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Neandertals and Modern Humans in Western Asia

Neandertals and Modern Humans in Western Asia Edited by

Takeru Akazawa International Research Center for Japanese Studies Kyoto, Japan

Kenichi Aoki University of Tokyo Tokyo, Japan

and

Ofer Bar-Yosef Harvard University Cambridge, Massachusetts

KLUWER ACADEMIC PUBLISHERS NEW YORK, BOSTON, DORDRECHT, LONDON, MOSCOW

H%RRN ,6%1 3ULQW,6%1

0-306-47153-1 59248

‹.OXZHU$FDGHPLF3XEOLVKHUV 1HZ95% of the globe/population, although what happened in western Europe probably reflects patterns in perhaps many other currently more poorly sampled parts of the Old World Returning to genetics, let us imagine taking a time machine and making a sampling

trip around the world around 45 ka ago. We would find “modern” populations throughout Africa (based on the admittedly relatively small portions of northwestern, eastern, and southern Africa that have been sampled), Southwest Asia, Australia, and probably some regions between eastern Africa/Southwest Asia and Australia. But in Europe and in many parts of South, East, and Southeast Asia we would find only nonmodern populations. The recent analysis of partial mtDNA of the Neandertal type specimen (Krings et al. 1997), although only a sample of one (and likely to remain so, Cooper et al. 1997), suggests that at least the Neandertal nonmodern population was genetically quite different from modern humans, at least as different as the central and eastern subspecies of chimpanzee are from each other (Pan troglodytes troglodytes and Pan troglodytes schweinfurthii) (Morin et al. 1994; Goldberg and Ruvolo 1997; Krings et al. 1997). That is, the living human populations (and their “modern” antecedents) would be equivalent genetically to one chimpanzee

subspecies (actually, part of one), while the various archaic human populations might have been equivalent genetically to other chimpanzee subspecies.

526

D. Pilbeam

Using morphological data, there are problems in comparing the degree and patterning of within-species variation between species, but enough has been published now to suggest that the cranial differences between human modern and nonmodern populations (whether or

not they are described as species or subspecies) were greater than those between subspecies of the chimpanzee or even the gorilla (Groves 1970; Van Vark 1984; Groves et al. 1992; Turbon et al. 1997). So, apparently paradoxically, we see genetic differences between moderns and nonmoderns (specifically Neandertals) equivalent to those between chimpanzee subspecies (implying equivalent time depths for the distinct clades) (Morin et al. 1994; Krings et al. 1997), whereas the morphological differences are (perhaps considerably) greater implying greater adaptive differences between the hominid populations than between the ape subspecies. Arguing over whether these are or are not species distinctions in either the hominids or the apes is pointless in my opinion, because these are matters of taste and taxonomic philosophy. (But this hominid case presents us with another example of the occasionally marked disjunction between “morphological” and “genetic” differences.) If we now consider the structure of arguments about modern human origins, although there is still a continuum from “regional continuity” to “replacement” models, with just about all intermediate positions, most would now be clustered toward the “replacement” end of the spectrum. Another interesting and increasingly discussed set of models are those that cover the behavior of these hominids. Were the Neandertals and other nonmoderns “just like us” in their behavioral potential; that is, did their biology allow the full living human range of “cultural” responses? Or was their behavioral capacity different, in that their “biology” was more constraining of behavioral potential, of “culture”? As behavior becomes biologically less constrained (that is, more like living humans) the behavioral repertoire can become more varied. But not all populations of equivalent biologically determined behavioral potential have the same set of adaptive-technological-ecological responses. To illustrate this, a good example would be the adaptive range of human behavior around 7 ka ago when some populations were sedentary agriculturalists, some were hunter-gatherers, and some pastoralists. Another example would be around 35 ka ago when most modern human populations across the globe would not have produced Upper Paleolithic cultures as defined in Europe and western Asia. Yet all these populations were equally human. The critical issue here is the nature of behavioral potential (“cultural capabilities”) in Neandertals and other nonmodern populations versus that (or those) in the various temporal stages of modern humans. The same or different? Were the differences equivalent to those between, say, Near Eastern agriculturalists and European late Paleolithic hunters or between European Upper Paleolithics and North African Aterians? That is, was the degree of biological-constraint on cultural behavior in Neandertals equivalent to that in modern humans? Or were these hominids different in ways analogous to, for example, “common” chimpanzees (Pan troglodytes) and “pygmy chimpanzees” or bonobos (Pan paniscus)? That is, were the differences more “biologically” based? If Neandertals and moderns were

indeed behaviorally different because of some “biological” difference, were they nonetheless equivalent in behavioral complexity (or grade) as are the two chimpanzee species, or chimps and gorillas? Or did they differ in grade, one being more “advanced” than the other, as we see in chimps versus baboons, for example? And what, if any, relevance do these issues have to do with “replacement”? (After all, chimps have “replaced” neither gorillas nor baboons.) Clearly the anatomical and archeological evidence is sufficiently ambiguous that essentially all possible combinations of these model features can be proposed at the moment by competent scholars. With that said though, I think there have been changes over the past decade that have moved the debate a little toward resolution. We have evidence for the youth and unity of living humans. Human genetic patterning, plus the one genetic fossil datum from

Afterword

527

the Neandertal type specimen, together with numerous morphological and morphometric studies, suggest that genetically and morphologically living humans are a descendant remnant of what was relatively recently (40 ka ago or even younger) a much more diverse cluster of populations, genetically and morphologically. Without more and different (nuclear as well as mitochondrial DNA) fossil genetic data (which will be difficult indeed to obtain), we shall have to rely on continuing analysis of living populations, but there is little if any sign of the existence of “ancient” lineages of the kind found in nonhuman hominoids (Ruvolo 1997). This is evidence for a “replacement” model. (Or, expressing this conservatively, in the one well-sampled, peripheral, and rather small area—western Europe—for which there is adequate fossil and archeological data, replacement is the more plausible option.). This still leaves us with some quite major questions to answer. Picking just three as examples: the pattern of evolution from nonmodern to modern morphologies and behaviors away from the periphery; the nature and timing of the behavioral transition to the current strongly culturally inflected biology from one in which the biological constraints on behavioral variation and change were less humanlike; and the dynamics of Eurasian population interactions between 45 ka and 25 ka ago. But a great deal of progress is being made.

