Hostname: page-component-586b7cd67f-rdxmf Total loading time: 0 Render date: 2024-11-26T04:39:48.691Z Has data issue: false hasContentIssue false

Genes, Fossils, and Culture. An Overview of the Evidence for Neandertal–Modern Human Interaction and Admixture

Published online by Cambridge University Press:  18 February 2014

João Zilhão
Affiliation:
Department of Archaeology & Anthroplogy, Bristol University, 43 Woodland Road, Bristol BS8 1UU, UK

Abstract

This paper re-examines current arguments concerning the evidence for Neandertal-modern human interaction and admixture. While most researchers now agree that the ancestry of all present day humans can be traced back to African late Middle Pleistocene populations, at a time when the remainder of Eurasia was inhabited by ‘archaic humans’, most notably the Neandertals, issues that remain to be resolved are the tempo and mode of early modern human dispersal and interaction with archaic humans.

This paper focuses on what happened at the time of contact in Europe, and assesses the level of admixture that may have occurred, as well as the extent to which such level may have varied in both time and space. It explains how the available mtDNA evidence does not preclude admixture at the time of contact, and is in fact consistent, depending on a number of parameters, with a possibly substantial Neandertal contribution to the initial modern human population of Europe. It is argued that the absence of Neandertal mtDNA lineages among present Europeans is likely, on dating evidence, to be simply a particular case of generalised loss of Pleistocene mtDNA lineages. Although the full range of interaction types (mutual avoidance, hostile confrontation, full integration) is conceivable, there is plenty of archaeological evidence to suggest that admixture must have been the general rule, and that the paleontological evidence for the generalised presence of archaic traits among Europe's earliest moderns implies the transmission of genes, and indicates that mixed groups should have been reproductively viable. In this context, it would seem that the most parsimonious explanation for the disappearance of the Neandertal mtDNA lineage is genetic swamping.

Résumé

Cette étude ré-éxamine les discussions actuelles concernant les témoignages d'interaction et d'intégration Néandertal -homme moderne. Tandis que la plupart des chercheurs est maintenant d'accord que l'ascendance de tous les humains actuels remonte aux populations africaines de la fin du Pléïstocène moyen, à une époque où le reste de l'Eurasie était habité par des ‘humains archaïques‘, plus précisément les Néandertaliens, les questions qui restent à résoudre sont le tempo et le mode de dispersion et d'interaction des humains modernes avec les humains archaïques.

Cette étude se concentre sur ce qui s'est passé au moment où le contact s'est établi en Europe, et évalue le niveau de mélange qui s'est peut-être produit, ainsi que l'étendue jusqu'à laquelle un tel niveau peut avoir varié aussi bien dans le temps que dans l'espace. Ce qui explique comment les témoignages disponibles de mtADN n'excluent pas un mélange au moment du contact, et sont en fait consistants, selon un certain nombre de paramètres, avec la possibilité d'une contribution néandertalienne substantielle à la population moderne initiale en Europe. On argumente qu'il est possible que l'absence de lignées de mtADN Néandertalien parmi les européens actuels, selon les témoignages de datation, soit tout simplement un cas particulier de perte généralisée des lignées de mtADN du pléïstocène. Bien que toute la gamme des types d'interaction, (mutuelle tenue à l'écart, confrontation hostile, complète intégration) soit concevable, de nombreux témoignages archéologiques suggèrent que le mélange doit avoir été la règle générale et que les témoignages paléontologiques de la présence généralisée de traits archaïques parmi les premiers modernes d'Europe implique la transmissiom de gènes et indique que des groupes mixtes devaient pouvoir se reproduire. Dans ce contexte, il semblerait que l'explication la plus parcimonieuse de la disparition de la lignée de mtADN Néandertalien soit la l'écrasement génétique.

