is constructed on a logic of unilinear, incremental notions. His answer to this shortfall is to combine structuralist evolution, which is what Shea (2011) is in principle trying to and ecological theory to recognize individual regional trajec- steer away from. We would add another remark: by placing tories that could be influenced in part by historical contin- the analysis of behavioral modernity in the realm of what he gencies. Belfer-Cohen and Hovers (2010) concur in that they perceives as a single species, Shea has unwittingly found the consider behavioral modernity to be a loose and poorly op- prime mover or “donor” of his underlying definition of be- erational concept that does not create a link between cognitive havioral modernity—the processes that led to the emergence theory and the typical work of archaeologists. They argue that of our species in Africa. modernity is multifaceted and cannot be boiled down to a rigid checklist of presence/absence characteristics, and they From Pars destruens to Pars construens propose that we should focus instead on the circumstances and contexts in which the phenotypic expressions of moder- The above review of the debate makes clear that behavioral nity, which were at first erratic, became fixed features in the modernity and varying lists of cultural traits associated with archaeological records of different regions of the world. it are not useful tools for establishing the way in which we Additional criticism has been leveled at this concept by became what we are. Some consensus now exists that the Langbroek (2011), who proposes that the concept of moder- evolution of human societies in the last 300 kyr has followed nity as it is typically applied leads us to frame the evolution a multitude of paths, not necessarily progressive in nature, in of cognition in a unilinear and exponential manner when we which the material expression of modern cognition is rep- should rather envision this process as a branching one that resented by different mosaics of cultural innovations. Focus- mirrors Darwinian evolution. Within such a framework, “cog- ing on regional trajectories appears to be the only way to nitions” associated with different members of our lineage document cultural changes and ultimately the mechanisms would be submitted to varying selective pressures, thereby behind such changes. In doing so, we must seek ways to creating a variety of cognitive outcomes that cannot be clas- integrate environmental, ecological, demographic, and social sified as more or less modern. factors as well as historical contingencies in order to under- Shea (2011), like many others, criticizes the fact that be- stand how human populations have developed and in some havioral modernity is derived from a trait list founded on the cases lost and reacquired cultural innovations that we rec- Upper Paleolithic archaeological record, and he echoes ognize to be the cornerstone of the human experience. Among d’Errico’s (2003:189) argument that archaeologists from dif- those that accept this frame of thinking (d’Errico 2009; Hovers ferent cultural backgrounds could propose different features 2009; Kuhn 2013; Stiner 2013; and comments in Shea 2011 to define modernity. Furthermore, he adds that the concept by Lawrence S. Barham, Nicholas J. Conard, James F. has been constructed on a paleoanthropological narrative tra- O’Connell, and Rick Potts), there is consensus that although dition that implies the transformation from an inferior state these are factors that played a role in the process of cultural to a superior one. His answer to overcome the behavioral innovation, the way that they were organized and the interplay modernity conundrum is to focus on “behavioral variability.” between them remains to be understood, and pertinent heu- The idea is to take into account “modality, variance, skew, ristic tools with which to interrogate the empirical evidence and other quantitative/statistical properties” (Shea 2011:2) to are lacking. measure how successful specific cultural adaptations were Some might still question whether this endeavor should be from a cost-benefit standpoint in specific environmental set- conducted only in archaeological records associated with tings. Shea’s view is anchored in the notion that whatever AMHs or should also include archaic hominins. The former approach one takes to address this issue, it is solely the busi- would be a clear mistake in our view because it would con- ness of Homo sapiens sapiens and does not concern archaic strain, whether one would admit it or not, the analysis of S374 Current Anthropology Volume 54, Supplement 8, December 2013 local trajectories within a conceptual framework in which key in the multitude of interactions that occur between them. The behavioral innovations can only be the consequence of an- second is more abstract and seeks to encompass interactions atomical modernity. By equating behavioral and anatomical between factors and components with the goal of explaining modernity, no matter what “variability” or “cost-benefit” bal- such interactions with a simple model. The final goal is to ance is found, we will not get rid of the mind-set that bio- move from complexity to the proposition of a general ex- logical change is the prime mover. Such a stance would also planatory law (e.g., the mechanism of natural selection in deprive us of examining a significant number of cultural tra- biological evolution). jectories, thus compromising our ability to compare how dif- When put in the context of the debate surrounding Middle ferent populations reacted to comparable suites of external Stone Age and Middle Paleolithic cultural trajectories, the first stimuli. Encompassing all local trajectories is in our view the concept of mechanism can be seen as a useful operational best way to obtain a full picture of the many rapid cultural tool, and the second points to the ultimate goal of identifying experiments that are key features of the cultural evolution of the long-term trends and rules that have shaped the cultural our lineage. evolution of our lineage. One of the interests in approaching With this paper, we propose a methodological framework cultural evolution from the standpoint of mechanism is that that moves away from “narrative” explanations toward a focus we can describe cultural change dynamically through the anal- on material culture and the evaluation of the potential in- ysis of setup and termination conditions (Machamer, Darden, terplay between cultural adaptation and environmental and Craver 2000). The former typically represent in the in- change. We think that by applying this approach to the many vestigation of mechanism the relevant components, their different instances of cultural change as well as stasis that “structural” properties, spatial relations, and the causal factors characterize the last 300 kyr of human societies, we may iden- thought to influence the relationships between different com- tify the mechanisms that have led us to become what we are ponents. Each component has some degree of variability and and the underlying trends, if any, that guided this process. independence such that identical setup conditions can result in two systems following different trajectories. Between setup Causes versus Mechanisms and termination, intervening factors or entities can influence the interactions between components within the mechanism, We argue that the primary problem with most of the scenarios thus influencing the direction the system follows. Termination proposed to explain the emergence of behavioral modernity conditions are idealized states that represent a point from is that they are based on single-cause models. Such models which one can infer how the mechanism functioned. In this are founded on the teleological notion that a unique cause context, “termination” is neither synonymous with equilib- will continuously act as the sole or the dominant factor in rium nor a moment in which the process has necessarily producing the observed outcome. Some of the proposed reached a terminal state. In reality, such states are idealized causes would have acted as long-term stimuli, such as altruism conditions, and their choice by researchers is often deter- or enhanced working memory. Others, such as genetic mu- mined by the heuristic and analytical tools they have at their tation or a bottleneck event (e.g., Toba super eruption: Am- disposal to understand the interplay between components and brose 1998), would have been short lived in nature and have causal factors within a perceivable time frame. This move had a relatively immediate effect. Both types of causes, how- between setup and termination is analogous to the concept ever, are not sufficient to explain the complex paths and mul- of “adaptive cycle” that is used in resilience theory to char- titude of adaptations that a growing body of archaeological acterize the dynamics of a socioecological system with respect data from Africa and Eurasia denotes. to external stimuli and internal processes (Holling 1973; Rather than causes, we need to focus on identifying the Schoon et al. 2011; Walker et al. 2004). mechanisms that have led different societies to develop spe- Setup conditions are of course the result of prior processes, cific cultural adaptations as a means of coping with external so in the study of mechanisms in a componential causal sys- stimuli (both environmental and cultural). “Mechanism” is tem framework one is beginning the investigation at a specific a term that has been defined in a number of ways, and Ku- point along a continuum. Human cultures follow continuous orikoski (2009) points out that it is difficult to come up with trajectories in which the various factors that play a role in a definition that satisfies all theoretical needs and potential producing change continuously interact. However, as archae- research practices. Drawing from the work of Bechtel and ologists, we identify in the archaeological record discrete and Abrahamsen (2005), Kuorikoski (2009), and MacHamer, Dar- recurrent associations of similar cultural items assumed to den, and Craver (2000), we define a mechanism as a con- reflect cohesive adaptive systems (CASs). We define a CAS as stellation of factors and components that through the process a cultural entity characterized by shared and transmitted of their interaction with one another stimulates the trajectory knowledge reflected by a recognizable suite of cultural traits of a system. The investigation of mechanisms functions at that a population uses to operate within both cultural and two different conceptual levels (Kuorikoski 2009). The first environmental contexts. The cultural traits used to define a consists of examining a componential causal system by dis- CAS can carry the same well-known ambiguities as those used assembling the role played by each component and factoring to define a techno-complex (Clarke 1968; Hodder 1991; Ren- d’Errico and Banks Mechanisms behind Cultural Trajectories S375 frew 1977). The archaeological record represents a very pale documented and seems to be clustered in regional traditions. reflection of the features that constituted a past cohesive cul- This is the case for personal ornamentation (Caron et al. 2011; tural group because human adaptation is shaped by cultural d’Errico et al. 2009; Peresani et al. 2011; Vanhaeren et al. rules that govern aspects of human behavior such as kinship 2006, 2013; Zilha˜o et al. 2010), symbolic use of pigments systems, marriage, gender politics, and symbolic or belief sys- (Roebroeks et al. 2012; Watts 2010; Zilha˜o et al. 2010), graphic tems. These rules define human societies as much as and expressions (d’Errico, Garcı´a Moreno, and Rifkin 2012; Hen- probably more than the environmental contexts they occupy. shilwood, d’Errico, and Watts 2009; Mackay and Welz 2008; As archaeologists we have limited means with which to infer Texier et al. 2010), and mortuary practices (Gru¨n et al. 2005; those rules for societies of the remote past. It is, however, Pettitt 2011). Although open to debate, we may assume that reasonable to think that those rules, or changes in those rules, such CASs also included behavioral features that have not shaped the material culture record that we have at our disposal survived in the archaeological record and that made each of and played a role in how a past group interacted with their these societies unique in the cultural history of our lineage. environment. The concept of CAS differs from that of a As discussed earlier, previous models proposed to explain techno-complex (Childe 1929) in that environmental param- the emergence of these cultural features and innovations have eters contribute to its definition. The difference lies in the typically been monocausal in nature and have not been geared fact that the approach that we detail below effectively explores toward identifying potential mechanisms and long-term potential links between cultural traits and ecological param- trends. The evolution of a human society cannot be reduced eters and evaluates this relationship through time, thereby to its demography; systems for transmitting and maintaining providing us the potential to identify general, long-term cultural innovations depend on a variety of factors. Recent trends. studies have pointed out that a number of ecological, his- Traditionally, archaeologists working with the Middle Pa- torical, and psychological variables appear to condition the leolithic and Middle Stone Age archaeological records have rules that societies impose on individuals and the degree of relied on lithic technology and stone tool typology to define tolerance a society has toward deviant behavior (Gelfand et past “cultures” and identify evolutionary trends. Regardless al. 2011; Henrich, Heine, and Norenzayan 2010; Norenzayan of the multitude of reasons for this focus, examinations of 2011). The cultural system that one inherits affects basic pro- ethnographically documented material cultures indicate that cesses such as perception, reasoning, motivation, and coop- stone tools represent just a small portion of the paraphernalia erative strategies. This seems to imply that each CAS is char- used by a hunter-gatherer population and do not necessarily acterized by a different potential for cultural transmission, reflect the complexity of cultural adaptation or its geography social learning, and the degree to which individuals are open (Hayden 1979). Relying on just a single element of a technical to accept, maintain, and communicate innovations (Gelfand system to represent or infer the complex suite of behaviors et al. 2011). It has also been argued that each individual and social rules that characterize a past cultural adaption is society’s ability to maintain effective social learning environ- clearly illusory. Data accumulated over the last decade in Af- ments is key (Sterelny 2011). It is still unclear whether the rica and Eurasia on populations that lived there during the reasons behind human behavioral variability are due to purely last 300 kyr have broadened our understanding of these CASs cultural processes leading to cultural divergence, to ecological by providing insight into a variety of domains beyond those constraints, to gene-culture coevolution (Norenzayan 2011), strictly related to lithic technology. These include technolog- or to some combination of these. Whatever the reason for ical behaviors that certainly are expressions of salient features these differences, each society can be seen as a complex system of those archaeological cultures. These behaviors include py- of attitudes and the potential they offer for change. In this rotechnology (Brown et al. 2009; Mourre, Villa, and Hen- context, we use attitude to refer to the way in which the shilwood 2010), mastic production (Caˆrciumaru 2012; Char- collective worldviews of individuals in a social group influence rie´-Duhaut et al. 2013; d’Errico et al. 2012; Lombard 2012; behavior. Bar-Yosef and Belfer-Cohen (2011) use such a ra- Pawlik and Thissen 2011; Villa et al. 2012; Wadley, Hodgskiss, tionale to explore patterns of human expansion out of Africa and Grant 2009), hafting techniques (Lombard 2005; Villa et with the debatable assumption that lithic technology is a fair al. 2009, 2012), projectile technology (Villa and Soriano 2010), reflection of these attitudes and therefore can be used as a techniques for small game capture (Stiner, Munro, and Su- proxy to trace successes and failures in hominin expansions. rovell 2000; Wadley 2010c), use of poison in hunting (d’Errico We argue that when identified and combined with the geo- et al. 2012), bone tool production (Backwell, d’Errico, and graphic and environmental settings in which societies oper- Wadley 2008; d’Errico, Backwell, and Wadley 2012; d’Errico, ated, an array of behaviors related to technical and symbolic Borgia, and Ronchitellli 2012; d’Errico and Henshilwood systems and reflecting inherited knowledge can be viewed as 2007), pigment processing and storage (d’Errico et al. 2010; the setup conditions for the processes at work behind the Henshilwood et al. 2011), and use of plants (Mercader 2009; suite of cultural experiments that took place in regions of Wadley et al. 2011). Symbolically mediated behavior, which Africa and Eurasia between 300 ka and 10 ka. appeared to be largely inexistent or largely out of our grasp The approach that we describe below entails means with a decade ago for the time periods in question, is now well which one can identify and follow the processes affecting CASs S376 Current Anthropology Volume 54, Supplement 8, December 2013 between setup and termination conditions with no a priori solve adaptive challenges. The problem one must address in assumption on the trajectories followed by the system being studies of the distant past is how to operationalize this frame examined. This approach is chosen to avoid the determinism of thinking such that one gives social dimensions and material inherent in a “single-cause” frame of thinking as well as the culture the attention they deserve. relativism implied in the behavioral variability approach. We As is already evident to most archaeologists, an important perceive the different regional trajectories as unique suites of method for inferring behavior and cognition is to view ar- cultural experiments with their own distinct setup conditions. chaeological remains as representing an ordered chain of While these experiments may have components and processes events, gestures, and processes belonging to a sequence of in common, they did not necessarily play the same role within actions that led to the transformation of a given material to each adaptive system. the finished form, that is, the chaıˆne ope´ratoire (Lemonnier Ideally in our field of study, understanding the processes 1986; Leroi-Gourhan 1964; Schlanger 1994). This concept is at work between setup and termination conditions would especially pertinent because it permits one to infer from the require one to understand the role played by each component finished artifact, production waste, and potentially missing and factor in a given region and time—environmental elements the socially shared and individual knowledge, co- changes, adaptive system-specific material culture character- operation, and amount of short- and long-term memory nec- istics, inferred differences in social rules and attitudes, ge- essary for the functioning, maintenance, and transmission of ography—and to understand how they interacted over time a given production sequence. It also allows one to understand to produce the outcome that we describe as the termination to some extent the mental template of the actors. conditions at a particular point in time. This would provide Different classes of material culture possess different po- insight into the internal functioning of regional or individual tentials to inform us on cultural cohesiveness, shared knowl- componential causal systems. A second goal would be to un- edge, and underlying cognitive processes. Until 10 years ago, derstand why for a different componential causal system in identifying setup conditions of Middle Stone Age and Middle a different region and time similar factors interacted in dif- Paleolithic “cultural experiments” would have meant focusing ferent ways to produce a different outcome. Through the almost exclusively on lithic technology, but now this has be- identification of commonalities and differences and by eval- come a more complex and potentially informative endeavor uating the role played by specific components and factors considering the many categories of material remains that we within individual trajectories as well as the interplay between have recognized since then (bone tools, pigments, personal them, one should be able to identify overarching trends in ornaments, engravings, mastic compounds, poisons, contain- the way in which systems operated. This process would allow ers, use of plants and feathers, etc.). Because of their ubiquity us to move toward the goal of finding “the general law,” if and durability, lithic artifacts remain a valuable class of ma- any, that operated behind the evolution of componential terial for identifying the geographic extent of a CAS. In par- causal systems. ticular, consistencies in lithic technologies and formal tool types have allowed researchers to identify discrete cultural Regional Trajectories as Cultural Experiments adaptations and begin to piece together their geographic dis- tribution and chronological context in parts of Africa and How do we put this approach into practice with the archae- Eurasia (Barham 2001; Belfer-Cohen and Hovers 2010; Del- ological, chronological, and paleoenvironmental data that we agnes and Meignen 2006; Discamps, Jaubert, and Bachellerie have at hand? Gaining insight into the more remote aspects 2011; Kuhn 2013; Soriano, Villa, and Wadley 2007; Villa et of cultural systems, socially shared knowledge, and attitudes al. 2010, 2012; Wurz et al. 2003). The form that lithic raw toward innovations is typically thought of as being a difficult material acquisition takes (local or long-distance direct ac- endeavor for archaeologists. We argue, however, that useful quisition, trade, etc.) can be a reliable proxy for how an adap- inferences can be made pertaining to these cognitive and so- tive system is linked to a given territory. The importance cial domains through the detailed analysis of a wide range of assigned to specific types of raw material can provide insight material culture. When such inferences are placed within pa- into how rigid or flexible a lithic technical system is and how leoenvironmental and landscape contexts, we possess an array natural resources became key elements in the cultural system. of data that represent the setup conditions from which an Acquisition patterns can also reflect the presence of social investigation into mechanism can be launched. By applying networks and their complexity. Analysis of debitage tech- methods that we detail below, one has the ability to identify niques can hint at the form and duration of apprenticeship the key features and the degree of cohesiveness of these sys- likely necessary to produce certain classes of tools. Shaping tems and to track the way in which they evolved and possibly techniques to transform knapped blanks into finished tools, responded to environmental change through time. An interest the degree of conformity to strict stylistic rules, and whether in culture-environment interactions is by no means new, and formal tools were used for single or multiple functions allow a number of scholars have already argued that a culture’s us to understand the technical system’s degree of plasticity “core” (Steward 1955; see also Odum 1971, and for a synthesis and to gauge the nature of information needed to maintain Johnson and Earle 2000) can be seen as a society’s means to and transmit the required know-how. Finally, whether each d’Errico and Banks Mechanisms behind Cultural Trajectories S377 phase of the chaıˆne ope´ratoire is found in a unique location past. Taphonomic analyses of the various categories of sym- or multiple locations can inform us not only about how a bolic items may help to identify which classes of artifacts are population organized its activities across the landscape but especially affected by taphonomic processes and infer whether also about the planning involved and the complexity of the such processes may have led to their disappearance at some cognitive maps that characterized an adaptive system. sites. Cross-cultural analyses of the raw materials used to During the last decade, improvements in excavation tech- produce symbolic items and the evaluation of their respective niques and reappraisal of existing collections have identified durability are also means to address the issue of the loss of a variety of material culture remains other than lithics, and some elements of symbolic material culture, particularly in methods have been developed to better infer their behavioral contexts in which we observe other material remains pointing significance. During the Middle Stone Age and the end of the to the presence of symbolic mediated behaviors. These be- Middle Paleolithic, we observe for the first time bone tools haviors may exist in a society that does not express them produced with techniques specifically conceived for this me- through purely symbolic items but rather embodies them in dium—such as grinding, scraping, and polishing—that al- functional items. These items express symbolic meaning by lowed their final shape and size to be achieved with a high their adherence to strict stylistic norms that archaeologists degree of accuracy. This class of tool is considered particularly perceive as modern without having the means to disentangle appropriate for the characterization of technical systems, functional from symbolic traits (Barham 2007). Taking into tracking technical changes through time, identifying regional account such items as components of a CAS is a means of variability, disentangling style from function, and inferring incorporating them, including their potential symbolic value, the complexity inherent in a given adaptive system (Backwell among the factors that played a role in the relationship be- and d’Errico 2005). tween culture and environment. In this way, potentially sym- The production of complex compounds entails the ordered bolic aspects of past material culture become full actors in combination and modification, often with the use of pyro- the exploration of mechanisms governing the evolution of technology, of a variety of raw materials in order to produce cultural systems. an end product that has physical properties not found in the Second, advanced analytical techniques contribute to the natural world. Beyond the important ability to develop such disentanglement of accidental from purposeful behavior, thus technologies through experimentation, what is paramount is allowing a precise documentation of the operational chain the ability to maintain and transmit these innovations. Once and underlying cognitive processes. Actualistic studies con- adopted, each recipe and the way it is employed can become ducted with the aim of verifying the purposeful alteration of key features of a CAS and have a feedback effect on other pigment (Wadley 2010b) or shell bead color (d’Errico et al. aspects of the technical system as well as on how knowledge 2009; Kandel and Conard 2005) as well as assessing whether is accumulated and shared. We now possess means to infer plant remains are present naturally or because of specific hu- how these compounds were made and used by specific human man actions such as use as bedding and as insect repellent populations in the Middle Paleolithic and the Middle Stone (Wadley et al. 2011) are good examples of evaluating the Age (Bo¨eda et al. 1996; Caˆrciumaru et al. 2012; Charrie´- behavioral significance of past human agency. Archaeozoo- Duhaut et al. 2013; d’Errico et al. 2012; Hauck et al. 2013; logical, taphonomic, chemical, technological, and functional Henshilwood et al. 2011; Mazza et al. 2006; Pawlik and This- analysis of symbolic items relying on actualistically established sen 2011; Villa et al. 2012; Wadley 2010a; Wadley, Hodgskiss, criteria, when placed within the framework of a chaıˆne op- and Grant 2009). Bitumen, birchbark tar, wood resin mixed e´ratoire, have provided new means to identify and analyze with hematite, hematite powder mixed with animal fat: we early instances of symbolically mediated behavior. This pro- are starting to understand how different CASs have created vides insights into the way in which material expressions of and incorporated comparable compound technologies as a symbols were created, assembled, and displayed, how and for response to specific needs in different environmental settings. how long they were used, and to what degree those early We are also moving beyond the simple recognition that symbolic systems are comparable with those created by eth- instances of symbolic material culture in the form of personal nographically documented human societies. Adapted theo- ornaments, engravings, pigment production and storage, and retical frameworks are proposed to understand the amount decorated bone items are present in the archaeological record and nature of information that one can convey through each of this period and reflect modern behavior to the exploration category of the identified type of symbolic material culture of the representativeness and significance of specific instances. (Kuhn and Stiner 2007) and the function personal ornaments First, one can wonder how much evidence has not been pre- may have played in prehistoric societies (d’Errico and Van- served or, if preserved, been destroyed during excavation or haeren 2007). By establishing to what degree specific instances gone unrecognized. It has been pointed out, for example, that of symbolic behavior are representative of a CAS and ex- the appearance of symbolic items in the archeological record ploring patterns of variability through time and space, we are may be largely conditioned by taphonomic processes (Barham able to verify whether these instances are inherent features of 2007). This problem can be partially overcome by critically the system or simply the expression of subregional or suc- examining excavation techniques used at specific sites in the cessive cultural trajectories (Vanhaeren et al. 2013). S378 Current Anthropology Volume 54, Supplement 8, December 2013