REFERENCES Barbujani, G., A. Magagni, E. Minch, and L. L. Cavalli-Sforza. 1997. An apportionment of human DNA diversity. Proceedings of the National Academy of Science USA 94:4516–19. Brauer, G., and K. Rimbach. 1990. Late archaic and modem Homo sapiens from Europe, Africa, and Southwest

Asia: Craniometric comparisons and phylogenetic implications. Journal of Human Evolution 19:789–807. Cavalli-Sforza, L., P. Menozzi, and A. Piazza. 1994. The History and Geography of Human Genes. Princeton: Princeton University Press. Cooper, A., H. N. Poinar, S. Pääbo, J. Radovcic, A. Debénath, M. Caparros, C. Barroso-Ruiz, J. Bertranpetit, C. Nielsen-Marsh, R. E. M. Hedges, and B. Sykes. 1997. Neandertal genetics. Science 277:1021–4. Goldberg, T., and M. Ruvolo. 1997. Molecular phylogenetics and historical biogeography of east African chimpanzees. Biological Journal of the Linnean Society 61:301–24. Goldstein, D. B., A. Ruiz Linares, L. L. Cavalli-Sforza, and M. W. Feldman. 1995. Genetic absolute dating based on microsatellites and the origin of modern humans. Proceedings of the National Academy of Science USA

92:6723–7. Groves, C. 1970. Population systematics of the gorilla. Journal of the Zoological Society of London 161:287–300.

Groves, C., C. Westwood, and B. T. Shea. 1992. Unfinished business: Mahalanobis and a clockwork orang. Journal of Human Evolution 22:327–40.

Holliday, T. 1997. Body proportions in Late Pleistocene Europe and modern human origins. Journal of Human Evolution 32:423–47. Krings, M., A. Stone, R. W. Schmitz, H. Krainitzki, M. Stoneking, and S. Pääbo. 1997. Neandertal DNA sequences and the origin of modern humans. Cell 90:19–30. Lewontin, R. 1972. The apportionment of human diversity. Evolutionary Biology 6:381–98. Morin, P. A., J. J. Moore, R. Chakraborty, L. Jin, J. Goodall, and D. S. Woodruff. 1994. Kin selection, social structure, gene flow, and the evolution of chimpanzees. Science 265:1193–1201. Penny, D., M. Steel, P. J. Waddell, and M. D. Hendy. 1995. Improved analyses of human mtDNA sequences support a recent African origin for Homo sapiens. Molecular Biology and Evolution 12:863–82.

Ruvolo, M. 1996. A new approach to studying modern human origins: Hypothesis testing with coalescence time distributions. Molecular Phylogenetics and Evolution 5:202-19. Ruvolo, M. 1997. Genetic diversity in hominoid primates. Annual Revue of Anthropology 26:515–40.

Turbon, D., A. Pérez-Pérez, and C. Stringer. 1997. A multivariate analysis of Pleistocene hominids: Testing hypotheses of European origins. Journal of Human Evolution 32:449–68. Van Vark, G. 1984. On the determination of hominid affinities, 323–50. In Multivariate Statistical Methods in Physical Anthropology, ed. G. van Vark and W. W. Howells. Boston: Reidel.

Zischler, H., H. Geisert, A. von Haeseler, and S. Pääbo. 1995. A nuclear “fossil” of the mitochondrial D-loop and the origin of modern humans. Nature 378:489–92.

CONTRIBUTORS

Takeru Akazawa International Research Center for Japanese Studies Kyoto 610-11, Japan Stanley H. Ambrose Department of Anthropology University of Illinois Urbana, Illinois 61801 Kenichi Aoki Department of Biological Sciences

University of Tokyo Tokyo 113-0033, Japan Baruch Arensburg Department of Anatomy and Anthropology Sackler School of Medicine Tel Aviv University Ramat Aviv 69978, Israel

Steven E. Churchill Department of Biological Anthropology and Anatomy Duke University Durham, North Carolina 27708 Jacques Connan Elf Exploration Production Direction Exploration, CSTJF Avenue Larribau, PAU 64018, France Yukio Dodo Department of Anatomy Tohoku University School of Medicine Sendai 980, Japan Paul Goldberg Department of Archaeology Boston University Boston, Massachusetts 02215

Ofer Bar-Yosef Department of Anthropology Peabody Museum Harvard University Cambridge, Massachusetts 02138

Naama Goren-Inbar Institute of Archaeology Hebrew University Mt. Scopus, Jerusalem 91905, Israel

Anna Belfer-Cohen Institute of Archaeology Hebrew University Mt. Scopus, Jerusalem 91905, Israel

Donald O. Henry Department of Anthropology University of Tulsa Tulsa, Oklahoma 74104

Eric Boëda Department of Ethnology and Prehistory Université de Paris X Nanterre 92001, France

Erella Hovers Institute of Archaeology Hebrew University Mt. Scopus, Jerusalem 91905, Israel 529

530

Contributors

F. Clark Howell

Paul Mellars

Laboratory for Human Evolutionary

Department of Archaeology Cambridge CB2 3DZ, United Kingdom

Studies/Museum of Vertebrate Zoology University of California Berkeley, California 94720 Jean-Jacques Hublin UMR 152 du CNRS Laboratoire d’Anthropologie Musée de l’Homme Paris 75116, France

Jean-Louis Joron Groupe des Sciences de la Terre Laboratoire Pierre Süe CEN Saclay Gif sur Yvette Cedex 91191, France

Richard G. Klein Department of Anthropology Stanford University Stanford, California 94305 Osamu Kondo Department of Anatomy Tohoku University School of Medicine Sendai 980, Japan Janusz K. Kozlowski Institute of Archaeology Jagellonian University

Kraków 31007, Poland Kevin N. Laland Sub-Department of Animal Behaviour University of Cambridge Cambridge CB3 8AA, United Kingdom Daniel E. Lieberman Department of Anthropology The George Washington University Washington, DC 20052 Liliane Meignen ERA 28 du Centre de Recherches Archeologiques CNRS, Sophia Antipolis Valbonne 06560, France

Norbert Mercier Centre de Faibles Radioactivites Laboratoire Mixte CNRS-CEA Gif sur Yvette Cedex 91198, France Nancy Minugh-Purvis Department of Neurobiology and Anatomy Allegheny University of the Health