Zusammenfassung

Dieser Artikel untersucht erneut die aktuellen Argumente zu Nachweisen von Interaktionen und der Vermischung von Neandertalern und modernen Menschen. Während die meisten Forscher mittlerweile in ihrer Meinung übereinstimmen, dass die Herkunft aller heutigen Menschen auf die afrikanischen Populationen des späten Mittelpleistozäns zurückzuführen ist – einer Zeit, als der Rest von Eurasien von ,archaischen Menschen’ bevölkert war (vor allem der Neandertaler), müssen die Streitpunkte des Tempos und der Art und Weise der Verbreitung der frühen modernen Menschen und ihrer Interaktion mit den archaischen Menschen, noch gelöst werden. Der Schwerpunkt dieses Artikel liegt auf der Frage, was genau in der Zeit des Kontaktes in Europa passierte. Hier wird sowohl der Grad der Vermischung, die bestanden haben könnte, untersucht, als auch das Ausmaß, in dem dieser Vermischungsgrad in Zeit und Raum variiert haben könnte. Dbaie wird dargelegt, dass die vorhandene mtDNA Evidenz die Vermischung in der Zeit des Kontaktes nicht ausschließt; sie ist, abhängig von der Anzahl der Parameter, mit einem möglicherweise substantiellen Beitrag der Neandertaler zur anfänglichen Population moderner Menschen in Europa, tatsächlich konsistent. Es wird weiterhin erläutert, dass die Abwesenheit von Neandertaler mtDNA unter den heutigen Europäern sogar wahrscheinlich ist, und wegen der zeitlichen Tiefe als Beispiel eines allgemeinen Verlusts von Pleistozäner mtDNA darstellt. Obwohl die volle Bandbreite an Interaktionsarten (gegenseitiges sich Vermeiden, feindliche Konfrontation, volle Integration) denkbar ist, gibt es zahlreiche archäologische Nachweise, die suggerieren, dass Vermischung die allgemeine Regel gewesen sein muss, und dass die paläontologischen Nachweise für ein allgemeines Vorhandensein archaischer Spuren unter Europas ersten modernen Menschen die Transmission von Genen impliziert, und damit andeutet, dass Mischgruppen bezüglich der Fortpflanzung lebensfähig gewesen sein sollten. Vor diesem Hintergrund scheint die einfachste Erklärung für das Verschwinden der Neandertaler mtDNA genetische Verdrängung zu sein.

Résumen

Este artículo re-examina los argumentos actuales sobre la evidencia para interacción y mezcla entre Neandertal – hombre moderno. La mayoría de los investigadores hoy en día están de acuerdo en que los antecesores del hombre actual se encuentran entre las poblaciones africanas del Pleistoceno Medio en un momento en que el resto de Eurasia estaba habitado por ‘hombres arcaicos‘, de modo especial los Neandertales. Sin embargo, quedan aún por resolver una serie de factores como el tempo y modo de dispersión inicial del hombre moderno y su interacción con el hombre arcaico. Este trabajo se concentra en lo que ocurrió en el momento de contacto en Europa, y examina el nivel de mezcla que puede haberse dado, así como las posibilidades de variación en tiempo y espacio de ese nivel. Explica cómo la evidencia disponible de mtDNA no descarta la posibilidad de mezcla en el momento de contacto, y es de hecho consistente, dependiendo de una serie de parámetros, con una posible y substancial contribución del Neandertal a la población inicial de hombre moderno en Europa. Se mantiene que la ausencia de linajes de mtDNA de Neandertal en el europeo actual es probablemente, de acuerdo con la evidencia cronológica, simplemente un caso especial de una pérdida generalizada de linajes de mtDNA del Pleistoceno. Aunque se puede imaginar la serie completa de tipos de interacción (exclusión mutua, confrontación hostil, integración plena), hay mucha evidencia arqueológica que sugiere que la mezcla entre las dos poblaciones parece haber sido la regla general, que la evidencia paleontológica de la presencia generalizada de características arcaicas entre las primeras poblaciones de hombres modernos en Europa indica la transmisión de genes, y que grupos mixtos deben haber sido reproductivamente viables. En este contexto, parecería que la explicación más parsimoniosa para la desaparición del linaje mtDNA Neandertal es inundación genética.