A key additional dimension that must be considered when realistically incorporate the environmental conditions of each establishing setup conditions of any cultural experiment is phase that characterize the variable climatic conditions of this space. This is particularly relevant in the case of hunter-gath- period. erer economies because we know, apart from a few cases in In a series of papers published during the last few years, which particular adaptations allowed them to be sedentary or we have outlined an approach with which we can explore the semisedentary, that one key feature of these populations is interactions between CASs and paleoenvironment and un- that they were mobile. Although hunter-gatherer economies derstand how environmental dynamics may have influenced and their organization across the landscape can be broadly these adaptations and the distribution of prehistoric hunter- classified as logistical or residential (Binford 1980; Foley gatherer populations (Banks et al. 2008b, 2009, 2011; d’Errico 1992), we know that reality is more complex and must have and Stringer 2011). This approach, termed eco-cultural niche been so in the past. We must delve deeper into each individual modeling (ECNM), integrates archaeological, chronological, case and attempt to understand the unique logic behind each geographic, and paleoclimatic data sets via biocomputational settlement-subsistence system within its particular landscape architectures derived from biodiversity studies (Peterson et framework. We now have the means to reconstruct, at both al. 2011) to estimate ecological niches and distributional areas continental and regional scales, the effects that ice-sheet and occupied by prehistoric hunter-gatherer populations and to sea-level changes (Lambeck, Esat, and Potter 2002) had on identify and quantify the environmental factors that shaped the landscape. Information pertaining to environmental con- these niches. ditions has traditionally been reconstructed on the basis of An eco-cultural niche is defined as the range of environ- proxies from individual sites or groups of sites such as faunal mental conditions (i.e., the ecological niche) exploited by a or plant remains and soil characteristics, among others. These CAS (see Banks, d’Errico, and Zilha˜o 2013 for a detailed certainly provide important information, but if one wishes to discussion of eco-cultural niches). ECNM assumes that we explore the systemic and dynamic relationship between cul- can characterize a past cultural niche by employing methods ture and environment, one must move beyond the site scale used to reconstruct and study ecological niches. A key feature and find a way to examine this relationship at regional and of such predictive architectures is that they can project the continental scales. ecological niche predicted for a climatic phase onto the en- vironmental conditions of a subsequent period. The resulting An Integrative Approach niche projection is compared with the locations of known occurrences for the latter period to see whether or not it Research seeking to understand the relationship between hu- successfully predicts their presence within the niche. In this man and/or cultural evolution and climate change during the way, one can evaluate whether an adaptive system, in the event Middle and Upper Pleistocene is becoming an increasingly of its persistence, exploited the same ecological niche across widespread field of study (Bocquet-Appel et al. 2005; Carto different climatic phases or significantly expanded or con- et al. 2009; Compton 2011; deMenocal 2011; d’Errico and tracted it. This approach parallels the inquiry into the mech- Sa´nchez-Gon˜i 2003; Discamps, Jaubert, and Bachellerie 2011; anism driving the evolution of a componential causal system Gamble et al. 2004; Jacobs et al. 2008; Lowe et al. 2012; Maslin because it allows one to define the setup conditions of the and Christensen 2007; Osborne et al. 2008; Richerson, Boyd, process at work, in our case an individual cultural experiment, and Bettinger 2009; Sepulchre et al. 2007; Van Andel and and evaluate its termination conditions (i.e., the attributes Davies 2003). Most of this research has attempted to explore and distribution of a CAS at the end of the process) within the role played by climate on either long-term or sudden either the framework of environmental change or relative sta- evolutionary/cultural changes/replacements, but they do not sis. detail means with which to verify the proposed causal con- For data inputs, ECNM requires the geographic coordinates nection or test alternative hypotheses. Only a handful of stud- of archaeological sites bearing cultural features that are con- ies have designed tools to test the relationship between these sidered distinctive of a particular CAS or consistent subsets factors and explore their interaction at regional scales (e.g., within that system along with a set of raster geographic in- Bocquet-Appel and Tuffreau 2009). These studies attempt to formation system data layers summarizing environmental di- test the correlation between a single and often qualitative mensions potentially relevant to shaping the eco-cultural climatic variable (e.g., isotopic stage, cold/warm) and the niche exploited by the CAS as well as its spatial expression function of sites or the nature of the material record in a during a specific climatic phase. Geographic variables are as- given region. Such an approach, however, does not evaluate sumed to have remained relatively constant over the past 300 the various components of a given CAS against multiple and kyr, and thus one can use high-resolution present-day data quantitative climatic environmental variables. Other studies (e.g., ETOPO1). Reconstructions of past sea-level fluctuations use a variety of modeling techniques to contrast Neanderthal at both general and regional scales are available and can be and AMH population dynamics and interactions (Barton et used to reconstruct coastlines and related paleogeography for al. 2011; Fabre et al. 2011), but they produce results that are the region of study. Reconstructions of ice-sheet volume and difficult to test against the archaeological record and do not location are available for most of the last climatic cycle and d’Errico and Banks Mechanisms behind Cultural Trajectories S379

Figure 1. Schematic rendition of how one of the predictive architectures, in this case genetic algorithm for rule-set prediction, reconstructs an eco-cultural niche. 1, Occurrence data (i.e., location of archaeological sites belonging to a cohesive adaptive system) are resampled randomly by the algorithm to create training (b) and test data sets. An iterative process of rule generation and improvement then follows in which an inferential tool is chosen from a suite of rule types and applied to the training data (S1–Si) and paleoenvironmental raster data layers (a) to develop specific rules (Stockwell and Peters 1999). These rules evolve to maximize predictivity by several means (e.g., crossing over among rules), mimicking chromosomal evolution. Predictive accuracy is evaluated based on an independent subsample of presence data and a set of points sampled randomly from regions where the species has not been detected. 2, The resulting rule set defines the distribution of the subject in environmental dimensions (i.e., the ecological niche; Sobero´n and Peterson 2005), which is projected onto the landscape to estimate a potential geographic distribution (Peterson 2003). can be inferred, with some incertitude, for more ancient pe- obtain reconstructions of vegetation cover compatible with riods. With respect to paleoclimatic variables (temperature the targeted climate state. In this way, one obtains values for and precipitation), there exists a variety of modeling tech- precipitation, temperature (mean annual, coldest month, niques for obtaining reconstructions that can be integrated warmest month), and broad vegetation types. During this into a niche modeling approach. One can run (1) a coupled process, outputs are compared with paleoenvironmental data ocean-atmosphere general circulation model (e.g., IPSL CM4 to test whether the simulations capture past conditions sat- and CM5A; Dufresne et al. 2013; Kageyama et al. 2013), (2) isfactorily, and if they do not, there exist means to improve an atmosphere-only model with a slab ocean component (rep- subsequent generations of simulations in an effort to better resenting the top 50 m of the water column; Kang et al. 2008), capture past paleoclimatic conditions. or (3) an atmosphere-only model with imposed sea surface Other methods exist, such as using statistical techniques to temperature (SST) values (Kageyama et al. 2005). With all infer past climatic conditions from a variety of vegetation three, boundary conditions (orbital parameters, greenhouse data (for a review, see Tingley et al. 2012), but so far they gas concentrations, ice-sheet volume) appropriate for the tar- lack the spatial resolution required for our purposes. Of geted climatic event are assigned. The atmosphere-only model course, models are just that, models, and one must keep in with imposed SSTs also can be run with a refined resolution mind that they only approximate past climatic conditions. (∼50 km) over the region(s) of interest (see Banks et al. 2008a; Also, our goals are different from those of paleoclimatologists Sepulchre et al. 2007). who seek to understand the functioning and evolution of the The results from the different methods listed above can be earth’s climate system. We wish to have at our disposal the statistically downscaled (e.g., Vrac, Stein, and Hayhoe 2007) most accurate simulation for paleoclimatic conditions at a to increase the resolution of the simulated paleoclimatic data. specific time in the past and seek to improve our means for A final option is to use a regional model forced by general evaluating the pertinence of paleoclimatic simulations. Be- circulation model outputs as boundary conditions, thereby cause of the pressure of the threat of global warming, climate producing climatic simulations with a resolution as fine as 5 modeling is a rapidly evolving field of research, and it is clear km (Frei et al. 2006). This high level of resolution, or even that means to evaluate the pertinence of high-resolution sim- finer when possible, is most appropriate for examining cul- ulations at regional scales will rapidly improve in coming tural and niche trajectories at a regional scale. The outputs years. We will greatly benefit from such improvements. of this simulation process can be used to force a dynamic A number of predictive modeling approaches are available global vegetation model (e.g., Orchidee, Spitfire) in order to (climatic envelope range, generalized linear model, general- S380 Current Anthropology Volume 54, Supplement 8, December 2013

they are interpredictive either within a single climatic event or between two different events. Two niches are interpredictive when their observed degree of similarity is greater than would be expected by chance. The interest in following this approach lies in that we move beyond the analysis of site distributions to that of niches and can evaluate and quantify patterns of continuity or niche shifts. Predictive algorithms allow us to evaluate whether a CAS has conserved, expanded, or contracted its ecological niche in the time span between two different points in time. It is noteworthy that such changes may not necessarily be reflected in observed changes of their geographic range. In the event of climatic change, if an adaptive system conserves its ecological niche and simply tracks its shifting footprint, this can result in either an expansion or contraction of its geographic range depending on the range that the related climatic envelope occupies following the climatic shift (fig. 2). With this approach, changes in eco-cultural niches are assessed without a priori assumptions on the role played by environment. ECNM can identify cases in which significant cultural change, potentially reflecting changes in social rules Figure 2. Schematic illustration of how, following a climatic change, the conservation of an eco-cultural niche (a) may result and related organization along with niche shifts, appear to be in either a contraction (b) or an expansion (c) of the niche’s unrelated to environmental variability. Also, it is often in- geographic range. tuitively assumed that cultural innovations indicate an ability to better exploit environments and increase a population’s geographic range. However, in some instances innovations ized additive model, genetic algorithm for rule-set prediction, may reflect responses to environmental change such that they maximum entropy, ensemble approach; for a review see Ar- allow a population to maintain its niche and avoid niche au´jo and New 2007 as well as Pearson et al. 2006) for re- contraction. Situations may exist in which cultural innova- constructing the ecological niche of a CAS and its geographic tions are associated with a niche contraction. This may occur distribution using the above data. Arau´jo and New (2007) either because a CAS copes with climate change by targeting point out that ideally one should use multiple modeling meth- a smaller subset of its previous niche or because, in spite of ods and compare their outputs. Therefore, in our previous innovations, the CAS is unable to maintain its previous niche. applications of ECNM we have employed genetic algorithm We can envision four different scenarios in the evolution (genetic algorithm for rule-set prediction) and maximum en- of a CAS between setup and termination conditions (fig. 3). tropy (Maxent) methods. At a very general level, these ar- In the first scenario, the material culture characterizing a CAS chitectures first identify shared paleoenvironmental parame- remains the same between these two points in time. The ters among the geographic locations of archaeological sites second scenario features changes in some aspects of the ma- belonging to the same culture and then find other geographic terial culture, but one can identify a clear continuity through regions where these parameters are present, thus predicting time. From the perspective of resilience theory, these two the total ecological range of the target population (fig. 1). situations can be characterized as ecological resilience (Pee- When estimating ecological niches, it is important to consider ples, Barton, and Schmich 2006), which describes the situa- the geographic areas that would have been accessible via dis- tion in which an adaptive system is changed and reorganized persal to the population in question and that have been sam- while maintaining its key features and functions. For the third pled such that occurrences could have been detected (Barve scenario, material culture disappears from the archaeological et al. 2011; see Banks, d’Errico, and Zilha˜o 2013 and Banks record, indicating that populations are no longer present in et al. 2013 for archaeological examples); this area is termed the region or that they are archaeologically invisible. In the “M” in the BAM framework of Sobero´n and Peterson (2005). fourth scenario, the material culture associated with the ter- Once meaningful ecological niche estimations have been pro- mination conditions is clearly different from that of setup duced, statistical methods are used to identify the environ- conditions (e.g., a different techno-complex). Of course, the mental factors that shaped these niches and to measure their recognition of scenarios 1, 2, and 4 assumes that archaeol- breadth. Similarly, a variety of methods exist (e.g., background ogists have the means to identify and follow the cultural pro- similarity test: Warren, Glor, and Turelli 2010; partial-ROC cesses at work in the various aspects of the technological and test: Peterson, Papes¸, and Sobero´n 2008) to test whether two symbolic domains. Scenarios 3 and 4 imply that at some CASs’ eco-cultural niches are significantly different or whether undetermined point in time between setup and termination Figure 3. Scenarios of a cohesive adaptive system’s (CAS’s) evolution between setup and termination conditions within the framework of climatic stability (A–D) or climate change (E–H). Columns depict setup conditions and one of four possible termination conditions: (1) niche conservation, (2) niche expansion, (3) niche contraction, or (4) disappearance. A and E illustrate situations in which material culture remains essentially unchanged (described as scenario 1 in the text). B and F depict situations in which the material culture changes but clear continuity between setup and termination conditions can be recognized (scenario 2). In instances C and G, a CAS disappears or becomes archaeologically invisible at termination, representing either an extreme instance of niche contraction or migration (scenario 3). D and H are instances in which clear discontinuities in the material culture are observed between setup and termination (scenario 4). The potential for shifts in geographic range is indicated by displacement of the eco-cultural niche within the frame representing termination conditions. S382 Current Anthropology Volume 54, Supplement 8, December 2013

Figure 4. Idealized example of a long-term regional cultural trajectory (a) composed of multiple stages in which termination conditions become setup conditions for the subsequent stage, all of this encompassing multiple periods that may be characterized by either relative climatic stability or climatic change. Trends in niche variability (b) synthesize long-term trends in the relationship between cohesive adaptive systems and environmental variability at a regional scale. conditions, a process or event occurred that led to the dis- narios taking place during a period of relative climatic stasis, appearance of the initial CAS or to its evolution into a sig- comparing the initial and final eco-cultural niches and quan- nificantly different cultural system, respectively. tifying potential differences can be accomplished with back- Climate change may or may not have occurred between ground similarity tests (Warren, Glor, and Turelli 2010; for setup and termination conditions for the above scenarios. The an archaeological application see Banks et al. 2011). For sce- outcome of these eight different trajectories (the four sce- narios that occurred across a period characterized by a cli- narios with or without climate change) may involve niche matic change, the setup eco-cultural niche is projected onto stability, contraction, or expansion (i.e., 20 possible different the climatic conditions of the termination period to evaluate outcomes; fig. 3). Once one has determined into which sce- whether or not it changed or remained stable. These initial nario the archaeological record under investigation falls, and projected eco-cultural niches can be compared with a ECNM tools allow one to determine whether ecological fac- variety of statistical methods (e.g., partial-ROC comparisons: tors played a role in a given CAS’s trajectory or whether it Peterson, Papes¸, and Sobero´n 2008; cumulative binomial sta- was influenced to a greater extent by cultural processes. More tistic: Banks et al. 2008a; background similarity test: Warren, importantly, in the first case, dedicated statistical tools allow Glor, and Turelli 2010), again with the goal of evaluating one to quantify whether these changes are significantly dif- whether or not there was an eco-cultural niche shift during ferent from what would be expected by chance. For the sce- the process. d’Errico and Banks Mechanisms behind Cultural Trajectories S383

By determining whether or not an eco-cultural niche re- ture is viewed as unique, and one implicitly assumes that our mained stable, along with an identification of the key envi- job as archaeologists is to simply document cultural variability ronmental variables that played a role in the definition of the without the ability to identify underlying trends in the be- initial and final niches, and in conjunction with a detailed havioral evolution of our lineage. Alternatively, intrinsic in a reconstruction of the behaviors reflected in the material cul- number of regional examinations is the idea that one can ture record, we should be able to identify the different factors better identify, at that scale, one or more points in time during and components involved in the mechanism at work behind which significant behavioral transformations occurred that the evolution of a CAS. When one steps back to look at the lead human populations in that region to cross one or more bigger picture, each transition between setup and termination Rubicons on the path to modernity. Proposing a best-fit factor represents a single stage within a long-term regional process at play in those passages from the available menu (demog- in which the termination conditions of each stage become raphy, environment, cognitive changes, climate change, spe- the setup conditions of the following stage (fig. 4). This sub- ciation, etc.) often equates to transferring a single-cause sce- sequent stage in turn may not resemble the previous one and nario from a general to a local scale. The same holds true for can potentially fall into another of the 20 possible scenarios more environmentally deterministic approaches seeking to outlined above. When viewed at a regional scale and across reconstruct climatic changes at a regional scale with the aim a long span of time encompassing a number of climatic of identifying a local prime mover behind a behavioral shift phases, this process may reveal trends (e.g., random, punc- identified in that region: contemporaneity does not equate to tuated, unilinear, exponential) in the way CASs respond to a causal link. climate variability. In this way trends may be revealed that We argue that meaningful advances in this field of study would not be evident at the stage level of analysis. We may, cannot be achieved without integrating detailed information for example, face at the stage level minor niche expansion on past human behavior into a research strategy that allows that is not statistically significant but that may become so one to interrogate, rather than simply document, past material when multiple, successive stages are considered. This could culture with the aim of identifying short- and long-term reveal trends that underlie a long-term regional trajectory. We mechanisms at work in the evolution of CASs within their do not expect regional trends to mirror one another. It is respective, dynamic paleoenvironmental frameworks. To ef- possible, however, that by comparing them, consistencies may fectively do so, we must apply the same methods to individual become apparent. The identification of such consistencies regional trajectories and conceive heuristic tools that enable may allow us to move from the analysis of multiple successive us to quantitatively compare and evaluate different regional componential causal systems to the formulation of a general trajectories and their associated behavioral changes through or overarching theory that explains the mechanisms that gov- time. Integration of paleoanthropological and paleogenetic erned the evolution of human cultures and their relationship data can be important but should probably represent a later to the environment before the development of production- stage of the inquiry into mechanisms, as implied by Lalueza- based economies. Fox (2013), rather than being used as a prime mover as is now the case in the more popular single-cause models. Cog- Conclusions nition does not exist in nature as a given but rather as the result of a continuous interaction between conspecifics as well Documenting regional cultural trajectories is worthwhile, but as between them and the environment. Hypotheses on the it is not enough. While the direction that archaeological in- behavioral implications of genomic variability need to be vestigations into behavioral modernity have taken in recent tested by finding ways to explore possible interactions between years has been useful in that it has provided us with a wealth aptitudes and genes rather than attributing to our ancestors of detailed empirical data, this research has so far fallen short an assumed cognitive potential based on our taxonomic read- of examining the dynamic relationships between the multi- ing of the fossil record. tude of factors that were at play between human populations and the environments within which they operated. Focusing on the various paths that human populations around the world have taken to become what we are today certainly Acknowledgments presents the advantage of escaping previous gross approxi- mations or misleading and generalized scenarios accounting We thank Erella Hovers and Steve Kuhn for their invita- for the origin of modernity. A growing body of evidence is tion to participate in the “Alternative Pathways to Complex- demonstrating that such scenarios, founded on single causes, ity” symposium and Laurie Obbink for her valuable assis- do not account for the complex processes at work in each tance. We also wish to thank Gauthier Devilder for his help region of the world. This raises the question of how detailed in creating figures 2–4. Finally, we thank the editors and an compilations of existing data and results from new research anonymous reviewer for their constructive comments. The on individual regions can lead us forward. Such approaches research that led us to the writing of this paper was supported can quickly fall into the trap of cultural relativism: each cul- by the European Research Council (FP7/2007/2013/ERC S384 Current Anthropology Volume 54, Supplement 8, December 2013