Sciences Philadelphia, Pennsylvania 19129 University Museum of Archaeology and Anthropology University of Pennsylvania Philadelphia, Pennsylvania 19104 Division of Plastic Surgery The Children’s Hospital of Philadelphia Philadelphia, Pennsylvania 19104 Sultan Muheson Director-General of Antiquities and Museums Damascus, Syrian Arab Republic Marcel Otte Service de Préhistoire University of Liege Liège B-4000, Belgium

David Pilbeam Department of Anthropology Peabody Museum Harvard University Cambridge, Massachusetts 02138 Rolf M. Quam Department of Anthropology State University of New York Binghamton, New York 13902

Yoel Rak Department of Anatomy and Anthropology Sackler School of Medicine Tel Aviv University Ramat Aviv 69978, Israel

Contributors

W. J. Rink

531

Department of Geology McMaster University Hamilton, Ontario L8S 4M1, Canada

John D. Speth Museum of Anthropology University of Michigan Ann Arbor, Michigan 48109

Avraham Ronen Zinman Institute of Archaeology University of Haifa Haifa 31905, Israel

Department of Anthropology University of Arizona Tucson, Arizona 85721

Karen R. Rosenberg Department of Anthropology University of Delaware Newark, Delaware 19716

Chris Stringer Department of Palaeontology The Natural History Museum London SW7 5BD, United Kingdom

Christopher B. Ruff Department of Cell Biology and Anatomy Johns Hopkins University School of Medicine

Baltimore, Maryland 21205 Kathy D. Schick Anthropology Department and CRAFT Research Center Indiana University Bloomington, Indiana 47405 Henry P. Schwarcz

Mary C. Stiner

Eitan Tchernov Department of Evolution, Systematics, and Ecology Hebrew University Jerusalem 91904, Israel Anne-marie Tillier URA 376 CNRS Laboratoire d’Anthropologie

Université de Bordeaux I Talence 33405, France

Department of Geology McMaster University Hamilton, Ontario L8S 4M1, Canada

Erik Trinkaus Department of Anthropology Washington University St. Louis, Missouri 63130

Fred H. Smith Department of Anthropology Northern Illinois University DeKalb, Illinois 60115

Hélène Valladas Centre de Faibles Radioactivités Laboratoire Mixte CNRS-CEA Gif sur Yvette Cedex 91198, France

INDEX

AAR, 39 Abadzekh, 469 Abbevilian, 98 Abkhazia, 475 Abou Obeida, 102 Abri Pataud, 14, 497, 500 Abu el-Khass, 104 Abu Habil, 104 Abu Sif, 41, 44, 82, 157, 177, 208 Abu Sifian, 44 Achenheim, 305 Acheulean (Acheulian), 4, 7, 8, 9, 12, 17, 18, 40, 49,

58, 78, 82, 92, 93, 97, 99, 103, 104, 184, 206, 211, 214, 423, 424, 439, 440, 445, 451, 452, 453, 454, 455, 456, 457, 462, 463, 466, 469, 470, 480, 484, 490

Acheulo-Yabrudian, 4, 40, 41, 49, 78, 82, 84, 408, 464, 470 Activity areas, 132, 133, 134, 137, 138, 139, 140 Adlun, 41, 91 Aeolian, 117, 118, 119, 120 Afro-Arabian, 4, 78, 79, 84, 85, 86, 254 Ahmarian, 16, 23, 177, 184, 185, 476, 480, 487 Ain Aqev, 50 Ain Difla, 176 Akhschtyr Cave, 475 Akkaya, 472 Alpine, 6, 24, 79, 86, 97, 303 Altaï, 487 Amino acid racemization, 39 AMS, 6, 12, 13, 14, 20, 186 Amud, 1, 3, 10, 11, 33, 34, 43, 48, 49, 58, 63, 64, 65, 70, 74, 80, 86, 87, 91, 92, 93, 118, 119, 121, 122, 144, 145, 146, 148, 151, 152, 153, 154, 155, 156, 157, 158, 216, 224, 267, 272, 291, 292, 313, 314, 315, 318, 319, 330, 332, 355, 357, 358, 363, 364, 381, 386, 392, 393, 395, 397, 399, 406, 408, 409, 413 Amudian, 176, 177, 191 Anatolia, 23, 304, 306, 424, 461, 463, 466, 471, 472, 476, 480, 483, 488, 490

Apatite, 117, 278, 279, 280, 281, 282, 283, 284, 285,

286, 287 Apiancha, 478 Apomorphic, 8, 358 Apomorphies, 31, 32, 296, 302, 315, 413 Arago, 2, 8, 31, 299, 300, 301, 315, 373, 418 Archaic behavior, 127, 132, 135, 139 Arcy-sur-Cure, 13, 22, 304, 497, 499, 501 Ard Habibe, 102, 103 Ariendorf, 305 Armenia, 472, 474 Ash, 50, 60, 121, 122, 167, 227, 228, 242, 244, 258 Ashy deposits, 114, 121 Atapuerca, 7, 8, 29, 31, 32, 298, 299, 300, 301, 414, 454 Aurignacian, 2, 13, 14, 15, 16, 21, 22, 23, 24, 25, 26, 58, 114, 184, 185, 186, 190, 247, 257, 385, 424, 476, 478, 480, 487, 488, 490, 496, 497, 498, 499, 500, 501, 502

Auroch, 4, 93, 223, 231, 232, 236. 247, 250, 500 Australia, 1, 485, 523, 525 Australian, 315, 317, 355, 362, 372 Azov, 303, 470 Azykh, 9, 463, 466 Bacho Kiro, 14, 476, 497 Bachokirian, 14, 23, 476 Bagnoles, 301 Baksa-Idriss, 97 Balkans, 24, 304, 306, 424, 461, 466, 471, 472, 476, 478, 480, 488, 490, 496, 501

Baradostian, 16, 23, 476 Barakai, 332, 384, 385 Barakayevska, 472 Barma del Caviglione, 374 Base camp, 94, 130, 132, 139 Bassit, 97 Bayer, 97 Bezez, 47, 108 Bi-directional, 153, 184 Biache, 11, 305

533

534

Index

Biache-Saint-Vaast, 300, 301 Bilzingsleben, 8, 9, 299, 300, 463 Biogenic, 50, 51, 92, 108, 113, 120, 121