Type
Research Article
Copyright
Copyright © The Prehistoric Society 2006

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

BIBLIOGRAPHY

Abbott, A. 2003. Anthropologists cast doubt on human DNA evidence. Nature 423, 468CrossRefGoogle ScholarPubMed
Adovasio, J.M., Soffer, O., Hyland, D.C., Illingworth, J.S., Klíma, B. & Svoboda, J. 2001. Perishable industries from Dolní Vestonice I: new insights into the nature and origin of the Gravettian. Archaeology, Ethnology & Anthropology of Eurasia 2, 4865Google Scholar
Aiello, L., & Wheeler, P. 2003. Neandertal Thermoregulation and the Glacial Climate. In van Andel, T.H. & Davies, W. (eds), Neanderthals and modern humans in the European landscape during the last glaciation, 147–66. Cambridge: McDonald Institute for Archaeological ResearchGoogle Scholar
Andel, T.H. van & Davies, W. (eds). 2003. Neanderthals and Modern Humans in the European Landscape During the Last Glaciation. Cambridge: McDonald Institute for Archaeological ResearchGoogle Scholar
Baffier, D. 1999. Les Derniers Néandertaliens. Le Châtelperronien. Paris: La Maison des RochesGoogle Scholar
Beauval, C., Maureiile, B., Lacrampe-Cuyaubère, F., Serre, D., Peressinotto, D., Bordes, J.-G., Cochard, D., Couchoud, I., Dubrasquet, D., Laroulandie, V., Lenoble, A., Mallye, J.-B., Pasty, S., Primault, J., Rohland, N., Pääbo, S. & Trinkaus, E. 2005. A late Neandertal femur from Les Rochers-de-Villeneuve, France. Proceedings of the National Academy of Sciences USA 102, 7085–90CrossRefGoogle ScholarPubMed
Binford, L. 1983. In Pursuit of the Past. London: Thames & HudsonGoogle Scholar
Caramelli, D., Lalueza-Fox, C., Vernesi, C., Lari, M., Casoli, A., Mallegnii, F., Chiarelli, B., Dupanloup, I., Bertranpetit, J., Barbujani, G. & Bertorelle, G. 2003. Evidence for a genetic discontinuity between Neandertals and 24,000-year-old anatomically modern Europeans. Proceedings of the National Academy of Sciences USA 100, 6593–7CrossRefGoogle ScholarPubMed
Churchill, S.E. & Smith, F.H. 2000. Makers of the Early Aurignacian of Europe. Yearbook of Physical Anthropology 43, 611153.0.CO;2-3>CrossRefGoogle Scholar
Clarke, D. 1978. Mesolithic Europe: the economic basis. London: DuckworthGoogle Scholar
Conard, N.J., Grootes, P.M. & Smith, F.H. 2004. Unexpectedly recent dates for human remains from Vogelherd. Nature 430, 198201CrossRefGoogle ScholarPubMed
Cooper, A., Drummond, A.J. & Willerslev, E. 2004. Ancient DNA: would the real Neandertal please stand up? Current Biology 14, R4313CrossRefGoogle ScholarPubMed
Currat, M. & Excoffier, L. 2004. Modern humans did not admix with Neanderthals during their range expansion into Europe. PLoS Biology, 2, 2264–74CrossRefGoogle Scholar
d'Errico, F. 2003. The invisible frontier. A multiple species model for the origin of behavioral modernity. Evolutionary Anthropology 12, 188202CrossRefGoogle Scholar
d'Errico, F., Zilhão, J., Baffier, D., Julien, M. & Pelegrin, J. 1998. Neanderthal acculturation in Western Europe? A critical review of the evidence and its interpretation. Current Anthropology 39, S144CrossRefGoogle Scholar
d'Errico, F. & Vanhaeren, M. 2005. Evolution or revolution? New evidence for the origin of symbolic behavior in Africa and Europe. In Mellars, P., Stringer, C., Bar-Yosef, O. & Boyle, K. (eds), Rethinking the Human Revolution: new behavioural perspectives on the origins and dispersal of modern humans, 31–3. Cambridge: McDonald Institute for Archeological Research/American School of Prehistoric Research (book of abstracts of the conference, September 7–11)Google Scholar