TRACSYMBOLS 249587) and the Project Origine II, Aqui- tlement systems and archaeological site formation. American Antiquity 45: 4–20. taine Region. Bocquet-Appel, Jean-Pierre, Pierre-Yves Demars, Lorette Noiret, and Dmitry Dobrowsky. 2005. Estimates of Upper Paleolithic meta-population size in Europe from archaeological data. Journal of Archaeological Science 32:1656– References Cited 1668. Bocquet-Appel, Jean-Pierre, and Alain Tuffreau. 2009. Technological responses Ambrose, Stanley H. 1998. Late Pleistocene human population bottlenecks, of Neanderthals to macroclimatic variations (240,000–40,000 BP). Human volcanic winter, and differentiation of modern humans. Journal of Human Biology 81:287–307. Evolution 34:623–651. Boe¨da, Eric, Jacques Connan, Daniel Dessort, Sultan Muhesen, Norbert Mer- Arau´jo, Miguel B., and Mark New. 2007. Ensemble forecasting of species cier, He´le`ne Valladas, and Nadine Tisne´rat. 1996. Bitumen as a hafting distributions. Trends in Ecology and Evolution 22:42–47. material on Middle Palaeolithic artefacts. Nature 380:336–338. Backwell, Lucinda, and Francesco d’Errico. 2005. The origin of bone tool Bowles, Samuel. 2009. Did warfare among ancestral hunter-gatherers affect technology and the identification of early hominid cultural traditions. In the evolution of human social behaviors? Science 324:1293–1298. From tools to symbols: from early hominids to modern humans. Francesco Brown, Kyle S., Curtis W. Marean, Andy I. R. Herries, Zenobia Jacobs, Chantal d’Errico and Lucinda Backwell, eds. Pp. 238–275. Johannesburg: Witwa- Tribolo, David Braun, David L. Roberts, Michael C. Meyer, and Jocelyn tersrand University Press. Bernatchez. 2009. Fire as an engineering tool of early modern humans. Backwell, Lucinda, Francesco d’Errico, and Lyn Wadley. 2008. Middle Stone Science 325:859–862. Age bone tools from the Howiesons Poort layers, Sibudu Cave, South Africa. Caˆrciumaru, Marin, Rodica-Mariana Ion, Elena-Cristina Nit¸u, and Radu S¸te- Journal of Archaeological Science 35:1566–1580. fa˘nescu. 2012. New evidence of adhesive as hafting material on Middle and Banks, William E., Nicolas Antunes, Solange Rigaud, and Francesco d’Errico. Upper Paleolithic artefacts from Gura Cheii-Raˆs¸nov Cave (Romania). Jour- 2013. Ecological constraints on the first prehistoric farmers in Europe. nal of Archaeological Science 39:1942–1950. Journal of Archaeological Science 40:2746–2753. Caron, Franc¸ois, Francesco d’Errico, Pierre Del Moral, Fre´de´ric Santos, and Banks, William E., Thierry Aubry, Francesco d’Errico, Joa˜o Zilha˜o, Andre´s Joa˜o Zilha˜o. 2011. The reality of Neanderthal symbolic behavior at the Lira-Noriega, and A. Townsend Peterson. 2011. Eco-cultural niches of the Grotte du Renne, Arcy-sur-Cure, France. PLoS ONE 6(6):e21545. Badegoulian: unraveling links between cultural adaptation and ecology dur- Carto, Shannon L., Andrew J. Weaver, Rene´e Hetherington, Yin Lam, and ing the Last Glacial Maximum in France. Journal of Anthropological Ar- Edward C. Wiebe. 2009. Out of Africa and into an ice age: on the role of chaeology 30:359–374. global climate change in the late Pleistocene migration of early modern Banks, William E., Francesco d’Errico, A. TownsendPeterson, Masa Kageyama, humans out of Africa. Journal of Human Evolution 56:139–151. Adriana Sima, and Maria Sa´nchez-Gon˜i. 2008a. Neanderthal extinction by Charrie´-Duhaut, Armelle, Guillaume Porraz, Caroline R. Cartwright, Marina competitive exclusion. PLoS ONE 3(12):e3972. Igreja, Jacques Connan, Cedric Poggenpoel, and Pirre-Jean Texier. 2013. Banks, William E., Francesco d’Errico, A. Townsend Peterson, Marian Van- First molecular identification of a hafting adhesive in the Late Howiesons haeren, Masa Kageyama, Pierre Sepulchre, Gilles Ramstein, Anne Jost, and Poort at Diepkloof Rock Shelter (Western Cape, South Africa). Journal of Daniel Lunt. 2008b. Human ecological niches and ranges during the LGM Archaeological Science 40:3506–3518. in Europe derived from an application of eco-cultural niche modeling. Childe, V. Gordon. 1929. The Danube in prehistory. Oxford: Oxford University Journal of Archaeological Science 35:481–491. Press. Banks, William E., Francesco d’Errico, and Joa˜o Zilha˜o. 2013. Human-climate Clarke, David L. 1968. Analytical archaeology. London: Methuen. interaction during the Early Upper Paleolithic: testing the hypothesis of an Compton, John S. 2011. Pleistocene sea-level fluctuations and human evo- adaptive shift between the Proto-Aurignacian and the Early Aurignacian. lution on the southern coastal plain of South Africa. Quaternary Science Journal of Human Evolution 64:39–55. Reviews 30:506–527. Banks, William E., Joa˜o Zilha˜o, Francesco d’Errico, Masa Kageyama, Adriana Conard, Nicholas J. 2008. A critical view of the evidence for a southern African Sima, and Annamaria Ronchitelli. 2009. Investigating links between ecology origin of behavioural modernity. South African Archaeological Society Good- and bifacial tool types in Western Europe during the Last Glacial Maximum. win Series 10:175–179. Journal of Archaeological Science 36:2853–2867. Deacon, Janette. 1990. Weaving the fabric of Stone Age research in southern Barham, Lawrence S. 2001. Central Africa and the emergence of regional Africa. In A history of African archaeology. Peter Robertshaw, ed. Pp. 39– identity in the Middle Pleistocene. In Human roots: Africa and Asia in the 58. London: James Currey. Middle Pleistocene. Lawrence S. Barham and Kate Robson-Brown, eds. Pp. Delagnes, Anne, and Liliane Meignen. 2006. Diversity of lithic production 65–80. Bristol: Western Academic and Specialist Press. systems during the Middle Paleolithic in France. In Transitions before the ———. 2007. Modern is as modern does? technological trends and thresholds transition: evolution and stability in the Middle Paleolithic and Middle Stone in the south-central African record. In Rethinking the human revolution. Age. Erella Hovers and Steven L. Kuhn, eds. Pp. 85–107. New York: Springer. Paul Mellars, Katie Boyle, Ofer Bar-Yosef, and Chris Stringer, eds. Pp. 165– deMenocal, Peter B. 2011. Climate and human evolution. Science 331:540– 176. McDonald Institute Monographs. Cambridge: McDonald Institute for 542. Archaeological Research. d’Errico, Francesco. 2003. The invisible frontier: a multiple species model for Barton, C. Michael, Julien Riel-Salvatore, John M. Anderies, and Gabriel the origin of behavioral modernity. Evolutionary Anthropology 12:188–202. Popescu. 2011. Modeling human ecodynamics and biocultural interactions ———. 2009. Le rouge et le noir: implications of early pigment use in Africa, in the Late Pleistocene of western Eurasia. Human Ecology 39:705–725. the Near East, and Europe for the origin of cultural modernity. South African Barve, Narayani, Vijay Barve, Alberto Jime´nez-Valverde, Andre´s Lira-Noriega, Archaeological Bulletin 10:157–174. Sean P. Maher, A. Townsend Peterson, Jorge Sobero´n, and Fabricio Villa- d’Errico, Francesco, Lucinda Backwell, Paola Villa, Ilaria Degano, Jeannette lobos. 2011. The crucial role of the accessible area in ecological niche mod- J. Lucejko, Marion K. Bamford, Thomas F. G. Higham, Maria Perla Col- eling and species distribution modeling. Ecological Modelling 222:1810– ombini, and Peter B. Beaumont. 2012. Artifacts from Border Cave: earliest 1819. evidence of San material culture. Proceedings of the National Academy of Bar-Yosef, Ofer, and Anna Belfer-Cohen. 2011. Following Pleistocene road Sciences of the USA 109:13214–13219. signs of human dispersals across Eurasia. Quaternary International 285:30– d’Errico, Francesco, Lucinda Backwell, and Lyn Wadley. 2012. Indentifying 43. regional variability in Middle Stone Age bone technology: the case of Sibudu Bechtel, William, and Adele Abrahamsen. 2005. Explanation: a mechanistic Cave. Journal of Archaeological Science 39:2479–2495. alternative. Studies in History and Philosophy of Biological and Biomedical d’Errico, Francesco, Valentina Borgia, and Annamaria Ronchitelli. 2012. Uluz- Sciences 36:421–441. zian bone technology and its implications for the origin of behavioural Belfer-Cohen, Anna, and Erella Hovers. 2010. Modernity, enhanced working modernity. Quaternary International 259:59–71. memory, and the Middle to Upper Paleolithic record in the Levant. Current d’Errico, Francesco, and Christopher S. Henshilwood. 2007. Additional evi- Anthropology 51(suppl. 1):S167–S175. dence for bone technology in the southern African Middle Stone Age. Bickerton, Derek. 1990. Language and species. Chicago: University of Chicago Journal of Human Evolution 52:142–163. Press. ———. 2011. The origin of symbolically mediated behavior. In Homo sym- Binford, Lewis R. 1980. Willow smoke and dogs’ tails: hunter-gatherer set- bolicus: the dawn of language, imagination, and spirituality. Christopher S. d’Errico and Banks Mechanisms behind Cultural Trajectories S385

Henshilwood and Francesco d’Errico, eds. Pp. 49–73, Amsterdam: John J. M. Macpherson, Jeffrey M. Kidd, Laura Rodrı´guez-Botigue´, et al. 2011. Benjamins. Hunter-gatherer genomic diversity suggests a southern African origin for d’Errico, Francesco, Renata Garcı´a Moreno, and Riaan Rifkin. 2012. Tech- modern humans. Proceedings of the National Academy of Sciences of the USA nological, elemental and colorimetric analysis of an engraved ochre frag- 108:5154–5162. ment from the Middle Stone Age levels of Klasies River Cave 1, South Henrich, Joseph. 2004. Demography and cultural evolution: how adaptive Africa. Journal of Archaeological Science 39:942–952. cultural processes can produce maladaptive losses: the Tasmanian case. d’Errico, Francesco, He´le`ne Salomon, Colette Vignaud, and Chris Stringer. American Antiquity 69:197–214. 2010. Pigments from the Middle Palaeolithic levels of Es-Skhul (Mount Henrich, Joseph, Steven J. Heine, and Ara Norenzayan. 2010. The weirdest Carmel, Israel). Journal of Archaeological Science 37:3099–3110. people in the world? Behavioral and Brain Sciences 33:61–135. d’Errico, Francesco, and Maria Fernanda Sa´nchez Gon˜i. 2003. Neanderthal Henshilwood, Christopher S., Francesco d’Errico, Karen L. van Niekerk, Yvan extinction and the millennial scale climatic variability of the OIS 3. Qua- Coquinot, Zenobia Jacobs, Stein-Erik Lauritzen, Michel Menu, and Renata ternary Science Reviews 22:769–788. Garcı´a-Moreno. 2011. A 100,000-year-old ochre-processing workshop at d’Errico, Francesco, and Chris B. Stringer. 2011. Evolution, revolution or Blombos Cave, South Africa. Science 334:219–222. saltation scenario for the emergence of modern cultures. Philosophical Henshilwood, Christopher S., Francesco d’Errico, and Ian Watts. 2009. En- Transactions of the Royal Society B 366:1060–1069. graved ochres from the Middle Stone Age levels at Blombos Cave, South d’Errico, Francesco, and Marian Vanhaeren. 2007. Evolution or revolution? Africa. Journal of Human Evolution 57:27–47. new evidence for the origin of symbolic behavior in and out of Africa. In Henshilwood, Christopher S., and Curtis W. Marean. 2003. The origin of Rethinking the human revolution. Paul Mellars, Katie Boyle, Ofer Bar-Yosef, modern human behaviour: a review and critique of models and test im- and Chris Stringer, eds. Pp. 275–286. McDonald Institute Monographs. plications. Current Anthropology 44:627–665. Cambridge: McDonald Institute for Archaeological Research. Hodder, Ian. 1991. Reading the past: current approaches to interpretation in d’Errico, Francesco, Marian Vanhaeren, Nick Barton, Abdeljalil Bouzouggar, archaeology. 2nd edition. Cambridge: Cambridge University Press. Henk Mienis, Daniel Richter, Jean-Jacques Hublin, Shannon P. McPherron, Holling C. S. 1973. Resilience and stability of ecological systems. Annual and Pierre Lozouet. 2009. Additional evidence on the use of personal or- Review of Ecology and Systematics 4:1–23. naments in the Middle Paleolithic of North Africa. Proceedings of the Na- Hovers, Erella. 2009. The Middle-to-Upper Paleolithic transition: what news? tional Academy of Sciences of the USA 106:16051–16056. In Sourcebook of Paleolithic transitions: methods, theories, and interpretations. Discamps, Emmanuel, Jacques Jaubert, and Franc¸ois Bachellerie. 2011. Hu- Marta Camps and Parth Chauhan, eds. Pp. 455–462. New York: Springer. man choices and environmental constraints: deciphering the variability of Hovers, Erella, and Anna Belfer-Cohen. 2006. “Now you see it, now you large game procurement from Mousterian to Aurignacian times (MIS 5– don’t”: modern human behavior in the Middle Paleolithic. In Transitions 3) in southwestern France. Quaternary Science Reviews 30:2755–2775. before the transition: evolution and stability in the Middle Paleolithic and Dufresne, Jean-Louis, Marie-Alice Foujols, Se´bastien Denvil, Arnaud Caubel, Middle Stone Age. Erella Hovers and Steven L. Kuhn, eds. Pp. 295–304. Olivier Marti, Yves Balkanski, Slimane Bekki, et al. 2013. Climate change New York: Springer. projections using the IPSL-CM5 earth system model: from CMIP3 to Jacobs, Zenobia, Richard G. Roberts, Rex F. Galbraith, Hilary J. Deacon, Rainer CMIP5. Climate Dynamics 40:2123–2165. Gru¨n, Alex Mackay, Peter Mitchell, Ralf Vogelsang, and Lyn Wadley. 2008. Dunbar, Robin I. M. 2003. The social brain: mind, language, and society in Ages for the Middle Stone Age of southern Africa: implications for human evolutionary perspective. Annual Review of Anthropology 32:163–181. behavior and dispersal. Science 322:733–735. Fabre, Virginie, Silvana Condemi, Anna Degioanni, and Estelle Herrscher. Johnson, Allen W., and Timothy Earle. 2000. The evolution of human societies: 2011. Neanderthal versus modern humans: evidence for resource compe- from foraging group to agrarian state. 2nd edition. Stanford, CA: Stanford tition from isotopic modelling. International Journal of Evolutionary Biology University Press. 2011, doi:10.4061/2011/689315. Kageyama, Masa, Pascale Braconnot, Laurent Bopp, Ve´ronique Mariotti, Tilla Foley, Robert L. 1992. Mobility/sedentism: concepts, archaeological measures, Roy, Marie-Noe¨lle Woillez, Arnaud Caubel, et al. 2013. Mid-Holocene and and effects. Annual Review of Anthropology 21:43–66. last glacial maximum climate simulations with the IPSL model. 2. Model- Frei, Christoph, Regina Scho¨ll, Sophie Fukutome, Ju¨rg Schmidli, and Pier L. data comparisons. Climate Dynamics 40:2469–2495. Vidale. 2006. Future change of precipitation extremes in Europe: intercom- Kageyama, Masa, Nathalie Combourieu Nebout, Pierre Sepulchre, Odile Pey- parison of scenarios from regional climate models. Journal of Geophysical ron, Gerhard Krinner, Gilles Ramstein, and Jean-Pierre Cazet. 2005. The Research 111, doi:10.1029/2005JD005965. Last Glacial Maximum and Heinrich Event 1 in terms of climate and veg- Gamble, Clive, William Davies, Paul Pettitt, and Martin Richards. 2004. Cli- etation around the Alboran Sea: a preliminary model-data comparison. mate change and evolving human diversity in Europe during the last glacial. Comptes Rendus Geoscience 337:983–992. Philosophical Transactions of the Royal Society B 359:243–254. Kandel, Andrew W., and Nicholas J. Conard. 2005. Production sequences of Gelfand, Michele J., Jana L. Raver, Lisa Nishii, Lisa M. Leslie, Janetta Lun, ostrich eggshell beads and settlement dynamics in the Geelbek Dunes of Beng Chong Lim, Lili Duan, Assaf Almaliach, Soon Ang, and Jakobina the Western Cape, South Africa. Journal of Archaeological Science 32:1711– Arnadottir. 2011. Differences between tight and loose cultures: a 33-nation 1721. study. Science 332:1100–1104. Kang, Sarah M., Isaac M. Held, Dargan M. W. Frierson, and Ming Zhao. Gibson, Kathleen R. 2007. Putting it all together: a constructionist approach 2008. The response of the ITCZ to extratropical thermal forcing: idealized to the evolution of human mental capabilities. In Rethinking the human slab-ocean experiments with a GCM. Journal of Climate 21:3521–3532. revolution. Paul Mellars, Katie Boyle, Ofer Bar-Yosef, and Chris Stringer, Klein, Richard G. 2000. Archaeology and the evolution of human behavior. eds. Pp. 67–77. McDonald Institute Monographs. Cambridge: McDonald Evolutionary Anthropology 9:17–36. Institute for Archaeological Research. ———. 2009. The human career: human biological and cultural origins. Chi- Green, Richard E., Johannes Krause, Adrian W. Briggs, Tomislav Maricic, Udo cago: University of Chicago Press. Stenzel, Martin Kircher, Nick Patterson, et al. 2010. A draft sequence of Kuhn, Steven L. 2013. Roots of the Middle Paleolithic in Eurasia. Current the Neandertal genome. Science 328:710–722. Anthropology 54(suppl. 8):S255–S268. Gru¨n, Rainer, Chris Stringer, Frank McDermott, Roger Nathan, Naomi Porat, Kuhn, Steven L., and Mary C. Stiner. 2007. Paleolithic ornaments: implications Steve Robertson, Lois Taylor, Graham Mortimer, Stephen Eggins, and Mal- for cognition, demography and identity. Diogenes 54:40–48. colm McCulloch. 2005. U-series and ESR analyses of bones and teeth re- Kuorikoski, Jaakko. 2009. Two concepts of mechanism: componential causal lating to the human burials from Skhul. Journal of Human Evolution 49: system and abstract form of interaction. International Studies in the Phi- 316–334. losophy of Science 23:143–160. Hauck, Thomas C., Jacques Connan, Armelle Charrie´-Duhaut, Jean-Marie Le Lalueza-Fox, Carles. 2013. Agreements and misunderstandings among three Tensorer, and Heba al Sakhel. 2013. Molecular evidence of bitumen in the scientific fields: paleogenomics, archaeology, and human paleontology. Cur- Mousterian lithic assemblage of Hummal (Central Syria). Journal of Ar- rent Anthropology 54(suppl. 8):S214–S220. chaeological Science 40:3252–3262. Lambeck, Kurt, Tezer M. Esat, and Emma-Kate Potter. 2002. Links between Hayden, Brian. 1979. Palaeolithic reflections: lithic technology and ethnographic climate and sea levels for the past three million years. Nature 419:199–206. excavation among Australian Aborigines. Englewood Cliffs, NJ: Humanities Langbroek, Marco. 2011. Trees and ladders: a critique of the theory of human Press. cognitive and behavioural evolution in Palaeolithic archaeology. Quaternary Henn, Brenna M., Christopher R. Gignoux, Matthew Jobin, Julie M. Granka, International 270:4–14. S386 Current Anthropology Volume 54, Supplement 8, December 2013

