Comparative morphology and paleobiology of MiddlePleistocene human remains from the Bau del’Aubesier, Vaucluse, FranceSerge Lebel*, Erik Trinkaus †‡§ , Martine Faure ¶ , Philippe Fernandez i , Claude Gue´rin i , Daniel Richter** †† ,Norbert Mercier ‡‡ , Hele`ne Valladas ‡‡ , and Gu¨nther A. Wagner***De´partement des Sciences de la Terre et de l’Atmosphe`re, Universite´ de Que´bec a` Montre´al, Casse Postale 8888, Succursale Centre Ville, Montre´al, QC,Canada H3C 3P8; † Department of Anthropology, Campus Box 1114, Washington University, St. Louis, MO 63130; ‡ Unite´ Mixte de Recherche 5809 duCentre National de la Recherche Scientifique, Laboratoire d’Anthropologie des Populations du Passe´, Universite´ de Bordeaux 1, 33405 Talence, France;¶ Unite´ Mixte de Recherche Pale´oenvironnements et Pale´obiosphe`re, Universite´ Lumie`re-Lyon 2, 7 Rue Raulin, 69007 Lyon, France; i Unite´ Mixte deRecherche Pale´oenvironnements et Pale´obiosphe`re du Centre National de la Recherche Scientifique, Unite´ de Formation et de Recherche desSciences de la Terre, Universite´ Claude Bernard-Lyon I, 27–43 Boulevard du 11 Novembre 1918, 69622 Villeurbanne, France; **ForschungsstelleArcha¨ometrie der Heidelberger Akademie der Wissenschaften am Max-Planck-Institut fu¨r Kernphysik, Postfach 10 39 80, 69029 Heidelberg,Germany; †† Instituto Tecnolo´gico e Nuclear, Estrada Nacional 10, 2686-953 Sacave´m, Portugal; and ‡‡ Laboratoire des Sciences du Climatet de l’Environnement, Avenue de la Terrasse, 91198 Gif-sur-Yvette, FranceContributed by Erik Trinkaus, July 10, 2001The discovery of later Middle Pleistocene human remains from theBau de l’Aubesier, France reinforces an evolutionary model of thegradual accumulation of Neandertal-derived facial and dentalfeatures during the Middle Pleistocene of the northwestern OldWorld. The pronounced maxillary incisor beveling of Aubesier 4helps to extend the antiquity of nondietary use of the anteriordentition. The interproximal ‘‘toothpick’’ groove on the Aubesier10 molar increases the sample for these lesions. The pathologicalloss of the mandibular dentition of Aubesier 11 indicates advancedantemortem masticatory impairment, at a level previously undoc-umented before the Late Pleistocene. These remains support aview of later Middle Pleistocene humans able to support debili-tated individuals despite the considerable use of their bodies toaccomplish routine activities.human paleontology u paleopathology u Neandertals u ArchaicHomo u EuropeIn 1975, the paleoanthropological record of the Middle Pleis-tocene was referred to as the ‘‘muddle in the middle’’ (1). Sincethat time, there have been major advances in our knowledge andperception of human biocultural evolution during the MiddlePleistocene. It has become increasingly recognized as the periodin human evolution during which several important aspects ofhuman biology and behavior evolved, including (among others)exaggerated encephalization, the controlled use of fire, temper-ate zone geographic dispersals, varieties of prepared core lithicreduction techniques, the development of effective (predatoryand defensive) weaponry, and regional differentiation of humanpopulations. At the same time, however, our paleoanthropologi-cal record for this period remains incomplete, and significantadditions to it have the potential to enlighten further ourperceptions of human evolutionary patterns.In the context of this research, recent discoveries of laterMiddle Pleistocene human remains at the Bau de l’Aubesier insoutheastern France, although limited anatomically, contributeto our perceptions of these later archaic humans. They provideadditional information relevant to the regional differentiation ofhuman populations during the Middle Pleistocene and thepaleobiological patterns of these archaic human groups. Inparticular, it has been argued that there was a gradual evolutionof a derived Neandertal morphological pattern through theMiddle Pleistocene in Europe and that this represents a periodof relative lineage stability combined with some degree ofgenetic isolation from other archaic human regional lineages inthe cul-de-sac of Europe (2, 3). It also has been noted that fewof these Middle Pleistocene human remains exhibit seriouspathological lesions (4, 5).The Bau de L’AubesierThe Bau de l’Aubesier is a large rock shelter in the Gorges dela Nesque, Monieux, Vaucluse, France (44° 059 N, 5° 219 E).Although previously investigated (6, 7), S.L. has excavated thesite since 1987 and discovered Middle Pleistocene human re-mains in 1994 and 2000.The upper portion of the Pleistocene deposits (13 m thick over.520 m 2 ) consists principally of Late Pleistocene levels contain-ing Middle Paleolithic archeological remains (Fig. 1). The lowerportion of the site, with a thickness . 10 m, derives primarilyfrom the Middle Pleistocene (levels G to K). Its succession ofstratifieddepositscontaininglimestonee ´boulis,blocks,andslabsis interspersed with a nonsystematic matrix of fine interstitialcalcareous sediment, much of which is brecciated in layer G. Thearcheological layer H-1, 55 cm thick, consists of a high densityof faunal remains, lithics, wood charcoal, and ash in a highlyorganic sediment, and the lithics and faunal remains are burnt todifferent degrees. On the basis of the lithostratigraphy, thereappears to have been a chronological hiatus between layer H-1and the underlying I series.The layers with human remains, I-2, I-3, and K-1, yielded highconcentrations of large herbivore remains and flint artifacts,each a palimpsest of repeated phases of site occupation. Theirlithic assemblages are typical Mousterian with abundant side-scrapers, Levallois flakes, and especially Levallois blades, similarto those from a growing number of late Middle PleistoceneMiddle Paleolithic site levels containing laminar assembla-ges (8).The fauna of level I includes Ursus arctos, Dicerorhinushemitoechus, Equus mosbachensis, Sus scrofa, Cervus elaphus,Dama dama, Capreolus capreolus, Megaceros giganteus, Bos pri-migenius, Hemitragus cedrensis, and Rupicapra rupicapra, withBos (43%) and Equus (31%) dominating. Well recognized in theMiddle Pleistocene, E. mosbachensis in particular indicates anagenolaterthanoxygenisotopestage(OIS)6(9);itissucceededbyE.cf.taubachensisintheoverlyinglayerH-1,whichcouldreferit to OIS 5e (9). The form of C. elaphus can be assigned to OISAbbreviations: ka, thousand years; OIS, oxygen isotope stage; AT-SH, Atapuerca–Sima delos Huesos.§ To whom reprint requests should be addressed. E-mail: trinkaus@artsci.wustl.edu.The publication costs of this article were defrayed in part by page charge payment. Thisarticle must therefore be hereby marked “advertisement” in accordance with 18 U.S.C.§1734 solely to indicate this fact.www.pnas.orgycgiydoiy10.1073ypnas.181353998 PNAS u September 25, 2001 u vol. 98 u no. 20 u 11097–11102ANTHROPOLOGY