de Vivo, B., Rolandi, G., Gans, P.B., Calvert, A., Bohrson, W.A., Spera, F.J. & Belkin, H.E. 2001. New constraints on the pyroclastic eruptive history of the Campanian volcanic Plain (Italy). Mineralogy & Petrology 73, 4765CrossRefGoogle Scholar
Duarte, C., Maurício, J., Pettitt, P.B., Souto, P., Trinkaus, E., Plicht, H. van der & Zilhão, J. 1999. The Early Upper Paleolithic human skeleton from the Abrigo do Lagar Velho (Portugal) and modern human emergence in Iberia. Proceedings of the National Academy of Sciences USA 96, 7604–9CrossRefGoogle ScholarPubMed
Fedele, F.G., Giaccio, B., Isaia, R. & Orsi, G. 2002. Ecosystem impact of the Campanian Ignimbrite eruption in Late Pleistocene Europe. Quaternary Research 57, 420–4CrossRefGoogle Scholar
Fedele, F.G., Giaccio, B., Isaia, R. & Orsi, G. 2003. The Campanian Ignimbrite Eruption, Heinrich Event 4, and Palaeolithic change in Europe: a high-resolution investigation. In Robock, A. & Oppenheimer, C. (eds), Volcanism and the Earth's Atmosphere, 301–25. Washington DC: Geophysical Monograph 139CrossRefGoogle Scholar
Forster, P. 2004. Ice Ages and the mitochondrial DNA chronology of human dispersals: a review. Philosophical Transactions of the Royal Society London B 359, 255–64CrossRefGoogle ScholarPubMed
Gamble, C. 1983. Culture and society in the Upper Paleolithic of Europe. In Bailey, G. (ed.), Hunter-gatherer Economy in Prehistory. A European Perspective, 201–11. Cambridge: University PressGoogle Scholar
Garrigan, D., Mobasher, Z., Severson, T., Wilder, J.A. & Hammer, M.F. 2005. Evidence for Archaic Asian Ancestry on the Human X Chromosome. Molecular Biology & Evolution 22, 189–92CrossRefGoogle ScholarPubMed
Gaudzinski, S. & Roebroeks, W. 2000. Adults only. Reindeer hunting at the Middle Palaeolithic site SalzgitterLebenstedt, Northern Germany. Journal of Human Evolution 38, 497521CrossRefGoogle Scholar
Gaudzinski, S. & Roebroeks, W. 2003. Profile analysis at Salzgitter-Lebenstedt. A reply to Munson & Marean. Journal of Human Evolution 44, 275–81Google Scholar
Gilman, A. 1984. Explaining the Upper Palaeolithic revolution. In Spriggs, M. (ed.), Marxist Perspectives in Archaeology, 115–26. Cambridge: University PressGoogle Scholar
Golovanova, L.V., Hoffecker, J.F., Kharitonov, V.M. & Romanova, G.P. 1999. Mezmaiskaya cave: a Neanderthal occupation in the northern Caucasus. Current Anthropology 40, 7786CrossRefGoogle Scholar
Grayson, D.K. & Delpech, F. 2002. Specialized Early Upper Palaeolithic Hunters in Southwestern France? Journal of Archaeological Science 29, 1439–49CrossRefGoogle Scholar
Grayson, D.K. & Delpech, F. 2003. Ungulates and the Middle-to-Upper Palaeolithic transition at Grotte XVI (Dordogne, France). Journal of Archaeological Science 30, 1633–48CrossRefGoogle Scholar
Hathaway, J. 2004. East African artifacts support evolution of symbolic thinking in Middle Stone Age (http://www.eurekalert.org/pub_releases/2004-03/asu-eaa032504.php).Google Scholar
Henry-Gambier, D. 2002. Les fossiles de Cro-Magnon (Les Eyzies-de-Tayac, Dordogne): nouvelles questions sur leur position chronologique et leur attribution culturelle. Bulletin et Mémoires de la Société d'Anthropologie de Paris 14, 89112Google Scholar
Henshilwood, C. & Marean, C. 2003. The origin of modem human behavior. Critique of the models and their test implications. Current Anthropology 44, 627–51CrossRefGoogle Scholar