Langley, Michelle C., Christopher Clarkson, and Sean Ulm. 2008. Behavioral P. Dawson, and David C. Lees. 2006. Model-based uncertainty in species complexity in Eurasian Neanderthal populations: a chronological exami- range prediction. Journal of Biogeography 33:1704–1711. nation of the archaeological record. Cambridge Archaeological Journal 18: Peeples, Matthew A., C. Michael Barton, and Steven Schmich. 2006. Resilience 289–307. lost: intersecting land use and landscape dynamics in the prehistoric south- Lemonnier, Pierre. 1986. The study of material culture today: toward an western United States. Ecology and Society 11(2), article 22. http:// anthropology of technical systems. Journal of Anthropological Archaeology www.ecologyandsociety.org/vol11/iss2/art22. 5:147–186. Peresani, Marco, Ivana Fiore, Monica Gala, Matteo Romandini, and Antonio Leroi-Gourhan, Andre´. 1964. Le geste et la parole I: technique et langage. Paris: Tagliacozzo. 2011. Late Neandertals and the intentional removal of feathers Albin Michel. as evidenced from bird bone taphonomy at Fumane Cave 44 ky B.P., Italy. Lombard, Marlize. 2005. Evidence for hunting and hafting during the Middle Proceedings of the National Academy of Sciences of the USA 108:3888–3893. Stone Age at Sibudu Cave, KwaZulu-Natal, South Africa. Journal of Human Peterson, A. Townsend. 2003. Predicting the geography of species’ invasions Evolution 48:279–300. via ecological niche modeling. Quarterly Review of Biology 78:419–433. ———. 2012. Thinking through the Middle Stone Age of sub-Saharan Africa. Peterson, A. Townsend, Monica Papes¸, and Jorge Sobero´n. 2008. Rethinking Quaternary International 270:140–155. receiver operating characteristic analysis applications in ecological niche Lowe, John, Nick Barton, Simon Blockley, Christopher Bronk Ramsey, Victoria modeling. Ecological Modelling 213:63–72. L. Cullen, William Davis, Clive Gamble, et al. 2012. Volcanic ash layers Peterson, A. Townsend, Jorge Sobero´n, Richard G. Pearson, Robert P. An- illuminate the resilience of Neanderthals and early modern humans to derson, Enrique Martı´nez-Meyer, Miguel Nakamua, and Miguel B. Arau´jo. natural hazards. Proceedings of the National Academy of Sciences of the USA 2011. Ecological niches and geographic distributions. Monographs in Popu- 109:13532–13537. lation Biology, no. 49. Princeton, NJ: Princeton University Press. Machamer, Peter, Lindley Darden, and Carl F. Craver. 2000. Thinking about Pettitt, Paul B. 2011. The Palaeolithic origins of human burial. London: Rout- mechanisms. Philosophy of Science 67:1–25. ledge. Mackay, Alex, and Aara Welz. 2008. Engraved ochre from a Middle Stone Age Powell, Adam, Stephen Shennan, and Mark G. Thomas. 2009. Late Pleistocene context at Klein Kliphuis in the Western Cape of South Africa. Journal of demography and the appearance of modern human behavior. Science 324: Archaeological Science 35:1521–1532. 1298–1301. Marean, Curtis W., Miryam Bar-Matthews, Jocelyn Bernatchez, Erich Fisher, Reich, David, Richard E. Green, Martin Kircher, Johannes Krause, Nick Pat- Paul Goldberg, Andy I. R. Herries, Zenobia Jacobs, et al. 2007. Early human terson, Eric Y. Durand, Bence Viola, et al. 2010. Genetic history of an use of marine resources and pigment in South Africa during the Middle archaic hominin group from Denisova Cave in Siberia. Nature 468:1053– Pleistocene. Nature 449:905–908. 1060. Maslin, Mark A., and Beth Christensen. 2007. Tectonics, orbital forcing, global Renfrew, A. Colin. 1977. Space, time and polity. In The Evolution of social climate change, and human evolution in Africa: introduction to the African systems. John Friedman and Michael J. Rowlands, eds. Pp. 89–114. London: paleoclimate special volume. Journal of Human Evolution 53:443–464. Duckworth. Mazza, Paul Peter Anthony, Fabio Martini, Benedetto Sala, Maurizio Magi, Richerson, Peter J., Robert Boyd, and Robert L. Bettinger. 2009. Cultural Maria Perla Colombini, Gianna Giachi, Francesco Landucci, Cristina Le- innovations and demographic change. Human Biology 81:211–235. morini, Francesca Modugno, and Erika Ribechini. 2006. A new Palaeolithic Roebroeks, Wil, Mark J. Sier, Trine Kellberg Nielsen, Dimitri De Loecker, discovery: tar-hafted stone tools in a European Mid-Pleistocene bone-bear- Josep Maria Pare´s, Charles E. S. Arps, and Herman J. Mu¨cher. 2012. Use ing bed. Journal of Archaeological Science 33:1310–1318. of red ochre by early Neandertals. Proceedings of the National Academy of McBrearty, Sally, and Alison S. Brooks. 2000. The revolution that wasn’t: a Sciences of the USA 109:1889–1894. new interpretation of the origin of modern human behavior. Journal of Schlanger, Nathan. 1994. Mindful technology: unleashing the chaıˆne ope´ratoire Human Evolution 39:453–563. for an archaeology of mind. In The ancient mind: elements of cognitive Mellars, Paul. 1989. Major issues in the origin of modern humans. Current archaeology. Colin Renfrew and Ezra B. W. Zubrow, eds. Pp. 143–151. Anthropology 30:349–385. Cambridge: Cambridge University Press. ———. 2006. Why did modern human populations disperse from Africa ca. Schoon, Michael, Christo Fabricius, John M. Anderies, and Margaret Nelson. 60,000 years ago? a new model. Proceedings of the National Academy of 2011. Synthesis: vulnerability, traps, and transformations: long-term per- Sciences of the USA 103:9381–9386. spectives from archaeology. Ecology and Society 16(2), article 24. http:// Mellars, Paul, and Chris Stringer. 1989. The human revolution: behavioural www.ecologyandsociety.org/vol16/iss2/art24. and biological perspectives on the origins of modern humans. Princeton, NJ: Sepulchre, Pierre, Gilles Ramstein, Masa Kageyama, Marian Vanhaeren, Ger- Princeton University Press. hard Krinner, Marı´a-Fernanda Sa´nchez-Gon˜i, and Francesco d’Errico. 2007. Mercader, Julio. 2009. Mozambican grass seed consumption during the Middle H4 abrupt event and late Neanderthal presence in Iberia. Earth and Plan- Stone Age. Science 326:1680–1683. etary Science Letters 258:283–292. Mesoudi, Alex, Andrew Whiten, and Kevin N. Laland. 2006. Towards a unified Shea, John J. 2011. Homo sapiens is as Homo sapiens was. Current Anthropology science of cultural evolution. Behavioral Brain Sciences 29:329–383. 52:1–35. Meyer, M., M. Kircher, M.-T. Gansauge, H. Li, F. Racimo, S. Mallick, J. G. Shennan, Stephen. 2001. Demography and cultural innovation: a model and Schraiber, et al. 2012. A high-coverage genome sequence from an archaic its implications for the emergence of modern human culture. Cambridge Denisovan individual. Science: doi:10.1126/science.1224344. Archaeological Journal 11:5–16. Mourre, Vincent, Paola Villa, and Christopher S. Henshilwood. 2010. Early Sobero´n, Jorge, and A. Townsend Peterson. 2005. Interpretation of models of use of pressure flaking on lithic artifacts at Blombos Cave, South Africa. fundamental ecological niches and species’ distributional areas. Biodiversity Science 330:659–662. Informatics 2:1–10. Norenzayan, Ara. 2011. Explaining human behavioral diversity. Science 332: Soriano, Sylvain, Paola Villa, and Lyn Wadley. 2007. Blade technology and 1041–1042. tool forms in the Middle Stone Age of South Africa: the Howiesons Poort Nowell, April. 2010. Defining behavioral modernity in the context of Nean- and post-Howiesons Poort at Rose Cottage Cave. Journal of Archaeological dertal and anatomically modern human populations. Annual Review of Science 34:681–703. Anthropology 39:437–452. Sterelny, Kim. 2011. From hominins to humans: how sapiens became behav- Odum, Eugene. 1971. Fundamentals of ecology. 3rd edition. Philadelphia: Saun- iourally modern. Philosophical Transactions of the Royal Society B 366:809– ders. 822. Osborne, Anne H., Derek Vance, Eelco J. Rohling, Nick Barton, Mike Rog- Steward, Julian. 1955. Theory of culture change. Urbana: University of Illinois erson, and Nuri Fello. 2008. A humid corridor across the Sahara for the Press. migration of early modern humans out of Africa 120,000 years ago. Pro- Stiner, Mary C. 2013. An unshakable Middle Paleolithic? trends versus con- ceedings of the National Academy of Sciences of the USA 105:16444–16447. servatism in the predatory niche and their social ramifications. Current Pawlik, Alfred F., Ju¨rgen P. Thissen. 2011. Hafted armatures and multi-com- Anthropology 54(suppl. 8):S288–S304. ponent tool design at the Micoquian site of Inden-Altdorf, Germany. Journal Stiner, Mary C., Natalie D. Munro, and Todd A. Surovell. 2000. The tortoise of Archaeological Science 38:1699–1708. and the hare: small-game use, the broad-spectrum revolution, and Paleo- Pearson, Richard G., Wilfried Thuiller, Miguel B. Arau´jo, Enrique Martinez- lithic demography. Current Anthropology 41:39–73. Meyer, Lluı´s Brotons, Colin McClean, Lera Miles, Pedro Segurado, Terence Stockwell, David, and David Peters. 1999. The GARP modelling system: prob- d’Errico and Banks Mechanisms behind Cultural Trajectories S387

lems and solutions to automated spatial prediction. International Journal ———. 2010b. A taphonomic study of ochre demonstrates post-depositional of Geographic Information Systems 13:143–158. color transformations. Journal of Taphonomy 8:243–254. Texier, Pierre-Jean, Guillaume Porraz, John Parkington, Jean-Philippe Rigaud, ———. 2010c. Were snares used in the Middle Stone age and does it matter? Cedric Poggenpoel, Christopher Miller, Chantal Tribolo, et al. 2010. A How- a review and case study from Sibudu, South Africa. Journal of Human iesons Poort tradition of engraving ostrich eggshell containers dated to Evolution 58:179–192. 60,000 years ago at Diepkloof Rock Shelter, South Africa. Proceedings of the Wadley, Lyn, Tamaryn Hodgskiss, and Michael Grant. 2009. Implications for National Academy of Sciences of the USA 107:6180–6185. complex cognition from the hafting of tools with compound adhesives in Tingley, Martin P., Peter F. Craigmile, Murali Haran, Bo Li, Elizabeth Mann- the Middle Stone Age, South Africa. Proceedings of the National Academy shardt, and Bala Rajaratnam. 2012. Piecing together the past: statistical of Sciences of the USA 106:9590–9594. insights into paleoclimatic reconstructions. Quaternary Science Reviews 35: Wadley, Lyn, Christine Sievers, Marion Bamford, Paul Goldberg, Francesco 1–22. Berna, and Christopher Miller. 2011. Middle Stone Age bedding construc- Tomasello, Michael. 1994. The question of chimpanzee culture. In Chimpanzee tion and settlement patterns at Sibudu, South Africa. Science 334:1388– cultures. Richard W. Wrangham, W. C. McGrew, Frans B. M. de Waal, and 1391. Paul Heltne, eds. Pp. 301–317. Cambridge: Harvard University Press. Walker, Brian, C. S. Holling, Stephen R. Carpenter, and Ann Kinzig. 2004. Trinkaus, Erik. 2005. Early modern humans. Annual Review of Anthropology Resilience, adaptability and transformability in social-ecological systems. 34:207–230. Ecology and Society 9(2), article 5. http://www.ecologyandsociety.org/vol9 Van Andel, Tjeerd H., and William Davies, eds. 2003. Neanderthals and modern /iss2/art5. humans in the European landscape during the last glaciation: archaeological Warren, Dan L., Richard E. Glor, and Michael Turelli. 2010. ENMTools: a results of the stage 3 project. Cambridge: McDonald Institute for Archaeo- toolbox for comparative studies of environmental niche models. Ecography logical Research. 33:607–611. Vanhaeren, Marian, Francesco d’Errico, Chris Stringer, Sarah L. James, Jon- Watts, Ian. 2010. The pigments from Pinnacle Point Cave 13B, Western Cape, athan Todd, and Henk K. Mienis. 2006. Middle Paleolithic shell beads in South Africa. Journal of Human Evolution 59:392–411. Israel and Algeria. Science 312:1785–1788. Weaver, Timothy D., and Charles C. Roseman. 2008. New developments in Vanhaeren, Marian, Francesco d’Errico, Karen L. van Niekerk, Christopher S. the genetic evidence for modern human origins. Evolutionary Anthropology Henshilwood, and Rudolph M. Erasmus. 2013. Thinking strings: additional 17:69–80. evidence for personal ornament use in the Middle Stone Age at Blombos Wurz, Sara, N. J. le Roux, S. Gardner, and Hilary J. Deacon. 2003. Discrim- Cave, South Africa. Journal of Human Evolution 64:500–517. inating between the end products of the earlier Middle Stone Age sub- Villa, Paola, Marie Soressi, Christopher S. Henshilwood, and Vincent Mourre. stages at Klasies River using biplot methodology. Journal of Archaeological 2009. The Still Bay points of Blombos Cave (South Africa). Journal of Science 30:1107–1126. Archaeological Science 36:441–460. Wynn, Thomas, and Frederick L. Coolidge. 2010. Beyond symbolism and Villa, Paola, and Sylvain Soriano. 2010. Hunting weapons of Neanderthals language. Current Anthropology 51(suppl. 1):S5–S16. and early modern humans in South Africa. Journal of Anthropological Re- Zilha˜o, Joa˜o. 2001. Anatomically archaic, behaviorally modern: the last Nean- search 66:5–38. derthals and their destiny. Amsterdam: Stichting Nederlands Museum voor Villa, Paola, Sylvain Soriano, Nicolas Teyssandier, and Sara Wurz. 2010. The Anthropologie en Praehistoriae. Howiesons Poort and MSA III at Klasies River main site, Cave 1A. Journal ———. 2006. Neandertals and moderns mixed, and it matters. Evolutionary of Archaeological Science 37:630–655. Anthropology 15:183–195. Villa, Paola, Sylvain Soriano, Tsenka Tsanova, Ilaria Degano, Thomas F. G. ———. 2007. The emergence of ornaments and art: an archaeological per- Higham, Francesco d’Errico, Lucinda Backwell, Jeannette J. Luceiko, Maria spective on the origins of “behavioral modernity.” Journal of Archaeological Perla Colombini, and Peter B. Beaumont. 2012. Border Cave and the be- Research 15:1–54. ginning of the Later Stone Age in South Africa. Proceedings of the National ———. 2011. The emergency of language, art, and symbolic thinking. In Academy of Sciences of the USA 109:13208–13213. Homo symbolicus: the dawn of language, imagination and spirituality. Chris- Vrac, Mathieu, Michael Stein, and Katharine Hayhoe. 2007. Statistical down- topher S. Henshilwood and Francesco d’Errico, eds. Pp. 111–131. Amster- scaling of precipitation through nonhomogeneous stochastic weather typ- dam: John Benjamins. ing. Climate Research 34:169–184. Zilha˜o, Joa˜o, Diego E. Angelucci, Ernestina Badal-Garcı´a, Francesco d’Errico, Wadley, Lyn. 2010a. Compound-adhesive manufacture as a behavioral proxy Flore´al Daniel, Laure Dayet, Katerina Douka, et al. 2010. Symbolic use of for complex cognition in the Middle Stone Age. Current Anthropology marine shells and mineral pigments by Iberian Neandertals. Proceedings of 51(suppl. 1):S111–S119. the National Academy of Sciences of the USA 107:1023–1028. S388 Current Anthropology Volume 54, Supplement 8, December 2013

Population Size as an Explanation for Patterns in the Paleolithic Archaeological Record More Caution Is Needed

by Mark Collard, Briggs Buchanan, and Michael J. O’Brien

Recently it has become commonplace to use population size to explain patterns in the Paleolithic archaeological record. Several modeling studies support the idea that population size can affect cultural evolution, but the results of empirical studies are ambiguous. Here we report a study that used tool kit data from recent hunter-gatherers, in conjunction with correlation analysis and a global sample, a continental sample, and a regional sample. The results of the analyses do not support the hypothesis. Population size was correlated with some tool kit variables in the global sample, but these relationships disappeared when two factors that have previously been found to affect hunter-gatherer tool kits—risk of resource failure and mobility—were controlled for. Population size was not correlated with the tool kit variables in the other samples. The regression analyses also did not support the population size hypothesis. Together, these results challenge the use of population size to explain patterns in the Paleolithic archaeological record. Population size may explain some of the patterns in question, but this needs to be demonstrated through tests in which the population size hypothesis is explicitly pitted against competing hypotheses, such as adaptation to shifting ecological conditions.

Introduction novations seem to have appeared, disappeared, and then reappeared during the late Pleistocene. Premo and Kuhn (2010) have argued that two key features of the Middle Pa- Recently a number of researchers have argued that population leolithic and Middle Stone Age archaeological records—an size might explain several long-debated patterns in the Pa- absence of directional technological change and the reap- leolithic archaeological record. Shennan (2001), for example, pearance of previously existing cultural behaviors—might be has suggested that the so-called creative explosion of the late a function of a high rate of extirpation of small, isolated Middle Stone Age and Upper Paleolithic might have resulted groups and subsequent repopulation. from a large, climate-driven increase in population size. Sim- Support for these hypotheses comes from a number of ilarly, Riede (2008) has argued that the emergence of the formal models that suggest population size can have a sig- Bromme and Perstunian technocomplexes in Northern Eu- nificant effect on the evolution of fitness-relevant cultural rope during the Late Glacial period was driven by population traits. The earliest of these models was described by Shennan size reduction associated with the Laacher See eruption. Pow- (2001). Shennan modified a population-genetics model to ell, Shennan, and Thomas (2009) have proposed that pop- incorporate social learning among individuals and then car- ulation increase might also explain why many cultural in- ried out a series of simulation trials. He found that larger populations have a major advantage over smaller ones when Mark Collard is Canada Research Chair and Professor, and Briggs it comes to adaptive cultural innovation because of the de- Buchanan is Postdoctoral Fellow, at the Human Evolutionary Studies creasing role of sampling effects as populations grow. His Program and Department of Archaeology of Simon Fraser University results suggested that when effective population size is large, (8888 University Drive, Burnaby, British Columbia V5A 1S6, Canada there is a far greater probability of fitness-enhancing cultural [[email protected]]). Michael J. O’Brien is Dean of the College of innovations being maintained and deleterious ones being lost Arts and Science, Professor of Anthropology, and Director of the than when effective populations are small. In the latter sit- Museum of Anthropology in the Department of Anthropology of the University of Missouri (107 Swallow Hall, Columbia, Missouri uation, innovations that are maintained tend to be less ben- 65211-1440, U.S.A.). This paper was submitted 3 VII 13, accepted 4 eficial in terms of reproduction and also less attractive for IX 13, and electronically published 20 XII 13. imitators.