Cladism, 296, 297 Cladistic, 6, 296, 365 Cladogenesis, 6, 30, 365

Bioturbation, 92, 115, 117, 120, 121, 122, 123

Clastic sediments, 117 Climate, 59, 65, 78, 108, 118, 119, 120, 123, 242, 246, 254, 256, 286, 295, 406, 502, 513

Biozonation, 461 Bison, 494

Bitumen, 93, 182, 184, 186, 187, 189, 190, 191, 192, 193, 194, 195, 196, 200, 201, 202, 203

Black Sea, 304, 424, 461, 462, 464, 465, 468, 470, 471, 472, 473, 477, 478, 480, 488, 490 Boegoeberg, 511, 512, 514, 516, 519 Bohuslavice, 281, 282, 283, 286 Boker Tachtit, 15, 23, 64, 143, 211, 215, 216, 311 Border Cave, 319 Borj Qinnarit, 104 Bosphorus, 307, 308 Bovid, 34, 85, 500 Boxgrove, 8, 298 Broken Hill, 31, 315 Bronze Age, 408 Brown bear, 247

Brunhes-Matuyama, 454 Buffalo, 513 Bulgaria, 14, 23, 476, 478, 497

Burial, 16, 34, 59, 64, 65, 69, 70, 82, 86, 224, 227, 263, 264, 280, 281, 284, 306, 353, 406, 409, 423, 440, 441, 443, 469 Burrow, 112, 114, 115, 120, 435 Bush pigs, 512 Byneskranskop Cave, 511, 516 Calcareous, 112, 115, 117, 118, 123, 281 Calcite, 58, 59, 113, 117, 120 Canids, 247 Carbon, 197, 201, 278, 279, 280, 281, 282, 283, 284, 285, 286 Caspian, 302, 306, 308, 470, 471, 472

Castel de Guido, 300

Climatic oscillations, 78, 256, 303, 304, 494, 501 Cognitive, 42, 130, 152, 217, 218, 264, 272, 423, 427,

428, 432, 435, 436, 439, 440, 441, 445, 449, 453, 457, 458, 519

Collagen, 203, 265, 278, 279, 280, 281, 282, 283, 284, 285, 286, 287

Combe Saunière, 501 Commensal, 85, 86, 265 Computerized tomography, 30

Core modification, 93, 145, 208 Core reduction, 42, 44, 50, 51,93, 145, 147, 149, 150, 151, 152, 154, 157, 167, 168, 170, 172, 173, 176, 177, 184, 214, 272

Cricetines, 84, 85, 86 Crimea, 10, 11, 12, 19, 20, 384, 461, 463, 468, 472, 480, 488 Cro-Magnon, 2, 13, 15, 23, 26, 71, 291, 360, 374, 424, 484, 486, 495

Cromerian, 99, 298 rvena Stijena, 466 CT, 30, 33 Cultural capabilities, 526 Cultural transmission, 431, 432, 433, 436, 450, 456, 457 Dali, 373 Danube, 12, 19, 461, 463, 472, 478, 495 Débordant, 145, 150

Décapage, 226, 227, 229, 266 Dederiyeh, 10, 11, 30, 33, 34, 48, 65, 92, 95, 108, 278, 281, 282, 283, 284, 285, 291, 292, 323, 324,

325, 326, 327, 328, 329, 330, 331, 332, 333, 334, 335, 336, 368, 381, 382, 392

Castillo, 14, 497 Caucasus, 7, 9, 11, 12, 13, 19, 20, 302, 303, 332, 461, 463, 469, 470, 471, 475, 480, 488, 490 Cave bear, 281, 282, 285 Cementation, 113, 117, 118 Cementum, 51, 59, 94, 265, 266 Central Asia, 306, 308, 487, 488 Ceprano, 7, 29, 298 Chaine operatoire, 44 Chakhaty, 472 Charcoal, 110, 115, 116, 122 Charente, 13, 304 Châtelperron, 500

Deep-sea cores, 6, 493, 502 Deep Sounding, 114, 115

Chatelperronian, 13, 14, 15, 21, 497, 498, 499, 500, 501, 502

Dmanisi, 2, 7, 17, 29, 298, 453 DNA, 35, 223, 486, 495, 524, 527

Chimpanzee, 425, 429, 433, 434, 435, 441, 444, 450,

Dneper, 461, 463, 472

458, 524, 525, 526 China, 1, 29, 368, 370, 372, 454, 455, 456, 485 Choukoutien, 360 Clactonian, 463, 464 Clade, 2, 6, 18, 26, 30, 31, 32, 33, 358, 406, 524, 525, 526

Deir ez-Zor, 103 Devil’s Tower, 30, 326, 328, 331, 384 Diagenesis, 3, 92, 113, 115, 117, 118, 119, 121, 122, 123, 244, 258, 280, 285

Diagenetic alteration, 62, 119, 280 Die Kelders Cave, 511, 512, 513, 514, 516, 517 Diepkloof, 516 Diet reconstruction, 278, 280, 285, 286

Diffusion studies, 432 Dispersal events, 80 Dispersals, 2, 3, 6, 30, 80, 340, 424, 449

Dnester, 463, 472 Dobrogea, 468 Dolni Vestonice, 281, 282, 283, 285, 286 Domestic fire, 443, 445

Dordogne, 304, 500

Index

535

Dose-rate, 70, 72 Dosimeter, 60, 70, 72 Douara, 44, 82, 95, 108, 120, 121, 122, 123, 168, 176, 177 Dripline, 132, 133, 135, 136, 138

France, 11, 13, 14, 15, 20, 21, 22, 23, 24, 87, 177, 285, 297, 298, 300, 305, 384, 494, 496, 497, 499, 500, 501, 502, 509 Fur seal, 513, 514

Dubossary I, 463

Galilee, 3, 9, 44, 50, 60, 63, 70, 114, 241, 250, 254, 255, 256

Dukove, 382, 388 Dune Field Midden, 511 Dzhrutchula, 474, 475 Dzudzuana, 478, 479