Henshilwood, C., d'Errico, E., Vanhaeren, M., Niekerk, K. van & Jacobs, Z. 2004. Middle Stone Age Shell Beads from South Africa. Science 304, 404CrossRefGoogle ScholarPubMed
Hublin, J.-J. 2000. Modern-nonmodern hominid interactions: a mediterranean perspective. In Bar-Yosef, O. & Pilbeam, D. (eds), The Geography of Neandertals and Modern Humans in Europe and the Greater Mediterranean, 157–82. Cambridge MA: Peabody Museum Bulletin 8Google Scholar
Hublin, J.-J., Barroso Ruiz, C., Medina Lara, P., Fontugne, M. & Reyss, J.L. 1995. The Mousterian site of Zafarra (Andalucia, Spain): dating and implications on the paleolithic peopling processes of Western Europe. Comptes-Rendus de l'Académie des Sciences de Paris IIa 321, 931–37Google Scholar
Hublin, J.-J., Spoor, F., Braun, M., Zonneveld, F. & Condemi, S. 1996. A late Neanderthal associated with Upper Palaeolithic artefacts. Nature 381, 224–6CrossRefGoogle ScholarPubMed
Jolly, C. 2001. A proper study for mankind: analogies from the papionin monkeys and their implications for human evolution. Yearbook of Physical Anthropology 44, 177204CrossRefGoogle Scholar
Klein, R. 2003. Whither the Neanderthals? Science 299, 1525–7CrossRefGoogle ScholarPubMed
Kuhn, S. & Stiner, M. 1998. The earliest Aurignacian of Riparo Mochi (Liguria, Italy). Current Anthropology 39, S17589CrossRefGoogle Scholar
Kuhn, S.L., Stiner, M.C., Reese, D.S. & Gùleç, E. 2001. Ornaments of the earliest Upper Paleolithic: new insights from the Levant. Proceedings of the National Academy of Sciences USA 98, 7641–6CrossRefGoogle ScholarPubMed
Kurtén, B. 1980. Dance of the Tiger. A novel of the Ice Age. New York: Pantheon BooksGoogle Scholar
Kurtén, B. 1986. Singletusk. A novel of the Ice Age. New York: Pantheon BooksGoogle Scholar
Lahr, M.M. & Foley, R. 1998. Towards a theory of modern human origins: geography, demography, and diversity in recent human evolution. Yearbook of Physical Anthropology 41, 37176Google Scholar
Malmström, H., Storå, J., Dalén, L., Holmlund, G. & Götherström, A. 2005. Extensive human DNA contamination in extracts from ancient dog bones and teeth. Molecular Biology & Evolution 22, 2040–7CrossRefGoogle ScholarPubMed
McBrearty, S. & Brooks, A. 2000. The revolution that wasn't: a new interpretation of the origin of modern human behavior. Journal of Human Evolution 39, 453563CrossRefGoogle ScholarPubMed
Mellars, P. 1999. In The Neanderthal Problem Continued. Current Anthropology 40, 341–50CrossRefGoogle Scholar
Mellars, P. 2004a. Neanderthals and the modern human colonization of Europe. Nature 432, 461–5CrossRefGoogle ScholarPubMed
Mellars, P. 2004b. Reindeer specialization in the early Upper Palaeolithic: the evidence from south west France. Journal of Archaeological Science 31, 613–7CrossRefGoogle Scholar
Mellars, P. 2005. The impossible coincidence. A single-Species model for the origins of modern human behavior in Europe. Evolutionary Anthropology 14, 1227CrossRefGoogle Scholar
Morin, E. 2004. Late Pleistocene Population Interaction in Western Europe and Modern Human Origins: new insights based on the faunal remains from Saint-Césaire, southwestern France. University of Michigan: Ph. D. dissertationGoogle Scholar
O'Connell, J.F. 2006. How did modern humans displace Neanderthals? Insights from hunter-gatherer ethnography and archaeology. In Conard, N.J. (ed.), When Neanderthals and Modern Humans Met, 4364. Tübingen: Kerns VerlagGoogle Scholar