᭧ 2013 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved. 0011-3204/2013/54S8-0016$10.00. DOI: 10.1086/673881 Collard, Buchanan, and O’Brien Population Size and the Paleolithic Archaeological Record S389

Other models that demonstrate that population size can hypothesis (Collard, Kemery, and Banks 2005; Collard et al. affect the evolution of fitness-relevant traits have been re- 2013a, 2013b; Kline and Boyd 2010; Neiman 1995; Nelson et ported by Henrich (2004); Powell, Shennan, and Thomas al. 2011). Some studies support it. Neiman (1995) investigated (2009); Mesoudi (2011); and Kobayashi and Aoki (2012). the amount of variation to be expected in the decoration of Henrich (2004) argued that population size can affect the a pottery assemblage if the motifs are neutral in terms of probability of more complex skills being invented and main- adaptation. He then analyzed rim decoration variation among tained. In his model, learners preferentially copy the most seven successive phases of the Woodland period in Illinois, skilled practitioner in their population with some amount of United States, and found that it matched the expectations of error. The probability distribution that determines the his model. He concluded that the patterns of variation de- amount of error is such that a learner will only occasionally pended on changing levels of intergroup contact, which arrive at a behavior that gives a better result than the previous started low, increased, and then declined again. Kline and best. The likelihood of this occurring is dependent on pop- Boyd (2010) examined the effect of population size on marine ulation size because in large populations even improbable foraging tool kits of 10 recent nonindustrial farming popu- events occur now and again, and the larger the population, lations from Oceania and found that population size had a the more likely this is. Powell, Shennan, and Thomas (2009) significant effect on both the number of tools and the average implemented Henrich’s (2004) model with a spatially struc- number of parts per tool. Collard et al. (2013a) applied simple tured metapopulation and found that contact and migration linear and stepwise multiple regression analysis to data from affect cultural evolution in a similar manner to increase in 45 nonindustrial farming and pastoralist groups to test the population size. Mesoudi (2011) showed that Henrich’s hypothesis. Results of the analyses were consistent with the (2004) results could be replicated when acquisition costs are predictions of the hypothesis: both the number of tools and allowed to increase as skill level increases. Kobayashi and Aoki the number of tool parts were positively and significantly (2012) modified Henrich’s (2004) model to examine the ef- influenced by population size in the simple linear regression fects of overlapping generations and found that the effects of analyses. The multiple regression analyses demonstrated that population size on cultural evolution are amplified when gen- these correlations were independent of the effects of risk of erational overlap is taken into account. resource failure. Collard et al. (2013a) concluded from this The idea that population size can affect cultural evolution that population size influences cultural evolution in recent has also been supported by formal and agent-based models nonindustrial food-producing populations. involving selectively neutral traits. For example, Neiman Other empirical studies do not support the population size (1995) investigated the amount of variation to be expected hypothesis. Collard, Kemery, and Banks (2005) included pop- in the decoration of a pottery assemblage if the motifs are ulation size in a study designed to shed light on the drivers neutral in terms of adaptation and showed that random loss, of food-getting tool kit structure among recent hunter-gath- or “drift,” destroys variation more quickly in smaller popu- erers. The core of their data set comprised counts of the lations than in larger ones. More recently, Premo and Kuhn number of tools and tool parts in the tool kits of a worldwide (2010) used an agent-based model to show that local group sample of 20 recent hunter-gatherer populations. Their results extinction can reduce cultural richness and complexity even did not support the population size hypothesis. The only when cultural traits do not affect fitness. variables that had a significant effect on the tool kit structure The situation with regard to empirical support for the pop- measures were measures of risk of resource failure, effective ulation size hypothesis is more complicated. To date, only temperature, and net aboveground productivity. Read (2008) Powell, Shennan, and Thomas (2009) have attempted to test also tested the hypothesis as part of an investigation into the hypothesis with Paleolithic archaeological data. They used factors that drive variation in tool kit structure among recent molecular data to estimate when different regions of the world hunter-gatherers. Like Collard, Kemery, and Banks (2005), he would have reached the same population density as Europe found no support for the hypothesis. Nelson et al. (2011) at the start of the Upper Paleolithic and then compared those used archaeological data to examine the relationship between estimates with the timing of the appearance of markers of regional population density and the number of pottery wares modern behavior in the regions. Their results were mixed. in the U.S. Southwest between 1000 CE and 1600 CE. They They found a reasonable correspondence between the timing found population density and pottery richness to be inversely of the crossing of the density threshold and the timing of the correlated and argued that this indicates that social confor- appearance of markers of modern behavior in sub-Saharan mity becomes increasingly important as population density Africa, North Africa, and the Levant, but there was a consid- increases. Although Nelson and colleagues do not discuss the erable gap between the timing of the crossing of the density population size hypothesis, their results are clearly not con- threshold and the appearance of markers of modern behavior sistent with it. Finally, Collard et al. (2013b) tested the hy- in southern, northern, and central Asia. As such, Powell, pothesis as part of a study that focused on the drivers of Shennan, and Thomas’s (2009) results only partially support technological richness among 85 recent hunter-gatherer the population size hypothesis. groups from western North America. They found that the A number of other empirical studies have a bearing on the total number of material items and techniques was correlated S390 Current Anthropology Volume 54, Supplement 8, December 2013 with both a proxy for environmental risk—mean rainfall for man present and require only occasional monitoring (e.g., a the driest month—and population size. However, the direc- deadfall trap). Oswalt created a further distinction between tion of the relationship was the opposite of the one predicted simple and complex subsistants. Simple subsistants do not by the population size hypothesis (it was negative rather than change structurally during use, whereas complex subsistants positive). have multiple parts that change position relative to one an- Currently, then, the situation with respect to empirical sup- other during use. port for the population size hypothesis is mixed. Several stud- Oswalt (1973, 1976) devised three measures of tool kit ies support it, several refute it, and one study has yielded structure. One is the total number of subsistants, which Os- ambiguous results. Given this, and given the potential im- walt suggested is an indicator of the size of a tool kit. Other portance of the hypothesis for our understanding of the Pa- researchers have referred to the total number of subsistants leolithic and cultural evolution in general, we decided to re- as tool kit “diversity” (Collard, Kemery, and Banks 2005; Col- visit the relationship between population size and the number lard et al. 2011; Shott 1986; Torrence 1983, 1989), but this and intricacy of the food-getting tools used by recent hunter- term is potentially confusing. In ecology, “diversity” has two gatherer populations. Collard et al. (2011) recently reported dimensions: “richness” and “evenness.” The former refers to an analysis that suggests the effect of risk of resource failure the number of taxa in a community, landscape, or region; on hunter-gatherer food-getting technology is dependent on the latter refers to how close the taxa in a community, land- the scale of risk differences among populations. They found scape, or region are in terms of numbers of individuals (Col- that when there are large differences in risk of resource failure well 2009). The dimension of species diversity that the variable among populations, risk has a significant effect on the number “total number of subsistants present in a tool kit” is akin to and intricacy of the food-getting tools used by hunter-gath- is clearly “species richness.” Thus, to reduce the potential for erers. When differences in risk of resource failure are small, confusion, here we refer to the total number of subsistants in contrast, risk does not have a significant effect on the as “tool kit richness” rather than “tool kit diversity.” Another structure of hunter-gatherers’ food-getting tool kits. This find- of Oswalt’s measures of tool kit structure is the total number ing suggests that the conflicting results of studies that have of techno-units. Formally, a techno-unit is an “integrated, tested the population size hypothesis with ethnographic and physically distinct, and unique structural configuration that archaeological data may be more apparent than real. Specif- contributes to the form of a finished artifact” (Oswalt 1976: ically, it raises the possibility that the studies that have failed 38), but in simpler terms, techno-units are the different kinds to support the hypothesis have done so because they have of parts of a tool. The total number of techno-units included employed samples in which there are large risk differences, in a tool kit is a measure of its “complexity” (Collard, Kemery, and therefore the impact of population size on tool kit struc- and Banks 2005; Collard et al. 2011; Oswalt 1973, 1976; Torr- ture has been obscured by the impact of risk. With this in ence 1983, 1989). Oswalt’s third measure of tool kit structure mind, we tested the hypothesis with samples spanning three is the average number of techno-units per subsistant, which levels of among-population risk difference: a global sample is calculated by dividing the total number of techno-units in consisting of populations from several continents, a conti- a tool kit by its richness. Again, this is a measure of tool kit nental sample comprising populations from North America, complexity. and a regional sample made up of populations from the Pa- Using Oswalt’s (1973, 1976) method, we generated values cific Northwest. for total number of subsistants (STS), total number of techno- units (TTS), average number of techno-units per tool (AVE), Material and Methods and population size (POP) for a sample of 49 contact-era hunter-gatherer populations. Thirty of the populations are Oswalt (1973, 1976) developed the method we used in the from North America, five are from South America, five are study. The method focuses on tools employed directly in the from Africa, five are from Asia, and four are from Oceania. acquisition of food, which Oswalt termed “subsistants.” Os- The names and locations of the populations are given in table walt divided subsistants into four categories: instruments, 1. The majority of the tool kit data was taken from previous weapons, tended facilities, and untended facilities. Instru- studies that have used Oswalt’s (1973, 1976) method to quan- ments are used to procure food that cannot run away or tify tool kit structure (Collard, Kemery, and Banks 2005; Col- threaten its pursuer, such as plants or sessile animals. A dig- lard et al. 2011). These data were supplemented with STS, ging stick is an example of an instrument. Weapons are de- TTS, and AVE values generated specifically for this study. The signed to kill or maim potential prey that can escape or may sources from which the latter data were extracted vary in age harm its pursuer. Weapons include boomerangs, crossbows, from the late 1800s to the mid-20th century. The values for and harpoons. Facilities are structures that control the move- POP were taken from Binford (2001). The POP data were ment of animals or protect them to a human’s advantage, transformed to base e because the POP hypothesis predicts a such as a fish weir or a livestock pen. Tended facilities require concave relationship between POP and tool kit richness and continuous monitoring while in use (e.g., a fishhook), whereas complexity. Log-transforming POP made the expected rela- untended facilities are capable of functioning without a hu- tionship between POP and each tool kit measure a linear one. Collard, Buchanan, and O’Brien Population Size and the Paleolithic Archaeological Record S391

Table 1. Names and locations of hunter-gatherer pothesis found that POP did not affect tool kit structure when groups included in the samples measures of risk of resource failure and measures of mobility were included in the analysis (Collard, Kemery, and Banks Name Location 2005; Read 2008). The proxies for risk of resource failure were Mbuti Africa effective temperature (ET), net aboveground productivity Hadza Africa (NAGP), and mean rainfall for the wettest month of the year !Kung San Africa (RHIGH). Also known as “warmth,” ET was developed to Nharo Africa G/Wi Africa better understand the effect of temperature on the distribution Punan Asia of living and fossil plants (Bailey 1960). It is defined as the Great Andamanese Asia temperature characteristic of the start and finish of the period Veddas Asia in which plant growth occurs (Bailey 1960). NAGP is the Chenchu Asia amount of new cell life that is added to a given location by Yukaghir Asia Copper Inuit North America, arctic photosynthesis and growth in a year (measured in grams per Iglulik North America, arctic square meters per year; Binford 2001). The measures of res- Netsilik North America, arctic idential mobility we included were number of residential Angmagsalik North America, arctic moves per year (NOMOV) and total distance moved per year Tareumiut North America, arctic during residential moves (DISMOV). The values for ET, Twana North America, Pacific Northwest Nootka North America, Pacific Northwest NAGP, RHIGH, NOMOV, and DISMOV were obtained from Quinalt North America, Pacific Northwest Binford (2001). Upper Stalo North America, Pacific Northwest After compiling the data set, we used the Kolmogorov- Coast Salish North America, Pacific Northwest Smirnov test to assess how closely the variables approximate Makah North America, Pacific Northwest a normal distribution. In the global sample, ET, NAGP, Kwakiutl North America, Pacific Northwest Tlingit North America, Pacific Northwest RHIGH, and DISMOV were found to have distributions that Klamath North America, plateau departed significantly from normal and thus were trans- Lillooet North America, plateau formed. We transformed NAGP, RHIGH, and DISMOV using Coeur D’Alene North America, plateau the natural log. To transform ET, we applied the Box-Cox Okanagan North America, plateau power transformation in Minitab. Because a negative value Sanpoil-Nespelem North America, plateau Ϫ Shuswap North America, plateau was selected as the λ parameter ( 1.910), all values were then Owens Valley Paiute North America, Southwest subtracted from 1. In the North American sample, the only Surprise Valley Paiute North America, Southwest variable whose distribution departed significantly from a nor- Fort Nelson Slave North America, subarctic mal distribution was DISMOV. We transformed it using the Kaska North America, subarctic natural log. None of the distributions were significantly dif- Carrier North America, subarctic Lower Koyukon North America, subarctic ferent from normal in the Pacific Northwest sample. Chipewyan North America, subarctic After completing the transformations, we carried out three Ingalik North America, subarctic sets of analyses. In the first, we used simple correlation analysis Nabesna North America, subarctic to assess the direction and strength of the correlation between Tanaina North America, subarctic population size and each of the three tool kit variables (STS, Caribou Inuit North America, subarctic Tiwi Oceania TTS, AVE). Here, as in the other tests, we began with the Groote-eylandt Oceania global sample, then analyzed the North American sample, and Northern Arenda Oceania then the Pacific Northwest sample. The test prediction was Tasmanians Oceania that the relationships between tool kit variables and popu- Yaruro South America lation size should be both positive and statistically significant. Siriono South America Botocudo South America Because multiple tests were conducted, Benjamini and Yek- Ona South America utieli’s (2001) method of significance level correction was used Yahgan South America to reduce Type I error rates. We employed this method rather than the better-known Bonferroni correction because it has been shown to balance the reduction of Type I and Type II Some researchers contend that the technological variables error rates better than the Bonferroni correction (Narum should also be logged when testing the population size hy- 2006). pothesis. We tried this approach as well, and the results were In the second set of analyses, partial correlation analysis not qualitatively different. was used to assess the direction and strength of the correlation In addition to generating technological and POP data, we between population size and each of the three tool kit variables obtained values for three measures of risk of resource failure while controlling for the risk variables (ET, NAGP, RHIGH) and two measures of residential mobility. We did so because, and the mobility variables (NOMOV, DISMOV). The test as mentioned earlier, some previous tests of the POP hy- prediction was the same as the one in the previous set of S392 Current Anthropology Volume 54, Supplement 8, December 2013

Table 2. Summary of results of simple correla- tables 4–6. The only significant influences on tool kit richness tion analyses carried out to assess the strength of and diversity were the risk variables ET and RHIGH and the the relationship between population size (POP) mobility variables NOMOV and DISMOV. POP was not a and tool kit richness and complexity significant influence on any of the tool kit variables in any of the samples; it consistently had one of the lowest stan- Sample and variables dardized beta coefficients; and it was always either the fourth correlated rP or fifth lowest of the six independent variables. Global (n p 49 ): STS, POP .353 .013a TTS, POP .361 .011a Discussion and Conclusions AVE, POP .179 .219 North America (n p 30 ): POP was correlated with two of the three tool kit variables STS, POP .318 .086 in the global sample, but these relationships disappeared when TTS, POP .383 .037 partial correlation analysis was used to control for risk of AVE, POP .230 .221 resource failure and mobility, both of which have previously p Pacific Northwest (n 14 ): been found to influence tool kit richness and complexity STS, POP .455 .102 TTS, POP .525 .054 among hunter-gatherers. POP was not correlated with any of AVE, POP .204 .484 the tool kit variables in the North American and Pacific Northwest samples regardless of which form of correlation Note. STS p total number of subsistants; TTS p total number of techno-units; AVE p average number of techno- analysis was used. The regression analyses were consistent with units per tool. a Significant correlation using Benjamini and Yekutieli’s Table 3. Summary of results of partial correlation analyses (2001) alpha correction (the critical value for three tests is carried out to assess the strength of the relationship be- α p .027). tween population size (POP) and tool kit richness and complexity while controlling for variables that have previ- analyses: relationships between the tool kit variables and pop- ously been found to influence the richness and complexity ulation size should be both positive and statistically signifi- of hunter-gatherer tool kits cant. In the third set of analyses, we used standard multiple Sample, variables correlated, and regression analysis to assess the importance of population size variables controlled for rP as an influence on tool kit richness and complexity compared Global (n p 49 ): with the risk and mobility variables. The tool kit variables STS, POP: were the dependent variables, and population size, ET, NAGP, ET,a NAGP,b DISMOV,b RHIGH,b NOMOV .246 .107 RHIGH, DISMOV, and NOMOV were the independent var- TTS, POP: a b b b iables. When the variance inflation factor (VIF) for two or ET, NAGP, DISMOV, RHIGH, NOMOV .248 .104 AVE, POP: more variables exceeded 10, the variable with the highest VIF ET,a NAGP,b DISMOV,b RHIGH,b NOMOV Ϫ.017 .911 was removed and the analysis rerun. PASW (SPSS) 19 was North America (n p 30 ): used to carry out all the analyses. STS, POP: ET, NAGP, DISMOV,b RHIGH, NOMOV .289 .160 TTS, POP: Results ET, NAGP, DISMOV,b RHIGH, NOMOV .349 .087 Results of the simple correlation analyses are summarized in AVE, POP: ET, NAGP, DISMOV,b RHIGH, NOMOV .178 .395 table 2. In the analyses of the global sample, STS and TTS Pacific Northwest (n p 14 ): were significantly correlated with POP,but AVE was not. None STS, POP: of the tool kit variables were significantly correlated with POP ET, NAGP, DISMOV, RHIGH, NOMOV .428 .250 in the analyses of the North American sample or for the Pacific TTS, POP: Northwest sample. ET, NAGP, DISMOV, RHIGH, NOMOV .179 .491 AVE, POP: Table 3 summarizes results of the partial correlation anal- ET, NAGP, DISMOV, RHIGH, NOMOV .339 .372 yses. None of the tool kit variables were significantly corre- Note. STS p total number of subsistants; TTS p total number of lated with POP in the analyses of the global sample. The techno-units; AVE p average number of techno-units per tool; ET p results of the analyses using the North American sample were effective temperature; NAGP p net aboveground productivity; RHIGH similar: None of the tool kit variables were significantly cor- p rainfall for the wettest month of the year; NOMOV p number of related with POP. The results of the analyses that focused on residential moves per year; DISMOV p total distance moved per year the Pacific Northwest sample were consistent with the results during residential moves. a Transformed with Box-Cox method and reciprocal taken before anal- of the analyses of the other two samples. Once again, none ysis; see “Material and Methods” section for details. of the tool kit variables were significantly correlated with POP. b Converted to natural logarithm before analysis; see “Material and Meth- The results of the regression analyses are summarized in ods” section for details. Collard, Buchanan, and O’Brien Population Size and the Paleolithic Archaeological Record S393

Table 4. Summary of results of standard multiple regression analyses using the global sample (n p 49 ) carried out to assess the relative importance of various variables as drivers of tool kit richness and complexity

Dependent variable Full model POP ET NAGP RHIGH NOMOV DISMOV STS F p 6.199, β p .195, β p Ϫ.658, β p .204, β p Ϫ.264, β p Ϫ.265, β p .011, df p6, 42, P p .107, P p .002,b P p .379, P p .039,b P p .143, P p .954, P p .000,a VIF p 1.116 VIF p 3.310 VIF p 4.183 VIF p 1.215 VIF p 2.495 VIF p 2.940 r2 p .470 TTS F p 7.745, β p .187, β p Ϫ.740, β p .271, β p Ϫ.298, β p Ϫ.226, β p Ϫ.146, df p 6, 42, P p .104, P p .000,b P p .220, P p .015,b P p .186, P p .428, P p .000,a VIF p 1.116 VIF p 3.310 VIF p 4.183 VIF p 1.215 VIF p 2.495 VIF p 2.940 r2 p .525 AVE F p 5.469, β p Ϫ.014, β p Ϫ.544, β p Ϫ.012, β p Ϫ.222, β p Ϫ.097, β p Ϫ.409, df p 6, 42, P p .911, P p .013,b P p .959, P p .088, P p.598, P p .045,b P p .000,a VIF p 1.116 VIF p 3.310 VIF p 4.183 VIF p 1.215 VIF p 2.495 VIF p 2.940 r2 p .439 Note. STS p total number of subsistants; POP p population size; TTS p total number of techno-units; AVE p average number of techno-units per tool; ET p effective temperature; NAGP p net aboveground productivity; RHIGH p rainfall for the wettest month of the year; NOMOV p number of residential moves per year; DISMOV p total distance moved per year during residential moves; VIF p variance inflation factor. a Significant correlation using Benjamini and Yekutieli’s (2001) alpha correction (the critical value for three tests isα p .027 ). b Significant atP ≤ .05 . the results of the partial correlation analyses: POP was not a is adequate. Additionally, one of the other studies that have significant influence on any of the tool kit variables in any failed to support the POP hypothesis (Collard et al. 2013b) of the three samples and consistently had one of the lowest cannot be criticized for not taking into account cultural trans- standardized beta coefficients. Thus, the analyses did not sup- mission among populations. It controlled for cultural trans- port the POP hypothesis. Even when the influence of the mission and still failed to find support for the hypothesis. factor that previously has been found to most affect tool kit Thus, use of inadequate estimates of POP seems unlikely to richness and complexity among hunter-gatherers—risk of re- explain the failure of Collard, Kemery, and Banks’s (2005), source failure—was minimized, there was no evidence that Read’s (2008), Collard et al.’s (2013b), and this study to sup- POP influenced tool kit richness and complexity. port the POP hypothesis. This means that there are now four empirical studies that A second potential shortcoming that needs to be considered support the POP hypothesis (Collard et al. 2013a; Kline and is sample size. In principle, it is possible that the studies that Boyd 2010; Neiman 1995; Powell, Shennan, and Thomas have failed to support the hypothesis have done so because 2009) and four that do not (Collard, Kemery, and Banks 2005; the samples they used were too small to pick up the influence Collard et al. 2013b; Nelson et al. 2011; this study). There are of POP, but this seems unlikely. Samples employed in the two basic potential explanations for this disagreement. One studies that have tested the hypothesis with ethnographic data is that the studies that have failed to support the hypothesis and found support for it comprised 10 and 45 populations, suffer from shortcomings that are sufficiently serious to have respectively (Collard et al. 2013a; Kline and Boyd 2010). The resulted in Type II errors; that is, the studies’ failure to support majority of the samples that have failed to support the hy- the hypothesis is a false negative. The other is that the results pothesis are larger than Kline and Boyd’s (2010) sample, and of the studies that have failed to support the hypothesis are some of them are larger than Collard et al.’s (2013a) sample. reliable and the hypothesis needs modification. The samples used by Collard, Kemery, and Banks (2005) and Regarding the first possibility, there are three potential by Read (2008) comprised 20 hunter-gatherer populations. shortcomings that need to be evaluated. One is the accuracy Collard et al.’s (2013b) sample consisted of 85 populations. of the population estimates. Henrich (2006), Kline and Boyd Samples employed in the study reported here range in size (2010), and Boyd, Richerson, and Henrich (2013) have argued from 14 populations to 49 populations. As such, it is unlikely that the studies by Collard, Kemery, and Banks (2005) and that small sample size explains the failure of the studies of Read (2008) failed to support the hypothesis because they did Collard, Kemery, and Banks (2005), Read (2008), Collard et not take into account cultural transmission among popula- al. (2013b), and the one reported here to support the POP tions and therefore did not accurately measure the effective hypothesis. POP for cultural traits. This is unlikely. Population values for A third potential shortcoming concerns sample bias. In both studies were generated in the same way as the values addition to suggesting that Collard, Kemery, and Banks’s used by Collard et al. (2013a), which supported the hypoth- (2005) population estimates are inaccurate, Kline and Boyd esis. That Collard et al.’s (2013a) study supported the hy- (2010) and Boyd, Richerson, and Henrich (2013) claim that pothesis implies that the method of collecting population data Collard and colleagues’ results are unreliable because North S394 Current Anthropology Volume 54, Supplement 8, December 2013

Table 5. Summary of results of standard multiple regression analyses using the North American sample (n p 30 ) carried out to assess the relative importance of various variables as drivers of tool kit richness and complexity

Dependent variable Full model POP ET NAGP RHIGH NOMOV STS F p 1.526, β p .239, β p Ϫ.251, β p .810, β p Ϫ.479, β p Ϫ.116, df p 5, 24, P p .218, P p .319, P p .094, P p .241, P p .583, P p.219, VIF p 1.125 VIF p 1.923 VIF p 6.813 VIF p 5.026 VIF p 1.371 r2 p .241 TTS F p 3.351, β p .283, β p Ϫ.441, β p .675, β p Ϫ.291, β p Ϫ.337, df p 5, 24, P p .102, P p .054, P p .112, P p .416, P p .079, P p .020,a VIF p 1.125 VIF p 1.923 VIF p 6.813 VIF p 5.026 VIF p 1.371 r2 p .411 AVE F p 3.892, β p .146, β p Ϫ.493, β p .117, β p .151, β p Ϫ.500, df 5, 24, P p .372, P p .028,b P p .771, P p .662, P p .010,b P p .010,a VIF p 1.125 VIF p 1.923 VIF p 6.813 VIF p 5.026 VIF p 1.371 r2 p .448 Note. DISMOV was excluded from the analysis because of multicollinearity with NOMOV (see “Material and Methods” for details). STS p total number of subsistants; POP p population size; TTS p total number of techno-units; AVE p average number of techno-units per tool; ET p effective temperature; NAGP p net aboveground productivity; RHIGH p rainfall for the wettest month of the year; NOMOV p number of residential moves per year; DISMOV p total distance moved per year during residential moves; VIF p variance inflation factor. a Significant using Benjamini and Yekutieli’s (2001) alpha correction (the critical value for three tests isα p .027 ). b Significant atP ≤ .05 .

