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The Atapuerca Sites and Their Contribution to the Knowledge of Human Evolution in Europe J

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Evolutionary Anthropology 13:25–41 (2004) ARTICLES The Atapuerca Sites and Their Contribution to the Knowledge of Human Evolution in Europe J. M. BERMU´ DEZ DE CASTRO, M. MARTINO´ N-TORRES, E. CARBONELL, S. SARMIENTO, A. ROSAS, J. VAN DER MADE, AND M. LOZANO Over the last two decades, the Pleistocene sites of the Sierra de Atapuerca use of terms such as “advanced,” (Spain) have provided two extraordinary assemblages of hominin fossils that have “primitive,” “early archaic,” “archaic,” helped refine the evolutionary story of the genus Homo in Europe. The TD6 level of and “late archaic” to modify Homo the Gran Dolina site has yielded about one hundred remains belonging to a sapiens. These terms and other similar minimum of six individuals of the species Homo antecessor. These fossils, dated ones2,3 have no taxonomical validity, to the end of the Lower Pleistocene (800 kyr), provide the earliest evidence of but suggest a need to describe mor- hominin presence in Western Europe. The origin of these hominins is unknown, but phological diversity. they may represent a speciation event from Homo ergaster/Homo erectus. The The cladogenetic model also has TD6 fossils are characterized by a significant increase in cranial capacity as well as difficulties. Cladistics requires the es- the appearance of a “sapiens” pattern of craniofacial architecture. At the Sima de tablishment of character polarities los Huesos site, more than 4,000 human fossils belonging to a minimum of 28 (the phylogeny of the characters), the individuals of a Middle Pleistocene population (ca. 500–400 kyr) have been re- selection of independent traits, and covered. These hominins document some of the oldest evidence of the European the identification of homoplasies. roots of Neanderthals deep in the Middle Pleistocene. Their origin would be the These difficulties have opened several dispersal out of Africa of a hominin group carrying Mode 2 technologies to Europe. debates concerning the recognition of Comparative study of the TD6 and Sima de la Huesos hominins suggests a one or more species in a given hyp- replacement model for the European Lower Pleistocene population of Europe or odigm. One of the most prominent ex- interbreeding between this population and the new African emigrants. amples concerns the possible parti- tion of a large portion of the Early Pleistocene fossil record into two spe- Since the late 1980s, paleoanthro- the genus Homo.1 The anagenetic evo- cies, Homo erectus, Dubois 1892, and pology has witnessed a paradigm shift lutionary model, which postulate an Homo ergaster, Groves and Mazak in interpretation of the evolution of unbroken lineal temporal succession 1975. Homo ergaster would be re- of the reproductive continuity of stricted to Africa (if we consider the Homo habilis, Homo erectus, and hominins found in the Dmanisi site, Homo sapiens, representing three evo- Republic of Georgia, to belong to a Jose´ Marı´a Bermu´ dez de Castro is Re- lutionary grades, is gradually being different species4), while Homo erec- search Professor at the Consejo Superior replaced by the cladogenetic model of tus may have representatives in Asia de Investigaciones Cientı´ficas(CSIC), Spain 1 and co-director of the Atapuerca Research speciation, which recognizes several and Africa. The distinctions are Project. E-mail: [email protected] speciation events throughout the evo- based on the absence of some specific Eudald Carbonell is Professor at the Uni- lution of the genus Homo. These traits in the African fossils that are versity Rovira i Virgili of Tarragona, Spain and co-director of the Atapuerca Re- events (cladogenesis) might have been present in the Asian ones, to which the search Project. favored by successive dispersals of species name erectus applies. These Marı´a Martino´ n-Torres, Susana Sarmiento and Marina Lozano are researchers with hominins out of Africa and migratory include a midline keel on the vault, the Atapuerca team, having a primary inter- movements between Africa and Eur- strong reduction of the postglenoid est in dental anthropology. asia during the Pleistocene, as well as process, an angular torus at the pos- Antonio Rosas is Research Fellow at the CSIC, Spain and member of the Atapu- events of reproductive isolation due to terior-inferior corner of the parietal erca Research Team. climatic and ecological changes. bone, overall thickening of the brain- Jan van der Made is Research Fellow at the CSIC, Spain and member of the Ata- The anagenetic model divides the case, and reduced superstructures in puerca Research Team. fossil record, or the morphological the temporal-occipital region. More- space of the genus Homo, into three over, Homo erectus ss exhibits a Key words: Homo antecessor; ergaster; heidel- evolutionary grades. As the morphos- styloid process, as