Ovchinnikov, I., Götherström, A., Romanova, G.P., Kharitonov, V.M., Lidén, K. & Goodwin, W. 2000. Molecular analysis of Neandertal DNA from the northern Caucasus. Nature 404, 490–3CrossRefGoogle ScholarPubMed
Pääbo, S., Poinar, H., Serre, D., Jaenicke-Després, V., Hebler, J., Rohland, N., Kuch, M., Krause, J., Vigilant, L. & Hofreiter, M. 2004. Genetic analyses from ancient DNA. Annual Review of Genetics 38, 645–79CrossRefGoogle ScholarPubMed
Pike-Tay, A., Cabrera, V. & Bernaldo de Quirós, F. 1999. Seasonal variations of the Middle-Upper Paleolithic transition at El Castillo, Cueva Morin and El Pendo (Cantabria, Spain). Journal of Human Evolution 36, 283317CrossRefGoogle ScholarPubMed
Relethford, J. 2001. Absence of regional affinities of Neandertal DNA with living humans does not reject multiregional evolution. American Journal of Physical Anthropology 115, 95–8CrossRefGoogle Scholar
Relethford, J. 2003. Reflections of Our Past. Boulder: WestviewGoogle Scholar
Richards, M., Macaulay, V., Hickey, E., Vega, E., Sykes, B., Guida, V., Rengo, C., Sellitto, D., Cruciani, F., Kivisild, T., Villems, R., Thomas, M., Rychkov, S., Rychkov, O., Rychkov, Y., Gölge, M., Dimitrov, D., Hill, E., Bradley, D., Romano, V., Calì, F., Vona, G., Demaine, A., Papiha, S., Triantaphyllidis, C., Stefanescu, G., Hatina, J., Belledi, M.; di Rienzo, A., Novelletto, A., Oppenheim, A., Nørby, S., Al-Zaheri, N., Santachiara-benerecetti, S., Scozzari, S., Torroni, A. & Bandelt, H.-J. 2000. Tracing European founder lineages in the Near Eastern mtDNA pool. American Journal of Human Genetics 67, 1251–76CrossRefGoogle ScholarPubMed
Richards, M.P., Pettitt, P.B., Trinkaus, E., Stiner, M.C. & Trinkaus, E. 2001. Stable isotope evidence for increasing dietary breadth in the European mid-Upper Palaeolithic. Proceedings of the National Academy of Sciences USA 97, 6528–32CrossRefGoogle Scholar
Schmitz, R.W., Serre, D., Bonani, G., Feine, S., Hillgruber, F., Kraimtzki, H., Pääbo, S. & Smith, E.H. 2002. The Neandertal type site revisited: interdisciplinary investigations of skeletal remains from the Neander Valley, Germany. Proceedings of the National Academy of Sciences USA 99, 13342–7CrossRefGoogle ScholarPubMed
Serre, D., Langaney, A., Chech, M., Teschler-Nicola, M., Paunoviæ, M., Mennecier, Ph., Hofreiter, M., Possnert, G. & Pääbo, S. 2004. No evidence of Neandertal mtDNA contribution to early modern humans. PLoS Biology 2, 313–7CrossRefGoogle ScholarPubMed
Shennan, S. 2001. Demography and cultural innovation: a model and its implications for the emergence of modern human culture. Cambridge Archaeological Journal 11, 516CrossRefGoogle Scholar
Sinclair, A. 2003. Art of the ancients. Nature 426, 774–5CrossRefGoogle ScholarPubMed
Smith, C. 1992. Late Stone Age Hunters of the British Isles. London: RoutledgeCrossRefGoogle Scholar
Smith, E.H., Trinkaus, E., Pettitt, P.B. & Karavanić, I. 1999. Direct radiocarbon dates for Vindija G1 and Velika Pećina Late Pleistocene hominid remains. Proceedings of the National Academy of Sciences USA 96, 12281–6CrossRefGoogle ScholarPubMed
Soffer, O., Adovasio, J.M. & Hyland, D.C. 2000. The ‘Venus’ Figurines. Current Anthropology 41, 511–37CrossRefGoogle Scholar
Stiner, M. 1999. Palaeolithic mollusc exploitation at Riparo Mocchi (Balzi Rossi, Italy): food and ornaments from the Aurignacian through the Epigravettian. Antiquity 73, 735–54CrossRefGoogle Scholar