American populations dominate their sample. Collard et al.’s TTS (r p 0.185 ,P p .448 ) or AVE (r p Ϫ0.100 ,P p .683 ). (2013b) sample consists solely of North American popula- Thus, the balanced global sample did not support the POP tions, and North American populations also dominate the hypothesis. This indicates that the failure of this study to sample used in the study reported here. Thus, Kline and support the hypothesis cannot be explained away as a con- Boyd’s (2010) and Boyd, Richerson, and Henrich’s (2013) sequence of sample bias and suggests the same holds for the concerns can be extended to the other studies that have failed other studies that have not supported it. to support the hypothesis, but there are reasons to think their It appears, then, that the disagreement between the em- concerns are unwarranted. To begin with, there is an impor- pirical studies that support the hypothesis and those that do tant corollary to the idea that the studies that have failed to not is not a consequence of the latter studies suffering from support the hypothesis have done so because North American shortcomings that are sufficiently serious to have resulted in samples dominate the samples. The corollary is that the hy- Type II errors. Rather, it appears that the disagreement is pothesis does not apply to North American populations but substantive. rather to populations from other regions of the world. Thus, What might account for the disagreement? So far, we have even if the sample-bias argument were correct, it would re- been able to identify four potential answers to this question. quire us to revise the POP hypothesis to explain its failure to One concerns the mode of production. Samples that have apply to North America populations. In other words, the supported the hypothesis comprise populations that were sample-bias argument simply changes the scope of the ref- heavily dependent on domesticated species (Collard et al. utation of the hypothesis rather than explaining away the 2013a; Kline and Boyd 2010; Neiman 1995), whereas the ma- failure to support the hypothesis. jority of samples that have refuted the hypothesis consist of Another reason to reject the claims of Kline and Boyd populations that relied primarily on wild resources (Collard, (2010) and Boyd, Richerson, and Henrich (2013) is that we Kemery, and Banks 2005; Collard et al. 2013b; Read 2008). obtained similar results with a balanced global sample to our Consequently, it could be that mode of production mediates original global sample. We used a random-number generator the effect of POP on cultural evolution such that the tech- to select one population from each of five culture regions nology of food producers is more affected by POP than by represented among the North American populations in the risk, whereas the technology of hunter-gatherers is more af- sample. We then deleted the other 25 North American pop- fected by risk than by POP. The problem with this proposal ulations. This left us with 24 populations: five from North is that Nelson et al.’s (2011) data relate to small-scale farming America, five from South America, five from Africa, five from groups. This makes the idea that mode of production mediates Asia, and four from Oceania. We then repeated the partial the effect of POP on cultural evolution less plausible given correlation analyses in which we correlated STS, TTS, and that it means that groups with a food-producing mode of AVE with POP while controlling for ET, NAGP, RHIGH, production both support and refute the hypothesis. NOMOV, and DISMOV. STS was not positively and signifi- A second possibility is that there is a threshold effect in cantly correlated with POP (r p 0.193 ,P p .428 ), nor was the influence of POP on technology. Populations that support Collard, Buchanan, and O’Brien Population Size and the Paleolithic Archaeological Record S395

Table 6. Summary of results of standard multiple regression analyses using the Pacific Northwest sample (n p 14 ) carried out to assess the relative importance of various variables as drivers of tool kit richness and complexity

Dependent Full variable model POP ET NAGP RHIGH NOMOV DISMOV STS F p 1.753, β p .344, β p Ϫ.181, β p .348, β p .085, β p Ϫ1.028, β p 1.620, df p 6, 7, P p .250, P p .535, P p .553, P p .883, P p .160, P p .059, P p .240, VIF p 1.323 VIF p 1.354 VIF p 5.479 VIF p 5.444 VIF p 7.488 VIF p 9.034 r2 p .600 TTS F p 1.289, β p .447, β p .196, β p .839, β p Ϫ.345, β p Ϫ.803, β p 1.063, df p 6, 7, P p .179, P p .539, P p .211, P p .588, P p .297, P p .217, P .370, VIF p 1.323 VIF p 1.354 VIF p 5.479 VIF p 5.444 VIF p 7.488 VIF p 9.034 r2 p .525 AVE F p 1.862, β p .258, β p .608, β p .780, β p Ϫ.610, β p .210, β p Ϫ.681, df p 6, 7, P p .372, P p .061, P p .198, P p .302, P p .753, P p .366, P p .217, VIF p 1.323 VIF p 1.354 VIF p 5.479 VIF p 5.444 VIF p 7.488 VIF p 9.034 r2 p .615 Note. STS p total number of subsistants; POP p population size; TTS p total number of techno-units; AVE p average number of techno-units per tool; ET p effective temperature; NAGP p net aboveground productivity; RHIGH p rainfall for the wettest month of the year; NOMOV p number of residential moves per year; DISMOV p total distance moved per year during residential moves; VIF p variance inflation factor. the hypothesis and those that do not overlap in terms of size, about whom they copy. Indeed, they show that in certain but several of the former are much larger than the largest of circumstances (e.g., if individuals copy the most popular be- the latter. Thus, it could be that POP does not have a sig- havior or copy from each other at random), cultural com- nificant effect on cultural evolution until it is greater than a plexity can increase or decrease regardless of POP. One cor- value close to or above the upper end of the populations that ollary of their findings is that the POP effect is likely to be support the POP hypothesis, which is ∼12,000 people. How- mediated by degree of task specialization. Given that skill level ever, the modeling work of Shennan (2001) and Henrich is primarily a result of practice time (Ericsson and Charness (2004) suggests that the effect of POP on cultural evolution 1994), task specialization can be expected to increase the dif- should be greatest when POP is less than a few thousand, so ference in skill level between the most skilled individual within a threshold effect where the POP holds for larger populations a group and the majority of group members. This means that but not for smaller ones also seems unlikely to be the expla- the POP effect should be more pronounced in populations nation for the disagreement. with more task specialization than in populations with less The other two potential explanations involve social factors. task specialization. Thus, it is possible that the disagreement Henrich (2010) has argued that norms and institutions that among the studies that have tested the POP hypothesis is a foster sharing can positively affect the spread of inventions consequence of populations that support the hypothesis hav- within a population. Thus, it could be that sharing norms ing more task specialization than populations that do not and institutions can mediate the effects of POP on cultural support it. At the moment, we are not in a position to de- evolution such that a small population with numerous and/ termine which, if either, of these hypotheses is correct. Doing or strong sharing norms and institutions is equivalent or even so will require further modeling work and cross-cultural stud- better in terms of its ability to retain beneficial inventions ies. than a large population with few and/or weak sharing norms Together, the study reported here and the other studies that and institutions. If this is the case, then it is possible that the have failed to support the POP hypothesis have implications disagreement among the studies is the result of populations for interpreting the Paleolithic archaeological record. As we that support the hypothesis having fewer and/or weaker shar- noted earlier, it has become commonplace to use POP to ing norms and institutions than populations that do not sup- explain patterns in the record, but given that the record was port it. Another possibility is that the disagreement is a con- produced exclusively by hunter-gatherers, the failure of the sequence of differences in degree of task specialization. study reported here and the studies of Collard, Kemery, and Recently, Bentley and O’Brien (2011) argued that the effect Banks (2005), Read (2008), and Collard et al. (2013b)to of POP documented by Henrich (2004) and Powell, Shennan, support the hypothesis challenges these interpretations. If the and Thomas (2009) depends on two strong assumptions: (1) richness and complexity of the technology of recent hunter- the skill level of the most skilled member of the group is gatherers are not affected by POP, there would seem to be several times greater than the skill level of the average group little reason to expect changes in POP to be a broadly useful member, and (2) all learners can identify and copy the most hypothesis for explaining patterns in the Paleolithic archae- skilled member of the group. Bentley and O’Brien demon- ological record given that it was produced exclusively by strate that the POP effect is reduced if skill level is normally hunter-gatherers. The same holds for stability in POP. POP distributed within a group and/or if people are less selective change/stability may explain some of the patterns the Pale- S396 Current Anthropology Volume 54, Supplement 8, December 2013 olithic archaeological record, but this needs to be demon- in the Pacific Northwest. Philosophical Transactions of the Royal Society B 366:1129–1138. strated on a case-by-case basis through tests in which the POP Collard, Mark, Briggs Buchanan, Michael J. O’Brien, and Jonathan Scholnick. hypothesis is pitted against competing hypotheses. As men- 2013a. Risk, mobility, or population size? drivers of technological diversity tioned earlier, several studies have suggested that environ- among recent western North American hunter-gatherers. Philosophical Transactions of the Royal Society B 368:20120412. mental risk is the primary driver of technological richness Collard, Mark, Michael Kemery, and Samantha Banks. 2005. Causes of toolkit and complexity among recent hunter-gatherers (Collard, variation among hunter-gatherers: a test of four competing hypotheses. Kemery, and Banks 2005; Collard et al. 2013b;Read2008; Canadian Journal of Archaeology 29:1–19. Collard, Mark, April Ruttle, Briggs Buchanan, and Michael J. O’Brien. 2013b. Torrence 1983, 1989). Thus, adaptation to environmental Population size and cultural evolution in non-industrial food-producing conditions is one hypothesis that should be included in such societies. PLoS ONE 8:e72628. tests. Another factor that should probably be taken into ac- Colwell, Robert K. 2009. Biodiversity: concepts, patterns and measurement. In The Princeton guide to ecology. Simon A. Levin, ed. Pp. 257–263. Prince- count is social conformity, given that it has the capacity to ton, NJ: Princeton University Press. affect collective action (Nelson et al. 2011). Regardless of Ericsson, K. Anders, and Neil Charness. 1994. Expert performance: its struc- which competing hypotheses are considered, simply attrib- ture and acquisition. American Psychologist 49:725–747. Henrich, Joseph. 2004. Demography and cultural evolution: why adaptive uting patterns in the Paleolithic archaeological record to POP cultural processes produced maladaptive losses in Tasmania. American An- is not a defensible course of action. tiquity 69:197–218. ———. 2006. Understanding cultural evolutionary models: a reply to Read’s critique. American Antiquity 71:771–782. ———. 2010. The evolution of innovation-enhancing institutions. In Inno- Acknowledgments vation in cultural systems: contributions from evolutionary anthropology.Mi- chael J. O’Brien and Stephen J. Shennan, eds. Pp. 203–223. Cambridge, We thank Steven Kuhn and Erella Hovers for the invitation MA: MIT Press. Kline, Michelle A., and Robert Boyd. 2010. Population size predicts tech- to participate in the “Alternative Pathways to Complexity: nological complexity in Oceania. Proceedings of the Royal Society B 277: Evolutionary Trajectories in the Middle Paleolithic and Mid- 2559–2564. dle Stone Age” symposium and Leslie Aiello and Laurie Ob- Kobayashi, Yutaka, and Kenichi Aoki. 2012. Innovativeness, population size and cumulative cultural evolution. Theoretical Population Biology 82:38–47. bink of the Wenner-Gren Foundation and the other partic- Mesoudi, Alex. 2011. Variable cultural acquisition costs constrain cumulative ipants in the symposium for making it such an enjoyable and cultural evolution. PLoS ONE 6:e18239. intellectually stimulating experience. We also thank Michael Narum, Shawn R. 2006. Beyond Bonferroni: less conservative analyses for conservation genetics. Conservation Genetics 7:783–787. Kemery and Jesse Morin for assistance with data collection. Neiman, Fraser D. 1995. Stylistic variation in evolutionary perspective: in- Mark Collard is supported by the Canada Research Chairs ferences from decorative diversity and interassemblage distance in Illinois Program, the Social Sciences and Humanities Research Coun- Woodland ceramic assemblages. American Antiquity 60:7–36. Nelson, Margaret C., Michelle Hegmon, Stephanie R. Kulow, Matthew A. cil, the Canada Foundation for Innovation, the British Co- Peeples, Keith W. Kintigh, and Ann P. Kinzig. 2011. Resisting diversity: a lumbia Knowledge Development Fund, and Simon Fraser long-term archaeological study. Ecology and Society 16(1):25. University. Briggs Buchanan is supported by the University Oswalt, Wendell H. 1973. Habitat and technology: the evolution of hunting. New York: Holt, Rinehart, & Winston. of Missouri and Simon Fraser University. The opinions ex- ———. 1976. An anthropological analysis of food-getting technology. New York: pressed in this paper do not necessarily reflect the views of Wiley. the aforementioned funding bodies. Powell, Adam, Stephen J. Shennan, and Mark G. Thomas. 2009. Late Pleis- tocene demography and the appearance of modern human behavior. Science 324:1298–1301. References Cited Premo, Luke S., and Steven L. Kuhn. 2010. Modeling effects of local extinctions on culture change and diversity in the Paleolithic. PLoS ONE 5:e15582. Bailey, Harry P. 1960. A method of determining the warmth and temperateness Read, Dwight. 2008. An interaction model for resource implement complexity of climate. Geografiska Annaler 42:1–16. based on risk and number of annual moves. American Antiquity 73:599– Benjamini, Yoav, and Daniel Yekutieli. 2001. The control of false discovery 625. rate under dependency. Annals of Statistics 29:1165–1188. Riede, Felix. 2008. The Laacher See-eruption (12,920 BP) and material culture Bentley, R. Alexander, and Michael J. O’Brien. 2011. The selectivity of social change at the end of the Allerød in Northern Europe. Journal of Archae- learning and the tempo of cultural evolution. Journal of Evolutionary Psy- ological Science 35:591–599. chology 9:125–141. Shennan, Stephen J. 2001. Demography and cultural innovation: a model and Binford, Lewis R. 2001. Constructing frames of reference: an analytical method some implications for the emergence of modern human culture. Cambridge for theory building using ethnographic and environmental data sets. Berkeley: Archaeological Journal 11:5–16. University of California Press. Shott, Michael. 1986. Technological organization and settlement mobility: an Boyd, Robert, Peter J. Richerson, and Joseph Henrich. 2013. The cultural ethnographic examination. Journal of Anthropological Research 42:15–51. evolution of technology: facts and theories. In Cultural evolution: society, Torrence, Robin. 1983. Time budgeting and hunter-gatherer technology. In technology, language, and religion. Peter J. Richerson and Morten H. Chris- Hunter-gatherer economy in prehistory: a European perspective. Geoff Bailey, tiansen, eds. Pp. 119–142. Cambridge, MA: MIT Press. ed. Pp. 11–22. Cambridge: Cambridge University Press. Collard, Mark, Briggs Buchanan, Jesse Morin, and Andre Costopolous. 2011. ———. 1989. Re-tooling: towards a behavioral theory of stone tools. In Time, What drives the evolution of hunter-gatherer subsistence technology? a energy and stone tools. Robin Torrence, ed. Pp. 57–66. Cambridge: Cam- reanalysis of the risk hypothesis with data from early contact era populations bridge University Press. Current Anthropology Volume 54, Supplement 8, December 2013 S397

Measuring the Complexity of Lithic Technology

by Charles Perreault, P. Jeffrey Brantingham, Steven L. Kuhn, Sarah Wurz, and Xing Gao

CAϩ Online-Only Material: Supplementary Table

Assessments of the complexity of lithic technologies coming from different time periods, regions, or hominid species are recurrent features of the literature on Paleolithic archaeology. Yet the notion of lithic complexity is often defined intuitively and qualitatively, which can easily lead to circular arguments and makes difficult the comparison of assemblages across different regions and time periods. Here we propose, in the spirit of Oswalt’s techno-units, that the complexity of lithic technology can be quantified by counting the procedural units involved in tool manufacture. We define procedural units as mutually exclusive manufacturing steps that make a distinct contribution to the finished form of a technology. As a proof of concept, we use the procedural-unit approach to measure the complexity of 13 Paleolithic assemblages. While preliminary, these results provide a quantitative benchmark confirming that lithic technological complexity increased throughout the Paleolithic period. The method to measure lithic complexity outlined here will allow us to revisit several claims made about change in technological complexity during human evolution.

Introduction and Sisk 2010) or increasing the productivity and thereby aiding the dispersal of our species out of Africa (Mellars 2006b). Several processes can also drive changes in techno- Arguments about the complexity of lithic technologies coming logical complexity. Studying changes in technological com- from different time periods, regions, or hominid species are plexity thus allows us to make inferences about how these recurrent features of the literature on Paleolithic archaeology. processes unfolded during human evolution. For instance, Technological complexity—or technological “simplicity,” putative increases in technological complexity have been in- “crudeness,” “refinement,” “sophistication,” or “advance- terpreted as signaling changes in cognitive abilities (e.g., Am- ment”—are thought to have had important effects on forager brose 2001, 2010; Coolidge and Wynn 2009; de Beaune 2004; populations, including broadening of ecological niches (Shea Foley 1987; Foley and Lahr 2003; Haidle 2010; Mellars 1989, 2006b; Wadley 2010) as well as the extent to which our an- cestors relied on social learning (Foley and Lahr 2003; Richer- Charles Perreault is Assistant Professor in the Department of Anthropology at the University of Missouri, Columbia (107 Swallow son and Boyd 2005). In line with theories of artifact design Hall, Columbia, Missouri 65211, U.S.A. [[email protected]]). and technological organization—which predict that techno- P. Jeffrey Brantingham is Professor in the Department of logical complexity will vary with factors such as prey choice, Anthropology at the University of California, Los Angeles (341 time budgeting, risk, labor costs, and mobility pattern (e.g., Haines Hall, Box 951553, Los Angeles, California 90095, U.S.A. Bleed 1986; Bousman 1993; Kelly 1995; Osborne 1999; Oswalt [[email protected]]). Steven L. Kuhn is Professor in the School of 1976; Torrence 1983, 1989)—changes in lithic technology Anthropology at the University of Arizona (Emil W. Haury Building, complexity have also been seen as a response to change in P.O. Box 210030, Tucson, Arizona 85721, U.S.A. [skuhn@ climate and environment (Mellars 1989, 2006a, 2006b; Shea email.arizona.edu]). Sarah Wurz is Senior Researcher at the Institute and Sisk 2010), in variation in energetic constraints and time for Human Evolution of the University of the Witwatersrand (Private budgeting (Shea and Sisk 2010), in hunting strategies (Mellars Bag 3, PO WITS, 2050, Johannesburg, South Africa [sarah.wurz@ wits.ac.za]). Xing Gao is Vice Director of the Institute of Vertebrate 1989), or in lithic raw material quality (Brantingham et al. Paleontology and Paleoanthropology at the Chinese Academy of 2000; Mellars 2006a, 2006b; Pope 1989; Schick 1994). Cultural Sciences (P.O. Box 643, Beijing 100044, China [[email protected]]). traditions or ethnic groups have also been interpreted as the This paper was submitted 3 VII 13, accepted 18 VII 13, and cause of spatial variation in lithic complexity (Movius 1944, electronically published 24 X 13. 1948; Schick 1994). Finally, there is even greater need for