Ganovce, 469 Gard-Ardèche-Provence, 304

el Kown, 490

Gazelle, 4, 85, 94, 226, 229, 231, 234, 235, 236, 247, 249, 250, 253, 254, 265, 266, 267 Geissenklösterle, 14, 497 Gene flow, 32, 79, 292, 319, 320, 340, 449, 525 Genetic data, 272, 524, 525, 527 Genetic drift, 296, 305, 307 Geogenic, 92, 108, 117, 118, 122 Geometric Kebaran, 184, 190

el-Faraché, 102

Georgia, 29, 298, 453, 463, 472, 474, 475, 479, 488

el-Meira, 103

Engis 2, 328, 330, 331 Epigravetian, 304

Germany, 14, 21, 298, 299, 300, 304, 497 Geula, 80, 91 Gombe National Park, 429 Gran Dolina, 7, 29, 31, 32, 298 Gravettian, 281, 386 Greece, 8, 20, 298, 304 Grimaldi, 365

Eremian, 78 Erevan, 474

Gubskiy, 472 Günz, 97

Erq el Ahmar, 15, 48

Guomde, 32

Eryvan, 472 ESR, 3, 4, 6, 9, 10, 11, 12, 20, 32, 33, 34, 35, 39, 44,

Hamanaka 2, 335

Early Würm, 467, 470, 472, 480 East Africa, 1, 417, 450, 451 Eemian, 12, 301, 303, 305 Ehringsdorf, 11, 12, 32, 300, 301, 315

el Kowm (el-Kown), 93, 95, 103, 177, 181, 183

el-Wad, 15, 16, 24, 108, 118 Eland, 31, 511, 512, 513, 516, 519 Elandsfontein, 31 Election spin resonance, 405

45, 47, 48, 49, 51, 57, 58, 59, 60, 61, 62, 63,

64, 65, 70, 71, 72, 78, 80, 82, 86, 110, 293, 294, 340, 365, 373, 405, 406, 409, 464

Hamra, 118, 119 Hare, 247, 249, 255, 258, 281

Euphrates, 95, 97, 103, 104, 181 Eurasia, 1, 2, 5, 6, 7, 10, 14, 16, 17, 18, 20, 23, 26, 78,

Harhoura, 319 Haua Fteah, 10, 22, 319, 364, 418 Hayonim, 3, 4, 11, 16, 39, 44, 45, 47, 48, 49, 50, 51,

79, 93, 295, 406, 417, 424, 425, 454, 455, 457, 486, 487, 509, 519, 524 Eurasian, 2, 6, 16, 17, 22, 26, 78, 87, 250, 254, 292,

58, 60, 62, 63, 65, 70, 72, 80, 82, 84, 85, 86, 92, 93, 94, 108, 113, 117, 118, 119, 120, 121, 122, 123, 165, 167, 168, 169, 170, 171, 172,

296, 316, 319, 407, 416, 418, 425, 510, 519,

173, 174, 175, 176, 177, 208, 216, 241, 242,

525, 527

243, 244, 247, 248, 250, 251, 253, 254, 255,

Eve, 223, 319, 485

256, 257, 258, 259, 487

Hazar Merd, 91 Fallow deer, 4, 94, 226, 231, 233, 247, 249, 250, 253, 254

Far’ah II, 58, 65, 153, 155, 157, 207

Hearth, 22, 50, 51, 69, 70, 92, 110, 112, 113, 115, 116,

117, 120, 121, 122, 132, 133, 134, 135, 136, 138, 139, 140, 144, 167, 227, 228, 250, 263, 272

Fauresmith, 452

Heinrich event, 497, 502, 503

Fertile Crescent, 467

Hengelo interstadial, 497, 498 Hoedjies Punt, 511, 516 Holstein-Hoxnian, 299 Home base, 443 Horizontal transmission, 423 Horka, 469 Horse, 87, 247, 250, 281, 494, 500 Human cognition, 205, 423, 439 Hummalian, 44, 176, 182, 190, 191, 194, 202 Hunter-gatherer, 92, 128, 129, 130, 254, 258, 263,

Fidio, 99

Fire, 51, 69, 121, 250, 264, 273, 423, 442, 443, 444, 445, 502 Flaking, 44, 145, 146, 147, 149, 151, 152, 153, 154, 156, 157, 158, 167, 168, 171, 172, 176, 193, 210, 211, 214, 450, 451, 452, 454, 456 Florisbad, 32 Fodor, 93, 217 Fontechevade, 300 Fonténioux, 500

Foramen magnum, 330, 386, 387 Fox, 85, 281

264, 267, 268, 270, 271, 304, 516, 526

Hunting strategies, 25, 264, 272 Hviraty, 474

536

Index

Hyena, 85, 93, 228, 229, 230, 233, 236, 286, 444, 512, 513, 514

Khor el-Aassi, 102 Khrischtchi, 463

Kiik-Koba, 13, 469, 472 Ice core-sequences, 493

Klasies Cave, 319

Ignatienkov Kutok, 463 llskaya, 472

Klasies River Mouth, 32, 318, 414, 417, 425, 511,

Imitation, 428, 429, 430, 432, 435, 441 Imo, 428

Inuits, 395 Iraq, 91, 201, 318, 368, 381 Isle, 22, 500 Isochron analysis, 71 Isotope, 3, 17, 31, 32, 50, 58, 59, 61, 65, 72, 80, 84, 86, 94, 177, 272, 278, 279, 280, 281, 282, 283, 284, 285, 286, 287, 292, 313, 406, 424, 463,

512, 513, 514, 516, 519 Koobi Fora, 29, 451 Korolevo, 463, 476 Koshima, 428

Kostenki, 472, 480 Krapina, 11, 12, 292, 317, 340, 341, 342, 343, 344, 345, 346, 347, 348, 349, 350, 375, 414 Krems, 24, 497 Ksar ‘Akil (Ksar Akil), 11, 15, 23, 48, 85, 91, 118, 119, 168, 385, 386, 476

464, 466, 467, 470, 471, 472, 476, 480, 494,

Kudaro, 463, 466, 469, 470, 474, 475, 478

496, 497, 501, 502, 511

Istállóskö, 497

Kunji, 51, 120 Kurkar, 119

Italy, 7, 8, 12, 14, 20, 21, 22, 23, 29, 93, 224, 232, 258, 298, 300, 304, 496

L‘Arbreda, 497

La Chappelle, 315 Jabal Bicheri, 181, 201 Jabal Minshar, 181

La Chèvre, 500 La Cotte St Brelade, 305

Jabal Mqaïbara, 181

La Ferrassie 6, 333, 334, 336, 368

Jackass penguins, 512 Japanese macaques, 428 Jaramillo, 297 Java, 291, 454 Jebel Idriss, 99, 102 Jebel Irhoud, 2, 32, 319, 365 Jerf ‘Ajla, 44, 91, 108, 118, 119, 121, 123, 152