Stiner, M. 2001. Thirty years on the ‘Broad Spectrum Revolution’ and paleolithic demography. Proceedings of the National Academy of Sciences USA 98, 6993–6CrossRefGoogle ScholarPubMed
Stringer, C. 2002. Modern human origins: progress and prospects. Philosophical Transactions of the Royal Society London B 357, 563–79CrossRefGoogle ScholarPubMed
Stringer, C., Pälike, H., Andel, T.H. van; Huntley, B., Valdes, P. & Allen, J.R.M. 2003. Climatic stress and the extinction of the Neanderthals. In van Andel, & Davies, (eds) 2003, 233–40Google Scholar
Svoboda, J., Plicht, J. van der & Kuz̆elka, V. 2002. Upper Palaeolithic and Mesolithic human fossils from Moravia and Bohemia (Czech Republic): some new 14C dates. Antiquity 76, 957–62CrossRefGoogle Scholar
Svoboda, J. 2003. The Bohunician and the Aurignacian. In Zilhão, J. & d'Errico, F. (eds), The Chronology of the Aurignacian and of the Transitional Technocomplexes. Dating, Stratigraphies, Cultural Implications, 123–31. Lisboa: Instituto Português de ArqueologiaGoogle Scholar
Templeton, A. 2002. Out of Africa again and again. Nature 416, 4550CrossRefGoogle Scholar
Terberger, Th. & Street, M. 2003. Jungpaläolithische Menschenreste im westlichen Mitteleuropa und ihr Kontext. In Burdukiewicz, J.M., Fiedler, L. & Justus, A. (eds), Erkenntnisjäger: Kultur und Umwelt des frühen Menschen, 579–91. Halle: Veröffentlichungen des Landesamt für Archäologie Sachsen–Anhalt 57Google Scholar
Tindale, N.B. 1953. Tribal and intertribal marriage among the Australian aborigines. Human Biology 25, 169–90Google ScholarPubMed
Trinkaus, E. 2005a. Anatomical evidence for the antiquity of human footwear use. Journal of Archaeological Science 32, 1515–26CrossRefGoogle Scholar
Trinkaus, E. 2005b. Early Modern Humans. Annual Reviews of Anthropology 34, 207–30CrossRefGoogle Scholar
Trinkaus, E. & Zilhaão, J. 2002. Phylogenetic implications. In Zilhão, J. & Trinkaus, E. (eds), Portrait of the Artist as a Child. The Gravettian Human Skeleton from the Abrigo do Lagar Velho and its Archeological Context, 497518. Lisboa: Instituto Português de ArqueologiaGoogle Scholar
Trinkaus, E., Churchill, S.E., Ruff, Ch.B. & Vandermeersch, B. 1999. Long bone shaft robusticity and body proportions of the Saint-Césaire 1 Châtelperronian Neanderthal. Journal of Archaeological Science 26, 753–73CrossRefGoogle Scholar
Trinkaus, E., Milota, Ş., Rodrigo, R., Gherase, M. & Moldovan, O. 2003a. Early modern human cranial remains from the Peştera cu Oase, Romania. Journal of Human Evolution 45, 245–53.CrossRefGoogle ScholarPubMed
Trinkaus, E., Moldovan, O., Milota, S., Bîlgãr, A., Sarcinã, L., Athreya, S., Bailey, S.E., Rodrigo, R., Gherase, M., Higham, T., Bronk Ramsey, C. & Plicht, J. van der. 2003b. An early modern human from the Pestera cu Oase, Romania. Proceedings of the National Academy of Sciences USA 100, 11231–6CrossRefGoogle ScholarPubMed
Trinkaus, E., Ruff, C. B., Churchill, S.E. & Vandermeersch, B. 1998. Locomotion and body proportions of the Saint-Césaire 1 Châtelperronian Neandertal. Proceedings of the National Academy of Sciences USA 95, 5836–40CrossRefGoogle ScholarPubMed
Trinkaus, E., Zilhão, J., Rougier, H., Rodrigo, R., Milota, Ş., Gherase, M., Sarcinӑ, L., Moldovan, O., Baltean, I., Codrea, V., Bailey, S.E., Franciscus, R.G., Ponce de León, M. & Zollikofer, C.P. E. 2006. The Peştera cu Oase and early modern humans in southeastern Europe. In Conard, N.J. (ed.), When Neanderthals and Modern Humans Met, 145–64. Tübingen: Kerns VerlagGoogle Scholar