᭧ 2013 by The Wenner-Gren Foundation for Anthropological Research. All rights reserved. 0011-3204/2013/54S8-0017$10.00. DOI: 10.1086/673264 S398 Current Anthropology Volume 54, Supplement 8, December 2013 rigorous measures of technological complexity in order to test of Paleolithic assemblages as a proof of concept. Our goal increasingly influential models linking demography and cul- here is not to test any specific hypothesis about the mecha- tural evolution (Beatty 1995; Foley and Lahr 2003; Mellars nisms driving changes in technological complexity but rather 2006a; Powell, Shennan, and Thomas 2009; Premo and Kuhn to develop a tool that can be used to test such hypotheses in 2010; Reiter 2000; Shennan 2001). a more rigorous manner than has been done so far. Consideration of technological complexity in the Paleo- lithic record is weakened, however, by a lack of explicit def- Measuring Lithic Technology Complexity inition of complexity. Often one technology is simply de- scribed as more complex than another without explanation. We define technological complexity as the minimum amount Such intuitive notions of technological complexity are dan- of information that is needed to manufacture a product. This gerous because they lead easily to circular arguments. For definition is in line with other formalized definitions of com- instance, methods such as prismatic blade technology have plexity (Shannon and Weaver 1949). Computer scientists, for been assumed to be more complex that the Levallois method example, have defined the complexity of an algorithm as the or bifacial shaping by the simple virtues of appearing later in shortest string length, or the smallest number of bits of in- time or being associated with modern humans (see Bar-Yosef formation, that is necessary to describe it (Chaitin 1970). This and Kuhn 1999). information criterion is analogous to the various measures of The complexity of Paleolithic technologies is also rarely richness used to describe biological systems that are defined quantified. Complexity is sometimes defined by the presence as the number of unique types of some constituent present or absence of qualitative features, such as composite tools within an aggregate group. For instance, at the level of the (Ambrose 2001, 2010; Coolidge and Wynn 2009), the use of organism, biological complexity has been measured as the compound adhesives (Wadley 2010), the use of a composite count of cell types (Bonner 1988). At the level of an ecosystem, tool to make another composite tool (Lombard and Haidle biological complexity has been measured as the count of 2012), or technologies that store energy exosomatically, such unique species it contains (Bonner 1988). Finally, the com- as the bow and arrow (Shea and Sisk 2010). But the usefulness plexity of animal behavior has also been estimated by counting of these qualitative definitions of technological complexity is the number of elemental “building blocks” that is associated limited because they are based on time-specific, region-spe- with a specific behavior or, at a larger scale, as the number cific, or culture-specific traits. They do not allow for the com- of acts in a species’ behavioral repertoire (Sambrook and parison of two assemblages if both either lack or possess these Whiten 1997; Whiten et al. 1999). traits. In other words, if the transition from simple to complex In the same spirit, we argue that the complexity of a tech- technologies is marked by the advent of composite tools, then nology can be measured by counting the number of elemental how does the complexity of two assemblages that lack com- building blocks associated with it. We call these building posite tools compare? Finally, these qualitative definitions also blocks “procedural units.” We define procedural units as mu- fail to measure how much more complex these traits are. For tually exclusive manufacturing steps that make a distinct con- instance, how much of a leap in technological complexity tribution to the finished form of the product of a technology. does the use of a composite tool really imply? Focusing on lithic technology, the count of procedural units Given its importance in Paleolithic research, we have much present in a tool reduction sequence is a measure of com- to gain from developing a more objective way of measuring plexity because it reflects the minimum amount of infor- technological complexity that is quantitative and not bound mation that is needed to carry it out to a successful end. to a specific time period, region, culture, or indeed any specific This procedural-unit approach to stone-tool complexity technology. We need a way to measure technological com- parallels Oswalt’s “techno-units” (Oswalt 1976). Oswalt as- plexity that will allow us to detect an increase in complexity sessed the complexity of food-getting technologies by count- marked by, for example, the advent of composite tools but ing (1) the number of tool types present in a tool kit, which without treating composite tools as a qualitative break. Finally, he called “subsistants,” and (2) the number of integrated and our system of measurement should allow us to compare not physically distinct structures that contribute to the finished just the complexity of different lithic assemblages but also of form of a tool, which he called “techno-units.” Oswalt’s lithic and nonlithic technologies. We should be able to com- method is powerful because it allows for the measurement of pare the complexity of Oldowan tools not only with that of technological complexity cross-culturally. It has been applied European Upper Paleolithic blades but also with that of a to ethnographic data to test a wide range of hypotheses, in- Boeing 747. cluding hypotheses about the ecological determinants of tech- Here we present a quantitative and widely applicable ap- nological complexity (Collard, Kemery, and Banks 2005; Col- proach to measuring technological complexity in the archae- lard et al. 2011; Shott 1986; Torrence 1983, 1989, 2000) and ological record. We propose that technological complexity can the effect of demography on the evolution of technologies be measured by counting the procedural units, or manufac- (Collard, Kemery, and Banks 2005; Collard et al. 2011; Kline turing “building blocks,” represented in an assemblage. Below and Boyd 2010; Oswalt 1976). we describe this method, followed by its application to a series Oswalt (1976:229–230) recognized that there are problems Perreault et al. Measuring the Complexity of Lithic Technology S399 with applying his concept of complexity to the archaeological as a different procedural unit. Conversely, the repeated use record. First, the number of tool types present in a prehistoric of a hard hammer within the same reduction step, such as tool kit is difficult to assess because the actual function of in the shaping of a core, should not be counted as multiple tools is difficult to infer. Moreover, when tool function can independent procedural units; each hammer blow serves the be determined, there is often overlap in the functional tasks same function, and the number of blows struck during the accomplished with what appear to be morphologically distinct preparation of the core may vary from one core to another tool types. Second, the techno-unit approach will lead to un- without it affecting the nature of the technology. This is anal- derestimating the complexity of prehistoric technologies be- ogous to Oswalt’s (1976:52) recommendation that physically cause of preservation biases. This is especially true for the distinct elements serving the same purpose, such as the balls Paleolithic record, were the range of preserved material is of a bola or the teeth of a rake, be counted as only one techno- narrow. For instance, in most situations a complex technology unit. Researchers thus need to decide on a case-by-case basis such as the bow and arrow would leave in the archaeological whether the same procedural unit at different reduction steps record only one techno-unit, the stone projectile point. Yet constitutes functionally distinct manufacturing steps or not. with full preservation, it is clear that bow-and-arrow tech- Taking all these things into account and focusing on uncon- nologies are more complex technology than simple handheld troversial features that are commonly discussed in the lithic scrapers, a 1-techno-unit technology. Attempting to infer analysis literature, we have identified 35 procedural units that what techno-units are missing in an archaeological assemblage may be associated with lithic technology. for taphonomic reasons is not a solution because any given technology can contain a varying number of techno-units. Preliminary Steps For instance, Oswalt’s data set (1976) contains examples of bows that range in complexity from 2 to 10 techno-units and (1) Raw material treatment: evidence of heat treatment of arrows that range from 2 to 13 techno-units. Finally, be- cause Oswalt’s approach focuses on the finished product, it Core Preparation Techniques fails to capture variation in the complexity involved in the (2) Decortification: cortex is removed making of different tools. For example, producing a prismatic (3) Shaping of platform: platform intentionally prepared blade can be more complex than making a digging stick, and by flaking yet both could be seen as simple, 1-techno-unit technologies (4) Shaping of flaking surface: face of flake removal is in- when analyzed using Oswalt’s method. tentionally prepared by flaking We can avoid to a certain extent these problems by focusing (5) Shaping of nonflaking surface: nonactive part of the on the chaıˆnes ope´ratoires of technologies rather than on the coreisshaped finished products. Counting the procedural units present in (6) Blades: crested or de´bordante blades used to align face a reduction sequence or in an assemblage allows us to avoid of flake removal having to identify functionally distinct tool types as well as Core shaping techniques: (7) hard hammer percussion having to make inferences about tool parts that are missing used, (8) soft hammer percussion used, (9) bipolar hammer from the assemblage. The idea of comparing technologies percussion used, (10) indirect hammer percussion used, (11) based on the number of manufacture steps they contain is pressure hammer percussion used, (12) pecking hammer per- not new (see, e.g., Ambrose 2001; Gowlett 1996) and is similar cussion used to the “cognigrams” method developed by Haidle (Haidle 2009, 2010; Lombard and Haidle 2012). Cognigrams can pro- vide rich insights into the nature of technologies and human Blank Production Techniques behaviors, but their value in comparative research is limited (13) Hard hammer percussion used by the fact that they are not easily quantifiable. (14) Soft hammer percussion used (15) Bipolar hammer percussion used Lithic Procedural Units (16) Indirect hammer percussion used (17) Pressure hammer percussion used We have assembled a list of potential procedural units showing (18) Ochre applied (has traces of ochre) how the complexity of lithic technology can be measured. In Platform treatment: (19) abrasion (platform is rubbed/ accordance with our definition of procedural units, we have abraded), (20) overhang removal (small flakes are removed organized our list of lithic procedural units by reduction steps: from core face adjacent to platform), (21) faceting (small preliminary treatment of raw material, core preparation tech- flakes are removed from the platform) niques, blank production techniques, product shaping, and core rejuvenation. Such division is necessary because the same Product Shaping procedural unit, such as the use of a hard hammer, can serve distinct functions depending on the reduction step for which Edge shaping: (22) retouched edge (edge of final product it is called into action. Each distinct usage should be counted is retouched) S400 Current Anthropology Volume 54, Supplement 8, December 2013

Prehensile modification: (23) backing (a sharp era of final that makes a distinct contribution to the finished form of the product is backed for manual prehension or hafting), (24) product of a particular technology. It is this focus on the notching (final product is notched), (25) tanging (final prod- definition of a procedural unit rather than on its content that uct is tanged) allows for comparison among a wide range of lithic and non- Surface shaping: (26) unifacial retouch (overall morphol- lithic technologies across cultures, time, space, and species. ogy of the blank altered by unifacial flaking), (27) bifacial Many aspects of the way procedural units are counted de- retouch (overall morphology of the blank altered by bifacial pend on the research question asked, such as whether idio- flaking), (28) grinding (ventral/dorsal surface of the final syncratic units, or units that contribute to the decorative as- product is ground) pects of a technology, should be included or excluded from (29) Bulbar thinning: flat flakes removed from bulbar face the analysis. There is no single set of units that applies uni- of the blank versally, so it is important that these decisions be reported in publications in order to increase the replicability of analyses and to facilitate the comparison of published results. Finally, Core Rejuvenation the analysis of technological complexity can be conducted at (30) Tablette: platforms reshaped by removal of single large different scales because the procedural-unit approach can be flakes used to measure the complexity of individual chaıˆnes ope´ra- (31) Outrepasse´e: overpassed/plunging flakes struck inten- toires as well as that of assemblages as a whole. This flexibility tionally to shape distal end of core is useful because there are many archaeological contexts in (32) De´bordante: flakes or blades struck along edge of core which chaıˆnes ope´ratoires cannot be easily reconstructed. to reshape the flaking surface (33) Secondary crest: secondary crest used to reshape core Proof of Concept face (34) Rotation of core: subsequent flakes removed from We have conducted a series of experiments to evaluate the nonopposed platforms validity of the procedural-unit approach to lithic technology (35) Face shaping: core reshaped by lateral flaking between by measuring the complexity of 13 lithic assemblages. More removals or series of removals specifically, we are interested in detecting a temporal trend Using such a list, the complexity of chaıˆnes ope´ratoires can in lithic complexity through the Paleolithic period. We have be translated into a number, the sum of procedural units purposely tried to sample assemblages coming from a wide present in the reduction sequence, that summarizes their com- range of time periods and spatial locations. Our sample thus plexity and that can be compared across cases. includes sites dating from the Lower Paleolithic (Early Stone The list above is by no means exhaustive. Lithic technology Age) to the Upper Paleolithic (Late Stone Age) and ranging can potentially include more procedural units. For instance, from South Africa to Turkey. Given the temporal and spatial in some archaeological contexts it might be useful to also range it covers, our sample is too small to draw any definitive count the use of binding material for composite tools as well conclusion about patterns of complexity during the Paleo- as the type of binding material used (gum, ochre, fat, wax, lithic period. However, it is a useful proof of concept for the resin). The list can also be extended to encompass within the procedural-unit approach because it shows that (1) the same analysis other material and technologies, such as bone method can capture variation in lithic complexity and (2) the tools, ochre pigments, and shell beads. method is sufficiently robust in the face of variation in how In this regard, it is important to recognize that our unit procedural units are defined. of analysis is the procedural unit independent of its content. Different analysts may count procedural units in a lithic In that sense, it does not matter whether a core is shaped by assemblage differently. To examine this problem, we simulated a combination of hard hammer and indirect percussion or the noise that could be generated by different analysts with by a combination of hard and soft hammer: what matters is different views on what constitutes a procedural unit by lump- that in both cases, the shaping of the core involves two pro- ing and splitting the list presented above. In other words, we cedural units. This is analogous to what ecologists do when ask to what extent does the temporal pattern of lithic com- they contrast ecosystems by comparing their species richness, plexity that we can observe in the Paleolithic record depend that is, the total count of species present in each ecosystem. on whether the assemblages have been analyzed by a “splitter” This allows for ecosystems that have few or no species in or by a “lumper.” More saliently, what if a mixture of “lump- common to be compared. Similarly, and similar to Oswalt’s ers” and “splitters” analyzed the assemblages compared, as techno-units, the count of procedural units provides us with would be the case in a data set assembled from various a common measurement unit that allows for the comparison publications? To answer these questions, we counted the pro- of different technologies. This is also why the list above cannot cedural units of the 13 Paleolithic assemblages using two dif- be used as a universal checklist: whether or not an item on ferent lists. First, the assemblages were analyzed using the the list constitutes a procedural unit really depends on extended list presented above. In our view, this is the list that whether it constitutes a mutually exclusive manufacturing step best captures what we mean by procedural units in the context Perreault et al. Measuring the Complexity of Lithic Technology S401 of lithic technology. Second, we produced a second estimate data set thus contains a different mixture of conservative and of the complexity of the assemblages using a shorter version nonconservative analytical decisions simulating the effect of of the above list. This second estimate represents how a con- operator variation on estimates of lithic complexity. Running servative lithic analyst who prefers to err on the side of caution linear regression analysis on these data sets, we find that the might count procedural units. It excludes nine variables that results are affected by this sampling procedure but not so may be deemed ambiguous and too difficult to identify, such profoundly as to disguise the basic patterning. The effect size as heat treatment of raw material or the rotation of the core of age on the count of procedural units across these 1,000 during core rejuvenation. It also lumps 14 procedural units regression analyses ranges from Ϫ0.11 to Ϫ0.38 with an av- into four units. For instance, it lumps together hard hammer, erage of Ϫ0.25, and the adjusted R2’s of these different linear soft hammer, indirect, and pressure flaking into a single cat- models range from 0.13 to 0.73 with an average of 0.39 (fig. egory: “unipolar percussion.” This “conservative” list contains 2). a total of 16 procedural units as opposed to 38 procedural Although there is a general increase in lithic technological units for the “nonconservative” list. complexity over the Pleistocene, the rate of increase in com- Table 1 summarizes our sample of assemblages as well as plexity is greater for the subset of Middle Paleolithic and their complexity relative to the two lists (see supplementary Middle Stone Age assemblages (n p 8 ; see fig. 3). The slope material, available online, for details). The counts of proce- of the best-fit linear model is steeper (β p Ϫ0.46 and Ϫ0.7, dural units are reported for assemblages as a whole rather P p .004 and 0.003 for the conservative and nonconservative than for individual chaıˆnes ope´ratoires. These counts thus rep- counts, respectively), and the variance explained by the model resent the complexity of the tool kits used by the populations is greater (adjustedR 2 p 0.74 and 0.88, respectively). This that produced these assemblages rather than the complexity suggests that the increase in lithic complexity within the Mid- of specific tools they used. Table 1 shows that the procedural- dle Stone Age/Middle Paleolithic might have been more reg- unit approach does capture variation in the complexity of ular than it is across the whole Paleolithic period (thus better lithic technologies. Using the conservative list, the counts of described by a linear model on a log-log scale). This result procedural units range from four, with the Lower Paleolithic contrasts with other studies that have found no general tech- assemblage of Tabun Cave in Israel, to 11, with the Middle nological trends within the Middle Paleolithic period (see de Stone Age assemblage of Klasies River Mouth in South Africa. la Torre 2013; Kuhn 2013). Assuming that this long-term With the nonconservative list, the counts of procedural units trend for increasing complexity through time holds as more range from six, with the two Oldowan assemblages, to 23, assemblages are added to our sample, this analysis is in line with the Mousterian assemblage of Amud Cave. with the view that behavioral complexity increased gradually Figure 1 shows the count of procedural units of the assem- and cumulatively through the Middle Paleolithic/Middle blages plotted against the midpoint of their age on a log-log Stone Age and well before 50 ka (e.g., Brown et al. 2009; scale (the two Oldowan assemblages, A.L. 894 and A.L. 666, d’Errico and Henshilwood 2007; d’Errico et al. 2005; Marean are plotted against their terminus ante quem date, 12.35 mya). et al. 2007; McBrearty and Brooks 2000). A linear regression analysis in which the dependent variable Overall, our analysis suggests that trends such as the in- is the logarithm of the count of procedural units and the crease of lithic complexity through the Paleolithic period may independent variable is the logarithm of the midpoint of the be detected in a robust manner even in the face of divergence age of the assemblage suggests that the complexity of lithic between analysts in the definition of procedural units. Even technologies increases steadily through time. The slope β of though our results are preliminary, we find it intriguing that the best-fit linear model for the nonconservative count of the complexity of these lithic assemblages align along the same procedural units is Ϫ0.314 (P ! .001 ), which means that the trend line even though they were produced over thousands size of the procedural-unit inventory shrinks by about 3% as of years by different hominid species coming from various the age of the material increases by 10%. The model explains parts of Africa and Eurasia and given the presumed sensitivity 68% of the variance (adjustedR 2 p 0.68 ). The relationship of lithic technology to environmental, demographic, behav- between age and lithic complexity is also detectable when the ioral, economical, and cognitive factors. conservative view of procedural units is adopted, although it p Ϫ is not as strong: β 0.184 (a decrease in complexity of Discussion and Conclusion about 2% per 10% increase in age;P ! .005 , adjusted R 2 p 0.49). These results suggest that the temporal pattern of in- Our main argument in this paper is that the complexity of crease in complexity of lithic technology through time is lithic technology can be measured by counting the procedural strong enough to be detected by both conservative lumpers units, that is, the mutually exclusive manufacturing steps that and nonconservative splitters. But what if both conservative contribute to the finished form of the technology. Because it and nonconservative estimates were combined in the same is quantitative and independent of any specific technology, analysis? To test this possibility, we created 1,000 data sets by time period, region, or culture, this approach can also be selecting randomly, for each assemblage, a count of procedural applied to nonlithic technologies and could help solve many units from the conservative or the nonconservative view. Each of the issues associated with previous attempts at measuring Table 1. Count of procedural units of 13 Paleolithic assemblages

Count of procedural units Conservative Nonconservative Assemblage Period Age list list References A.L. 894, Ethiopia Lower Paleolithic (Oldowan) 12.35 mya 5 6 Hovers 2009a; E. Hovers and A. David- szon, unpublished manuscript A.L. 666, Ethiopia Lower Paleolithic (Oldowan) 12.35 mya 5 6 E. Hovers and A. Davidszon, unpublished manuscript; Kimbel et al. 1996 GnJh-03, Kenya Early Stone Age (Acheulean) ca. 500–284 ka 8 14 Cornelissen 1992; Leakey et al. 1969; McBrearty 1999; Tryon, McBrearty, and Texier 2005 Tabun Cave, Unit XI-XIV, Israel Early Paleolithic ca. 450–225 ka 4 7 Yanmburgaz Cave, Turkey Early Middle Paleolithic 275 ka 6 7 Kuhn, Arsebu¨k, and Howell 1996 Tabun Cave, Unit IX, Israel Early Middle Paleolithic (Mousterian) ca. 225–100 ka 6 13 R. Shimelmitz and S. L. Kuhn, unpublished manuscript GhJh-74, Kenya Middle Stone Age ca. 200 ka 5 11 Tryon 2003, 2006 Klasies River Mouth, Klasies stage, South Africa Middle Stone Age 115–100 ka 9 19 Wurz 2002a,2002b Qafzeh Cave (layers XXIV-III), Israel Middle Paleolithic (Mousterian) 92 ka 10 19 Hovers 2009b; Hovers and Raveh 2000 Klasies River Mouth, Mossel Bay stage, South Middle Stone Age 100–80 ka 9 15 Wurz 2002a,2002b Africa Amud Cave B4-B1, Israel Middle Paleolithic (Mousterian) 68–55 ka 11 23 Alperson-Afil and Hovers 2005; Ekshtain 2006; Goder 1997; Hovers 1998, 2007; Hovers et al. 2011 Klasies River Mouth, Howieson Poort stage, Middle Stone Age 62–58 ka 9 22 Wurz 2002a,2002b South Africa U¨ c¸ag˘ızlı Cave, Ahmarian layers B, B1-3, C, Turkey Upper Paleolithic ca. 35–32 ka 9 15 Kuhn et al. 2009 Perreault et al. Measuring the Complexity of Lithic Technology S403