La Gravette, 500 La Madeleine, 383, 384, 385 La Montagne de Segrez, 304 La Quina, 316, 326, 328, 330, 331, 413

La Rochette, 497 Laetoli, 319

Jinniushan, 293, 368, 370, 372, 373, 375 Jordan, 7, 9, 23, 41, 51, 91, 104, 128, 129, 141, 176, 202, 211, 369

Lagomorphs, 82, 242, 249, 250, 255, 256 Lake pollen sequences, 493 Laminar, 19, 21, 110, 155, 157, 158, 172, 173, 176, 177, 178,210,211,214,215 Lantian, 455

Kabazi, 20 Karain, 463, 464, 466, 467, 480, 484, 489, 490

Large game, 242, 253, 254, 255, 259, 286 Last Glacial Age, 39

Karangat transgression, 470 Karstic, 7, 63, 108, 225, 472

Lateralisation, 439

Jerzmanovizian, 499

Lapps, 395

Kasteelberg, 511 Kebara, 1, 3, 4, 10, 11, 16, 33, 34, 39, 41, 43, 47, 48,

49, 50, 51, 58, 65, 70, 72, 74, 80, 85, 86, 87,

91, 92, 93, 108, 110, 113, 114, 117, 118, 119, 120, 121, 122, 123, 143, 152, 153, 154, 155, 156, 157, 165, 168, 208, 210, 211, 214, 215,

216, 223, 224, 225, 226, 227, 228, 231, 232, 233, 234, 235, 236, 258, 266, 267, 272, 291, 292, 293, 313, 315, 317, 318, 319, 332, 340, 358, 368, 370, 371, 372, 373, 375, 381, 392, 393, 395, 397, 406, 408, 409, 413

Kebaran, 60, 94, 109, 114, 184, 190, 241, 242, 244, 245, 246, 247, 248, 249, 251, 253, 254, 255, 257, 258

Kefar Menahem, 104 Kenya, 29, 32, 94, 281, 284, 285, 525 Kharga oasis, 65 Khattab, 97, 102, 103, 104 Khergulis-klde, 478

Le Figuier, 383, 384, 385 Le Flageolet, 132, 497 Le Lazaret, 300 Le Piage, 15, 22, 499, 500, 501 Lebanon, 11, 23, 85, 91, 104, 385, 476 Leopard, 242, 243, 247 Les Cottés, 497, 500, 501 Liaoning Province, 370 Liashui, 373

Libya, 10, 22, 364, 418 Limpet, 516, 518, 519 Linguistic, 1, 43, 217, 264, 433, 436, 440 Lions, 444 Liujiang, 373 Lizard, 247, 249, 250, 255 Lochkovskoye, 469

Longgupo, 454 Lower Layer E (Lower E), 44, 48, 49, 72, 82, 84, 93, 167, 171, 172, 173, 174, 175, 176, 177 Lusakert, 472

Index

537

Magdalenian, 304 Mahardeh, 102

Obercassel, 318 Ohalo, 360

Mala Vorontsovska, 478

Oka, 463

Malaya Vorontskovskaya, 475 Mamia, 468

Oldowan, 7, 17, 423, 439, 440, 450, 451, 452, 453

Manganese, 117, 182, 184, 196, 198, 201, 202 Manych Inlet, 470, 471, 480 Marillac Cave, 285 Matuyama, 7 Mauer, 2, 8, 31, 32, 298, 299, 302, 418 Mediterranean sea, 78, 99, 303

Mental modules, 93, 217 Mental template, 51, 145, 206, 440

Mesopotamia, 490 Micherfeh, 97 Micoquian, 12, 13, 19, 472, 480

Microfauna, 51, 82, 86, 242, 251 Middle Awash Valley, 452 Migration hypothesis, 320, 406, 408, 409 Mikhaylovskoye, 463 Milovice, 281, 282, 283, 284, 286 Minatogawa, 373

Mlade , 14, 344, 476 Mobility, 3, 11, 19, 25, 49, 51, 52, 72, 92, 94, 128, 130, 264, 265, 266, 267, 268, 270, 271, 272, 277, 403, 502 Modern behavior, 2, 92, 128, 272, 425, 458, 510 Modern foragers, 92, 135, 136, 138, 139

Omo, 32, 319, 417 Onager, 247 Orangutan, 434 Orce, 29, 298 Organic matter, 92, 110, 118, 121, 122, 195, 202, 280, 281, 283 Orontes, 95, 97, 102, 103, 104 Ortvale, 472

Osetia, 9, 469, 474, 475 OSL, 6, 57 Ostrich, 4, 32, 85, 138, 247, 250, 251, 255, 514 Out of Africa, 35, 223 Oxygen-isotope stage, 79, 318, 493, 503, 511

Pair-non-Pair, 500 Palearctic, 4, 78, 82, 84, 85, 86 Paleoauxology, 388 Paleoclimate, 16 Paleodose, 48 Palestine, 104 Palmyra, 44, 181

Mollusk, 515, 516, 519

Paternoster Midden, 511 Paviland, 374 Pech de 1’Azé, 325, 326, 328, 330, 331, 332, 383, 385 Penultimate Glacial, 39 Perigord, 500

Monasheskiy, 472 Montenegro, 466, 468

Petralona, 2, 8, 31, 298, 299, 315 Phonemic speech, 425, 510

Montmaurin, 8, 300, 301 Morocco, 9, 87, 256, 365, 525 Mortuary, 86 Mount Circeo, 315 Mt. Carmel (Mount Carmel), 1, 3, 9, 10, 11, 23, 33, 50, 70, 80, 86, 91, 109, 112, 225, 292, 367, 406, 407 Multiregional evolution, 406 Multiregional model, 30 Multiregional perspective, 406 Mustelids, 247

Phosphate, 51, 60, 112, 113, 115, 117, 121, 123, 133, 140 Phosphatic, 115, 117, 121 Phreatic, 108, 117, 118 Pleitropic, 296 Plesiomorphic, 9, 296, 301, 313, 315, 316, 317, 318, 319, 358, 372, 374, 376 Plesiomorphies, 31, 315 Pogrebi I, 463