Vanhaeren, M. 2002. Les functions de la parure au Paléolithique supérieur: de l'individu à l‘umté culturelle. University of Bordeaux I: Ph. D. dissertationGoogle Scholar
Weaver, T.D. & Roseman, C.C. 2005. Ancient DNA, late Neandertal survival, and modern-human-Neandertal genetic admixture. Current Anthropology 46, 677–83CrossRefGoogle Scholar
Weninger, B. & Jöris, O. 2004. The Cologne Radiocarbon Calibration & Paleoclimate Research Package (http://www.calpal.de)Google Scholar
Wild, E.M., Teschler-Nicola, M., Kutschera, W., Steier, P., Trinkaus, E. & Wanek, W. 2005. Direct dating of Early Upper Palaeolithic human remains from Mladec̆. Nature 435, 332–5CrossRefGoogle ScholarPubMed
Willerslev, E. & Cooper, A. 2005. Ancient DNA. Proceedings of the Royal Society London B 272, 316Google ScholarPubMed
Wobst, M. 1974. Boundary conditions for palaeolithic social systems: a simulation approach. American Antiquity 39, 147–78CrossRefGoogle Scholar
Wobst, M. 1976. Locational relationships in Palaeolithic Society. Journal of Human Evolution 5, 4958CrossRefGoogle Scholar
Wolpoff, M. 2002. Human Paleontology. Ann Arbor: University of MichiganGoogle Scholar
Zilhão, J. 1993. Le passage du Paléolithique moyen au Paléolithique supérieur dans le Portugal. In Cabrera, V. (ed.), El Origen del Hombre Moderno en el Suroeste de Europa, 127–45. Madrid: Universidad Nacional de Educación a DistanciaGoogle Scholar
Zilhão, J. 1998. The extinction of Iberian Neandertals and its implications for the origins of modern humans in Europe. In Facchini, F., Palma di Cesnola, A., Piperno, M. & Peretto, C. (eds), XIII International Congress of Prehistoric and Protohistoric Sciences. Proceedings 2, 299312. Forlì: AbacoGoogle Scholar
Zilhão, J. 2000. The Ebro frontier: a model for the late extinction of Iberian Neanderthals. In Stringer, C., Barton, R.N.E. & Finlayson, C. (eds), Neanderthals on the Edge: 150th anniversary conference of the Forbes' Quarry discovery, Gibraltar, 111–21. Oxford, OxbowGoogle Scholar
Zilhão, J. 2001. Anatomically Archaic, Behaviorally Modern: the last Neanderthals and their destiny. Amsterdam: Stichting Nederlands Museum voor Anthropologic en PraehistoriaeGoogle Scholar
Zilhão, J. in press. The emergence of ornaments and art: an archaeological perspective on the origins of ‘behavioral modernity’. Journal of Archaeological ResearchGoogle Scholar
Zilhão, J. & d'Errico, F. 1999. The chronology and taphonomy of the earliest Aurignacian and its implications for the understanding of Neanderthal extinction. Journal of World Prehistory 13, 168CrossRefGoogle Scholar
Zilhão, J. & d'Errico, F. 2003. An Aurignacian ‘Garden of Eden’ in southern Germany? An alternative interpretation of the Geissenklösterle and a critique of the Kulturpumpe model. Paleo 15, 6986Google Scholar
Zilhão, J. & Trinkaus, E. 2002. Social implications. In Zilhão, & Trinkaus, (eds) 2002, 519–41Google Scholar
Zilhão, J. & Trinkaus, E. 2005. The earliest modern human settlement of Europe: anthropological and archaeological evidence. In Mellars, P., Stringer, C., Bar-Yosef, O. & Boyle, K. (eds), Rethinking the Human Revolution: new behavioral perspectives on the origins and dispersal of modern humans, 83–4. Cambridge: McDonald Institute for Archeological Research/American School of Prehistoric Research (book of abstracts of the conference, September 7–11)Google Scholar