well as differences bergensis; Neanderthals pace is filled with new findings, it is in the size and shape of the supraor- becoming increasingly difficult to bital torus.5–7 The debate about the © 2004 Wiley-Liss, Inc. agree on the exact boundaries be- partitioning of the fossil record is be- DOI 10.1002/evan.10130 Published online in Wiley InterScience tween species. The difficulties of this yond the scope of this paper but, from (www.interscience.wiley.com). model are exemplified by the frequent this point on, we assign the specimens 26 Bermu´dez de Castro et al. ARTICLES THE SIERRA DE ATAPUERCA SITES The Sierra de Atapuerca is a small hill of about twelve square kilometers that rises 1,079 meters above sea level and approximately 100 meters above the alluvial flatness of the Arlanzo´n river. The Sierra de Atapuerca is lo- cated in northern central Spain (Du- ero Basin) fourteen kilometers east of the historical city of Burgos. It con- tains numerous karst cavities in Cre- taceous limestones that are filled with Pleistocene sediments (Fig. 1). The Sima de los Huesos is a blind cavity of 8 ϫ 4m2 located well inside the Cueva Mayor-Cueva del Silo cave system of the karst of Sierra de Ata- puerca (Fig. 1). The access to this chamber is a vertical conduit 14 m deep, which ends with a steep inclined ramp about 9 m long. The first human remains, associated with a large as- semblage of cave-bear (Ursus deningeri) fossils, were found here in 1976.12 This site has been systemati- cally excavated since 1984. Over sev- eral decades, tons of sediment were turned over by cavers hunting for cave-bear teeth.13 A meticulous and methodical excavation began in Figure 1. Map of the Trinchera del Ferrocarril and Cueva Mayor, showing the location of the 1988.14 Arsuaga and coworkers14 de- main Atapuerca sites. TD: Gran Dolina; TG: Galerı´a; TN: Trinchera Norte; TZ: Covacha de los scribed the stratigraphy and many Zarpazos; TE: Sima del Elefante; SH: Sima de los Huesos. other aspects of the Sima de los Hue- sos site and other nearby cavities. They also published cross-sectional of the African Early Pleistocene, in- Despite the internal debate gener- diagrams of the Sima de los Huesos cluding KNM-ER 992 (the holotype of ated by the cladogenetic model, most site and a map of the excavation grid. the species), KNM-ER 3733, KNM-ER authors have abandoned the gradual- The stratigraphy, geological history, 3883, and KNW-WT 15000 (the Tur- ist model for the “phylogenetic species and preliminary dating of the site can kana boy) to the species Homo er- concept” and the recognition of a be found in Bischoff and coworkers.15 gaster. larger number of species in the fossil All the human fossils were recovered Most authors suggest the use of record of the genus Homo. This situ- from the same unit, which is shaped apomorphies (unique derived traits) ation has triggered an ongoing debate by breccias of clay-supporting bones, for the recognition of a species in a about the number of cladogenetic blocks, and clasts15 of varying thick- strict application of the cladogenetic events, the nomenclature of the iden- ness along the site’s profile. Appar- model.8 However, trait exclusiveness tified species, and possible evolution- ently, all fossils were deposited in the is not as common in the fossil record ary scenarios. For the past thirty site during the same sedimentation as it might seem. Given this situation, years, the research team of the Sierra period.15 criteria that are less strict, such as the de Atapuerca sites (Spain) has con- Previous radiometric and paleo- presence of a unique combination of tributed to this debate through the magnetic analyses, as well as biochro- apomorphic (derived) and plesiomor- discovery of two hominin fossil as- nological data, pointed to an age of phic (primitive) traits frequently has semblages from the Sima de los Hue- 200 to 320 kyr (oxygen isotope stages been used.9,10 Furthermore, due to the sos and Gran Dolina sites. These as- 7 to 9) for the Sima de los Huesos difficulties we have noted, some au- semblages, as well as other specimens hominins.15–17 However, recent radio- thors consider that the species diag- discovered in the last decade in Africa metric studies (U-series) of a 14-cm- nosis does not need a cladistic basis, and Europe, give insight into the rela- thick in-situ speleothem overlying the but can be made through the phenetic tionship between Early Pleistocene mud-breccia containing the human determination of a distinctive mor- Homo and the lineage that gave rise to bones has provided a minimum age of photype.11 Homo sapiens. 350 kyr for these hominins.18 An esti- ARTICLES The Atapuerca Sites 27 mated age of 400 to 500 kyr (oxygen continuous from TD3 to TD11 and per part of TD6, the Aurora stratum, isotope stages 12 to 14) for these some levels (TD4, 5, 6, 7, 10, and 11) corresponds to a Mediterranean forest hominins is based on the rate of contain abundant stone tools.
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