Modern Man, 483, 485, 487, 488, 490

Poland, 21, 304 Pontnewydd, 11, 300

Naamé, 47

Post-depositional, 49, 60, 107, 108, 113, 117, 184,

Nadaouiyeh, 95

186, 191, 227, 244 Postcranial, 7, 8, 9, 10, 13, 30, 235, 291, 332, 335, 376

Nahal Aqev, 44, 82, 153, 155, 208 Nahal Hemar, 202, 203 Nahr el-Kebir, 97, 99, 100, 103, 104 Nahr Ibrahim, 108 Ndutu, 31 Negev, 3, 15, 41, 44, 50, 58, 64, 91, 153, 154, 176, 208, 210, 211, 214, 216, 487 Nelson Bay Cave, 511, 512, 516, 519 New Guinea, 457 Ngaloba, 32 Ngandong, 35 Nihewan Basin, 455 Nitrogen, 278, 279, 281, 282, 283, 284, 286, 287 North Africa, 316, 319, 424, 452, 456, 485, 488

pre-Aurignacian, 21, 176

Predator, 13, 25, 51, 227, 228, 229, 230, 232, 233, 254, 277, 278, 286, 430, 433, 444 Predmosti, 374, 383, 384, 386 Prey, 13, 23, 25, 26, 224, 226, 229, 230, 231, 232, 233, 234, 236, 241, 242, 248, 251, 254, 255, 256, 258, 259, 263, 264, 273, 279, 434, 441

Procurement, 25, 42, 127, 128, 129, 130, 131, 167, 223, 224, 226, 231, 232, 234, 236, 259 Proto-Charentian, 463, 464 Protoculture, 423, 427, 429, 430 Provision system, 441, 442, 443, 445 Provisioning strategies, 128

538

Index

Prut, 463

Scanning Electron Microscope (SEM), 195, 196, 197,

Pyrenees, 304

198, 201 Sea Harvest, 511, 516, 519 Season of death, 265, 266, 267 Seasonal site occupation, 94, 265, 267 Seasonal site use, 265

Qafzeh, 1, 2, 3, 4, 10, 15, 32, 33, 34, 39, 41, 43, 47, 49, 58, 70, 72, 74, 80, 82, 84, 85, 86, 87, 91, 108, 117, 118, 119, 121, 123, 165, 167, 168, 208, 215, 216, 224, 266, 267, 272, 291, 292, 294, 312, 315, 317, 318, 319, 320, 326, 329, 340, 341, 343, 346, 358, 360, 361, 364, 365, 372, 374, 381, 382, 385, 388, 392, 393, 395, 397, 399, 405, 406, 407, 413, 417, 423, 441, 523 Qara Yakoub, 97, 103 Qdeir plateau, 181 Quincay, 500

Seclin, 177 Sedimentation, 20, 61, 70, 71, 80, 120, 122

Sefunim,47, 80, 91, 118, 119 Semiyblonovskoye, 469 Settlement-procurement, 130, 131, 132, 141 Shanidar, 1, 10, 11, 16, 51, 87, 91, 224, 291, 292, 306, 313, 314, 315, 318, 319, 364, 365, 368, 381, 392, 393, 395, 397, 399 Shellfish, 255, 519

Quneitra, 48, 152, 153, 154, 157, 168, 207, 214, 215, 216

Shovakh, 41 Shukbah, 41 Siberians, 395

Radiocarbon, 48, 57, 69, 110, 113, 114, 186, 373, 409, 495, 497, 499, 501, 502, 509 Radiometric dates, 104, 144, 157, 463, 478, 480

Sikh, 362

Ras el Kelb, 91

Siliceous, 50, 110, 115, 118, 121

Sima de Los Huesos, 299 Sinanthropus, 315

Red deer, 62, 87, 94, 231, 233, 243, 247, 249, 250,

Singa, 32

253, 254, 494, 496, 500, 501 Red Sea, 79

Site formation processes, 3, 41, 48, 50, 92, 107, 108, 123 Sitt-Markho, 97, 99, 100, 101, 104

Reduction sequence, 3, 19, 42, 44, 145, 146, 151, 152, 153, 154, 157, 158, 190, 192, 206, 207, 210, 215 Regionality hypothesis, 320

Régourdou, 357, 362 Reilingen, 299, 300

Reindeer, 87, 281, 285, 305, 494, 500, 501, 502 Replacement hypothesis, 509 Retouch, 174, 184, 186, 187, 188, 189, 192, 193, 271, 273, 456, 458, 463, 472, 474

Retouched tools, 44, 144, 151, 168, 170, 177, 466, 472

Siuren, 480 Skalistyi, 480 Skhul, 1, 2, 3, 10, 32, 33, 34, 47, 58, 70, 72, 74, 80, 82, 86, 108, 119, 120, 224, 266, 267, 272, 291, 292, 311, 312, 313, 314, 315, 316, 317, 318, 319, 320, 326, 328, 329, 333, 334, 336, 340, 341, 342, 343, 344, 345, 346, 347, 348, 349, 350, 358, 360, 361, 362, 363, 364, 365, 367, 372, 374, 375, 381, 382, 383, 384, 385, 386,

388, 392, 393, 397, 399, 406, 407, 413, 417

Rhinoceros, 247 Riencourt-lès-Bapaume, 177

Slovakia, 12, 382, 383, 469 Small game, 94, 242, 244, 247, 248, 249, 250, 251,

Roc de Combe, 15, 497, 499, 500, 501 Roc de Marsal, 328, 330, 332, 333, 334, 336, 358, 383, 384, 385 Rodents, 4, 242, 244, 279 Roe deer, 4, 85, 87, 247

254, 255, 256, 258, 259 Small prey, 255, 259 Social learning, 93, 158, 218, 423, 427, 428, 429, 430, 431, 432, 434, 435, 444, 450, 454 Solutrean, 304, 440 South Africa, 1, 32, 39, 278, 318, 452, 510, 512, 516, 518, 523, 525

Rosh Ein Mor, 44, 82, 153, 154, 155, 157, 158, 176, 177, 210, 214, 216 Russia, 386, 499