Figure 1. “Conservative” and “nonconservative” count of procedural units for 13 Paleolithic assemblages (see table 1) plotted against their age. Note the logarithmic scale (base 10) on both axes. Assemblages with a range of ages are plotted against the midpoint of the age range. The line represents the best-fit linear model using least squares regression (conservative: β p Ϫ.184, standard error [SE] p 0.05,P p .005 ,n p 13 , adjustedR2 p 0.49 ; nonconservative: B p β p Ϫ.314, SE p 0.06 ,P p .0003 ,n p 13 , adjusted R2 p 0.68). The dashed lines represent the 95% confidence bands, and the dotted lines represent the 95% prediction bands. complexity in the archaeological record. As such, this method semblages will allow us to verify this finding and compare will allow us to revisit several claims about the complexity of the long-term rates of change in technological complexity Paleolithic technologies. For instance, do Clark’s technological between Europe, Africa, and Asia as well as between different modes really represent an increase in complexity (Foley and periods of the Paleolithic. Lahr 2003)? Are Upper Paleolithic tools more complex than Nonetheless, there are several caveats to the procedural- Middle Paleolithic ones (Mellars 1989), or is the Levallois unit approach. For instance, it fails to capture some aspects method more complex than the production of blades from of what is commonly meant by “technological complexity,” prismatic cores (Bar-Yosef and Kuhn 1999)? Our preliminary such as the level of skills involved in the manufacturing of analysis suggests the existence of a long-term trend toward tools or the complexity that emerges from the hierarchical greater complexity in the evolution of lithic technologies organization of some manufacturing procedures (Byrne throughout the Paleolithic period but that there is a partic- 2007). A second caveat is that the procedural-unit approach ularly sharp increase in complexity within the Middle Pa- is not completely impervious to taphonomic issues. For in- leolithic and the Middle Stone Age. A larger sample of as- stance, the complexity of lithic technologies could plateau at

Figure 2. Relative frequency histogram of the effect size and variance explained of the best-fit linear model using least squares regression calculated over 1,000 data sets generated by randomly selecting for each of the 13 assemblages either the conservative or the nonconservative count of procedural units (see table 1). S404 Current Anthropology Volume 54, Supplement 8, December 2013

Figure 3. “Conservative” and “nonconservative” count of procedural units for eight Middle Paleolithic/Middle Stone Age assemblages plotted against their age. Note the logarithmic scale (base 10) on both axes. Assemblages with a range of ages are plotted against the midpoint of the age range. The line represents the best-fit linear model using least squares regression (conservative: β p Ϫ.46, standard error[SE] p 0.1 ,P p .004 ,n p 8 , adjustedR2 p 0.74 ; nonconservative: B p β p Ϫ.70, SE p 0.1 ,P p .0003 ,n p 13 , adjustedR2 p 0.88 ). The dashed lines represent the 95% confidence bands, and the dotted lines represent the 95% prediction bands. certain points in time while the complexity of other tech- quently discussed in the archaeological literature, it would be nological parts made of perishable material may continue to useful to verify the reliability of the approach by measuring increase. This means that the procedural-unit approach can the level of agreement between different operators and ex- be biased by false negatives and as such provides us only with perimental context. But the problem of subjectivity can also a lower bound for technological complexity. Moreover, dif- be addressed statistically. By analyzing large samples we can ferent classes of material, such as lithic and ceramic, can also average out the noise created by factors that are independent be subject to different taphonomic processes and therefore of the variable of interest, such as operator errors, between- be less amenable to comparison. And like other measures of operator disagreements, or differential preservation. This is richness in the archaeological record (e.g., Cannon 2001; not an attempt to avoid the issue of subjectivity in defining Cochrane 2003; Meltzer, Leonard, and Stratton 1992; Rhode procedural units. On the contrary, the problem of noise in 1988), the count of procedural units is likely dependent on empirical data is not specific to the procedural-unit approach: sample size. it is a problem that every scientific discipline faces, and the Another source of problems is the effect of subjective in- most powerful way we have to deal with it is to collect larger p ͱ ferences on the count of procedural units. Some procedural samples. The standard error of the mean,Sx¯ s / n , where units are harder than others to identify in an archaeological s is the standard deviation of the sample and n the size of assemblage, and not every analyst will be comfortable with the sample, is a good measure of the effect of errors on the the inferential leap required to mark them as “present”—the mean of a sample.1 Because the standard error of the mean use of a soft hammer versus indirect percussion comes to decreases in proportion to the square root of sample size, for mind here. Other operator errors can affect the replicability any given amount of error in a sample there will always be of the method, such as disagreement between operators on a sample size that is large enough to estimate accurately the whether something like rotating the core during core reju- venation constitutes a procedural unit or not. As such, the 1. In an ideal and purely mechanistic world,y p x , where y is the procedural-unit approach to lithic complexity suffers from variable of interest (e.g., lithic complexity) and x is the predictor (e.g., p ϩ the same kinds of subjectivity that prevail in lithic analysis in the age of assemblages). But we live in a world wherey x ε , where ε is the noise in the data. In most cases this error will be random with general. respect to x; some of us will tend to overestimate the number of pro- Our proof-of-concept study, however, does provide ten- cedural units present in assemblages while others will tend to underes- tative evidence that the method is sufficiently robust to sub- timate it. The standard error of the mean, a measure of the effects of jectivity in the definition of lithic procedural units. We were errors on the mean of a sample, is used to calculate the confidence interval able to detect a temporal trend in the complexity of Paleolithic within which the true mean of a population lies. For example, if the average complexity in a sample of Middle Stone Age assemblages of size lithic technology even when the list of procedural units ex- n is x with a standard deviation of s, then there is a 95% chance that amined was reduced by 54%. Even though the procedural the true mean complexity in the Middle Stone Age lies within the interval .(s /ͱn)1.96 ע units discussed in this paper are uncontroversial and fre- defined by x Perreault et al. Measuring the Complexity of Lithic Technology S405 true mean complexity of a sample. The same line of reasoning de Beaune, Sophie A. 2004. The invention of technology: prehistory and cognition. Current Anthropology 45(2):139–162. can be extended to regression analysis. In sum, the influence de la Torre, Ignacio, Jorge Martı´nez-Moreno, and Rafael Mora. 2013. Change of operator errors can be dealt with statistically. This allows and stasis in the Iberian Middle Paleolithic: considerations on the signifi- for the procedural-unit approach to remain a useful metric cance of Mousterian technological variability. Current Anthropology 54(suppl. 8):S320–S336. by which the complexity of prehistoric technologies can be d’Errico, Francesco, and Christopher Henshilwood. 2007. Additional evidence quantified and compared and to test a wide range of hy- for bone technology in the southern African Middle Stone Age. Journal of potheses about what drives changes in technological com- Human Evolution 52(2):142–163. d’Errico, Francesco, Christopher Henshilwood, Marian Vanhaeren, and Karen plexity. van Niekerk. 2005. Nassarius kraussianus shell beads from Blombos Cave: evidence for symbolic behaviour in the Middle Stone Age. Journal of Human Evolution 48(1):3–24. Ekshtain, R. 2006. A comparative study of knapping accidents in the Levantine Mousterian. Jerusalem: Hebrew University of Jerusalem. Acknowledgments Foley, Robert. 1987. Another unique species. New York: Longman Scientific & Technical. We thank Christian Tryon and Erella Hovers for providing Foley, Robert, and Marta Mirazo´n Lahr. 2003. On stony ground: lithic tech- us with their data. We also thank the Wenner-Gren Foun- nology, human evolution, and the emergence of culture. Evolutionary An- dation, Leslie Aiello, the participants of the Wenner-Gren thropology: Issues, News, and Reviews 12(3):109–122. Goder, M. 1997. Technological and typological aspects of Layer B1 in the Late symposium no. 145, and the reviewers for their comments Mousterian site at Amud Cave. MA thesis, Hebrew University of Jerusalem. and criticisms of our paper. Gowlett, John. 1996. The frameworks of early hominid social systems: how many useful parameters of archaeological evidence can we isolate? In The archaeology of human ancestry: power, sex and tradition. J. Steele and S. References Cited Shennan, eds. Pp. 135–183. London: Routledge. Haidle, Miriam Noe¨l. 2009. How to think a simple spear. In Cognitive ar- Alperson-Afil, N., and Erella Hovers. 2005. Differential use of space at the chaeology and human evolution. S. de Beaune, F. L. Coolidge, and T. Wynn, Neandertal site of Amud Cave, Israel. Eurasian Prehistory 3:3–22. eds. Pp. 57–73. Cambridge: Cambridge University Press. Ambrose, Stanley H. 2001. Paleolithic technology and human evolution. Sci- ———. 2010. Working-memory capacity and the evolution of modern cog- ence 291(5509):1748–1753. nitive potential. Current Anthropology 51(suppl. 1):S149–S166. ———. 2010. Coevolution of composite-tool technology, constructive mem- Hovers, Erella. 1998. The lithic assemblages of Amud Cave: implications for ory, and language. Current Anthropology 51(suppl. 1):S135–S147. the end of the Mousterian in the Levant. In Neanderthals and modern Bar-Yosef, O., and S. L. Kuhn. 1999. The big deal about blades: laminar humans in Western Asia. T. Akazawa, K. Aoki, and O. Bar-Yosef, eds. Pp. technologies and human evolution. American Anthropologist 101(2):322– 143–163. New York: Plenum. 338. ———. 2007. The many faces of cores-on-flakes: a perspective from the Beatty, J. 1995. The evolutionary contingency thesis. In Concepts, theories, and Levantine Mousterian. In Cores or tools? alternative approaches to stone tool rationality in the biological sciences. G. Wolters and J. G. Lennox, eds. Pp. analysis. S. P. McPherron, ed. Pp. 42–74. Cambridge: Cambridge Scholars. 45–81. Pittsburgh, PA: University of Pittsburgh Press. ———. 2009a. Learning from mistakes: flaking accidents and knapping skills Bleed, Peter. 1986. The optimal design of hunting weapons: maintainability in the assemblage of A. L. 894 (Hadar, Ethiopia). In The cutting edge: new or reliability. American Antiquity 51(4):737–747. approaches to the archaeology of human origins. K. Schick and N. Toth, eds. Bonner, John Tyler. 1988. The evolution of complexity by means of natural Pp. 137–150. Gosport, IN: Stone Age Institute. selection. Princeton, NJ: Princeton University Press. ———. 2009b. The lithic assemblages of Qafzeh Cave. New York: Oxford Bousman, C. Britt. 1993. Hunter-gatherer adaptations, economic risk and tool University Press. design. Lithic Technology 18(1/2):59–86. Hovers, Erella, A. Malinsky-Buler, M. Goder-Goldberger, and R. Ekshtain. Brantingham, P. Jeffrey, John W. Olsen, Jason A. Rech, and Andrei I. Kri- 2011. Capturing a moment: identifying short-lived activity locations in voshapkin. 2000. Raw material quality and prepared core technologies in Amud Cave, Israel. In The Lower and Middle Palaeolithic in the Middle East northeast Asia. Journal of Archaeological Science 27(3):255–271. and neighbouring regions. J.-M. Le Tensorer, R. Jagher, and M. Otte, eds. Brown, Kyle S., Curtis W. Marean, Andy I. R. Herries, Zenobia Jacobs, Chantal Pp. 110–114. Lie`ge: University of Lie`ge Press. Tribolo, David Braun, David L. Roberts, Michael C. Meyers, and Jocelyn Hovers, Erella, and A. Raveh. 2000. The use of a multivariate display technique Bernatchez. 2009. Fire as an engineering tool of early modern humans. in the analysis of inter-assemblage lithic variability: a case study from Qafzeh Science 325(5942):859–862. Cave, Israel. Journal of Archaeological Science 27(11):1023–1038. Byrne, R. W. 2007. Culture in great apes: using intricate complexity in feeding Kelly, Robert L. 1995. The foraging spectrum: diversity in hunter-gatherer life- skills to trace the evolutionary origin of human technical prowess. Philo- sophical Transactions of the Royal Society B 362(1480):577–585. ways. Washington, DC: Smithsonian Institution. Cannon, Michael D. 2001. Archaeofaunal relative abundance, sample size, and Kimbel, W. H., R. C. Walter, D. C. Johanson, K. E. Reed, J. E. Aronson, Z. statistical methods. Journal of Archaeological Science 28(2):185–195. Assefa, C. W. Marean, et al. 1996. Late Pliocene Homo and Oldowan stone Chaitin, G. J. 1970. On the difficulty of computations. IEEE Transactions on tools from the Hadar Formation (Kada Hadar Member), Ethiopia. Journal Information Theory IT-16:5–9. of Human Evolution 31:549–561. Cochrane, Grant W. G. 2003. Artefact attribute richness and sample size Kline, Michelle A., and Robert Boyd. 2010. Population size predicts tech- adequacy. Journal of Archaeological Science 30(7):837–848. nological complexity in Oceania. Proceedings of the Royal Society B 277 Collard, Mark, Briggs Buchanan, Jesse Morin, and Andre Costopoulos. 2011. (1693):2559–2564. What drives the evolution of hunter-gatherer subsistence technology? a Kuhn, Steven L. 2013. Roots of the Middle Paleolithic in Eurasia. Current reanalysis of the risk hypothesis with data from the Pacific Northwest. Anthropology 54(suppl. 8):S255–S268. Philosophical Transactions of the Royal Society B 366(1567):1129–1138. Kuhn, Steven L., Gu¨ven Arsebu¨k, and F. Clark Howell. 1996. The Middle Collard, Mark, Michael Kemery, and Samantha Banks. 2005. Causes of toolkit Pleistocene lithic assemblage from Yarimburgaz Cave, Turkey. Pale´orient variation among hunter-gatherers: a test of four competing hypotheses. 22(1):31–49. Canadian Journal of Archaeology 29:1–19. Kuhn, Steven L., Mary C. Stiner, Gu¨lec¸ Erksin, Ismail O¨ zer, Hakan Yılmaz, Coolidge, Frederick L., and Thomas Wynn. 2009. The rise of Homo sapiens: Ismail Baykara, Ays¸en Ac¸ıkkol, et al. 2009. The Early Upper Paleolithic the evolution of modern thinking. Malden, MA: Wiley-Blackwell. occupations at U¨ c¸ag˘ızlı Cave (Hatay, Turkey). Journal of Human Evolution Cornelissen, E. 1992. Site GnJh-17 and its implications for the archaeology of 56:87–113. the Middle Kapthurin Formation, Baringo, Kenya, vol. 133 of Annales, sciences Leakey, M., P. V. Tobias, J. E. Martyn, and R. E. F. Leakey. 1969. An Acheulian humaines. Tervuren: Muse´e Royale de l’Afrique Centrale. industry with prepared core technique and the discovery of a contemporary S406 Current Anthropology Volume 54, Supplement 8, December 2013

hominid mandible at Lake Baringo, Kenya. Proceedings of the Prehistoric Richerson, Peter J., and Robert Boyd. 2005. Not by genes alone: how culture Society 3:48–76. transformed human evolution. Chicago: University of Chicago Press. Lombard, Marlize, and Miriam Noe¨l Haidle. 2012. Thinking a bow-and-arrow Sambrook, T., and A. Whiten. 1997. On the nature of complexity in cognitive set: cognitive implications of Middle Stone Age bow and stone-tipped arrow and behavioural science. Theory and Psychology 7(2):191–213. technology. Cambridge Archaeological Journal 22(2):237–264. Schick, Kathy D. 1994. The Movius Line reconsidered: perspectives on the Marean, C. W., Miryam Bar-Matthews, Jocelyn Bernatchez, Eric Fisher, Paul earlier Paleolithic of eastern Asia. In Integrative paths to the past: paleo- Goldberg, Andy I. R. Herries, Zenobia Jacobs, et al. 2007. Early human use anthropological advances in honor of F. Clark Howell. R. S. Corruccini and of marine resources and pigment in South Africa during the Middle Pleis- R. L. Ciochon, eds. Pp. 569–596. Englewood Cliffs, NJ: Prentice Hall. tocene. Nature 449(7164):905–908. Shannon, C. E., and W. Weaver. 1949. The mathematical theory of commu- McBrearty, Sally. 1999. The archaeology of the Kapthurin Formation. In Late nication. Urbana: University of Illinois Press. Cenozoic environments and hominid evolution: a tribute to Bill Bishop.P. Shea, John J., and Matthew L. Sisk. 2010. Complex projectile technology and Andrews and P. Banham, eds. Pp. 143–156. London: Geological Society. Homo sapiens dispersal into western Eurasia. Paleoanthropology (2010):100– McBrearty, Sally, and Alison S. Brooks. 2000. The revolution that wasn’t: a 122. new interpretation of the origin of modern human behavior. Journal of Shennan, S. 2001. Demography and cultural innovation: a model and its Human Evolution 39(5):453–563. implications for the emergence of modern human culture. Cambridge Ar- Mellars, Paul. 1989. Technological changes across the Middle-Upper Palaeo- chaeological Journal 11(1):5–16. lithic transition: economic, social and cognitive perspectives. In The human revolution: behavioural and biological perspectives on the origins of modern Shott, Michael J. 1986. Technological organization and settlement mobility: humans. P. Mellars and C. Stringer, eds. Pp. 339–365. Princeton, NJ: Prince- an ethnographic examination. Journal of Anthropological Research 42:15– ton University Press. 51. ———. 2006a. Going east: new genetic and archaeological perspectives on Torrence, R. 1983. Time budgeting and hunter-gatherer technology. In Hunter- the modern human colonization of Eurasia. Science 313(5788):796–800. gatherer economy in prehistory: a European perspective. G. Bailey, ed. Pp. 11– ———. 2006b. Why did modern human populations disperse from Africa 22. Cambridge: Cambridge University Press. ca. 60,000 years ago? a new model. Proceedings of the National Academy of Torrence, Robin. 1989. Re-tooling: towards a behavioral theory of stone tools. Sciences, U.S.A. 103(45):9381–9386. In Time, energy and stone tools. R. Torrence, ed. Pp. 57–66. Cambridge: Meltzer, David J., Robert D. Leonard, and Susan K. Stratton. 1992. The re- Cambridge University Press. lationship between sample size and diversity in archaeological assemblages. ———. 2000. Hunter-gatherer technology: macro- and microscale ap- Journal of Archaeological Science 19(4):375–387. proaches. In Hunter-gatherers: an interdisciplinary perspective. C. Panter- Movius, Hallam, L. 1944. Early man and Pleistocene stratigraphy in southern Brick, R. H. Layton, and P. Rowley-Conwy, eds. Pp. 99–143. Cambridge: and eastern Asia. Papers of the Peabody Museum of Archaeology and Eth- Cambridge University Press. nology, vol. 19. Cambridge, MA: Peabody Museum of American Archae- Tryon, Christian A. 2003. The Acheulian to Middle Stone Age transition: ology and Ethnology. tephrostratigraphic context for archaeological change in the Kapthurin For- ———. 1948. The Lower Paleolithic cultures of southern and eastern Asia. mation. PhD dissertation, University of Connecticut, Storrs. Transactions of the American Philosophical Society 38:329–420. ———. 2006. “Early” Middle Stone Age lithic technology of the Kapthurin Osborne, A. J. 1999. From global models to regional patterns: possible de- Formation (Kenya). Current Anthropology 47:367–375. terminants of Folsom hunting weapon design diversity and complexity. In Tryon, Christian A., Sally McBrearty, and P. J. Texier. 2005. Levallois lithic Folsom lithic technology: explorations in structure and variation. D. S. Amick, technology from the Kapthurin Formation, Kenya: Acheulian origin and ed. Pp. 188–213. Ann Arbor, MI: International Monographs in Prehistory. Middle Stone Age diversity. African Archaeological Review 22(4):199–229. Oswalt, Wendell H. 1976. An anthropological analysis of food-getting technology. Wadley, Lyn. 2010. Compound-adhesive manufacture as a behavioral proxy New York: Wiley-Interscience. Pope, G. G. 1989. Bamboo and human evolution. Natural History 10:48–57. for complex cognition in the Middle Stone Age. Current Anthropology Powell, Adam, Stephen Shennan, and Mark G. Thomas. 2009. Late Pleistocene 51(suppl. 1):S111–S119. demography and the appearance of modern human behavior. Science Whiten, A., J. Goodall, W. C. McGrew, T. Nishida, V. Reynolds, Y. Sugiyama, 324(5932):1298–1301. C. E. G. Tutin, R. W. Wrangham, and C. Boesch. 1999. Cultures in chim- Premo, L. S., and Steven L. Kuhn. 2010. Modeling effects of local extinctions panzees. Nature 399(6737):682–685. on culture change and diversity in the Paleolithic. PLoS ONE 5(12):e15582. Wurz, Sarah. 2002a. The Middle Stone Age sequence at Klasies River, South Reiter, Paul. 2000. From Shakespeare to Defoe: malaria in England in the Africa. PhD dissertation, University of Stellenbosch. Little Ice Age. Emerging Infection Diseases 6(1):1–11. ———. 2002b. Variability in the Middle Stone Age lithic sequence, 115,000– Rhode, David. 1988. Measurement of archaeological diversity and the sample- 60,000 years ago at Klasies River, South Africa. Journal of Archaeological size effect. American Antiquity 53(4):708–716. Science 29(9):1001–1015.