Russian, 306, 307, 472, 480

Spain, 7, 13, 15, 21, 23, 29, 298, 299, 304, 496, 497, 500, 509

Stalingradskaya, 472 Saalian, 300, 307

Saccopastore II, 315 Sagvardzile, 478 Sahara-Arabian, 78, 79, 84, 91 Sahba, 41, 44

Saint-Césaire, 497, 499, 500 Sajour valley, 97 Sakhazia, 472, 476 Samarkand, 488

Standardization, 24, 206, 207, 208, 217, 440, 458, 510 Steinheim, 8, 299, 300

Steppe, 85, 488 Streletskayan, 499 sub-Saharan Africa, 3, 27, 318, 425, 456, 485

Sambungmachan, 35

Subalyuk 2, 328, 383 Sukhaya Mechetka, 472 Sungir, 386 Surozh transgression, 472 Swanscombe, 8, 9, 31, 299, 300

Samertskhle-klde, 478

Symbolic behavior, 52, 423, 439

Index

539

Symboling behavior, 439, 440, 443

Szeletian, 12, 21, 499

Trinchera Dolina, 454 Tskhaltsitalskaya, 472 Tsona, 474 Tsony, 475 Tsopi, 475

Tabelbala, 452

Tsutskhavat, 472 Tyrhenian, 303

Syria, 9, 11, 91, 92, 93, 95, 97, 98, 99, 102, 103, 104, 108, 176, 177, 181, 201, 202, 278, 280, 281,

292, 323, 335, 368, 381

Tabun B, 11, 34, 41, 47, 49, 51, 87, 157, 209, 267, 268, 271, 272, 281, 409 Tabun B-type, 3, 41, 47, 48, 49, 50, 51, 80, 86, 92, 93,

143, 144, 156, 157, 323, 408, 409 Tabun C, 10, 41, 49, 72, 87, 209, 266, 267, 268, 272, 294, 409, 417

Tabun C-type, 41, 47, 49, 50, 80, 82, 93, 168, 408, 409 Tabun Cl, 10, 11, 65, 369, 393, 395, 397, 399, 407, 408, 409, 413

Tabun C2, 10, 407, 408, 410, 411, 412, 413, 414, 416, 417, 418 Tabun C3, 393, 409 Tabun D, 41, 48, 49, 62, 65, 82, 84, 176, 209, 210, 211,214,216 Tabun D-type, 3, 23, 41, 44, 48, 50, 63, 65, 82, 84, 86,

U-series, 3, 6, 9, 10, 11, 20, 58, 59, 60, 61, 62, 65, 82

Ukraine, 463 Ukrainian, 306

Uluzzian, 14, 15, 21, 499 Um Naqus C, 41 Umm el Tlel, 93, 182, 185, 194, 202 Umm el-Tlel, 95 Umm Qatafa (Oumm-Qatafa), 82, 84, 118, 120, 123 Ungulate, 24, 26, 85, 94, 224, 225, 226, 229, 230, 232, 233, 236, 242, 246, 249, 250, 251, 253, 254, 258, 305, 512 Unidirectional, 43, 79, 145, 152, 153, 154, 168, 172, 174, 176, 188, 189, 210, 214, 457 Unipolar, 44, 47, 144, 145, 149, 153, 156, 157, 158, 184, 210, 215, 272

87, 93, 143, 144, 153, 156, 157, 159, 176, 177 Tabun I, 72, 87, 267, 272, 292, 293, 311, 312, 313, 315, 317, 318, 319, 353, 354, 355, 358, 363, 364 Tabun II, 72, 272, 292, 293, 294, 312, 313, 353, 354, 355, 358, 360, 361, 362, 363, 364

Upper E, 47, 72, 80, 82, 93, 167, 168, 169, 170, 173, 176 Upper Galilee, 60, 241, 250, 254, 255 Ural, 303, 308 Uranium series, 39

Tachenghit, 452

Urupskoye, 469

Taglar, 472

Uzbekistan, 12, 306

Takasago 10, 335 Tanzania, 429, 525 Taro-klede, 478 Taubachian, 12, 18, 469 Taurus, 41, 49, 463

Tautavel, 298 Technical innovation, 178, 259, 423, 432 Technologic innovations, 258

Technological complexity, 209, 424, 457, 458 Tetnara, 319 Temnata, 14, 24, 476, 478, 497 Ternifine, 358, 418

Terra rosa, 112, 118, 120 Teshik-Tash, 10, 11, 12, 306 TL, 3, 4, 6, 9, 33, 34, 39, 44, 45, 47, 48, 49, 51, 57, 58, 60, 62, 63, 64, 69, 70, 71, 72, 74, 78, 80, 86, 109, 113, 120, 144, 177, 186, 216, 293, 294,

340, 364, 365, 405, 409, 463, 464, 472 Tor Faraj, 48, 50, 51, 92, 122, 128, 129, 133, 135, 136,

Vertical transmission, 423, 430, 432, 435, 436 Vézére, 500 Vindija, 12, 281, 282, 283, 285, 286, 413, 414, 416 Visogliano, 298 Volga, 472 Vulture, 247

Weschelian, 301 West Asian, 15, 254 Wild goat, 85, 247 Wild pig, 247 Wildcat, 247 Willendorf, 14, 497 Wolf, 281, 285, 445 Wolves, 286, 444

Woolly mammoth, 281 Xerotropical, 78

138, 139, 140, 141, 152, 153, 154, 157, 168, 211 Tor Sabiha, 48, 50, 51, 128, 141, 152, 153, 157, 211 Tortoise, 242, 243, 247, 248, 249, 250, 251, 252, 255, 256, 257, 258, 515, 516, 517

Zafarraya, 13, 20, 501

Trans-Jordanian plateau, 108

Zagros, 11, 19, 23, 120, 306, 467, 472, 473, 474, 488,

Transcaucasia, 461, 463, 466, 469, 470, 472, 474, 476,

478, 479, 480 Transhumance, 128, 267

490 Zaqrounian, 104 Zarzi, 91

Transitional industries, 186, 206 Transmission chain studies, 432, 433 Travertine, 9, 58, 65, 117, 120, 153, 469

Zaskalnaya, 12, 20, 472 Zuttiyeh, 1, 3, 9, 10, 33, 58, 70, 74, 82, 84, 118, 120, 123, 291, 292, 306, 312, 313, 318, 406, 417, 486

Yabrud, 9, 40, 41, 44, 82, 91, 95, 118, 119, 123