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Paleobiology and Comparative Morphology of a Late Neandertal Sample from El Sidro´N, Asturias, Spain

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Paleobiology and Comparative Morphology of a Late Neandertal Sample from El Sidro´N, Asturias, Spain Paleobiology and comparative morphology of a late Neandertal sample from El Sidro´n, Asturias, Spain Antonio Rosasa,b, Cayetana Martı´nez-Mazaa, Markus Bastira,c, Antonio Garcı´a-Taberneroa, Carles Lalueza-Foxd, Rosa Huguete, Jose´ Eugenio Ortizf, Ramo´ n Julia` g, Vicente Solerh, Trinidad de Torresf, Enrique Martı´nezi, Juan Carlos Can˜ averasj, Sergio Sa´ nchez-Moralk, Soledad Cuezvak, Javier Lariol, David Santamarı´am, Marco de la Rasillam, and Javier Forteam aDepartamento de Paleobiologı´a,Museo Nacional de Ciencias Naturales, Consejo Superior de Investigaciones Cientificas (CSIC), Calle Jose´Gutierrez Abascal 2, 28006 Madrid, Spain; dDepartament de Biologia Animal, Facultat de Biologia, Universitat de Barcelona, Avenida Diagonal 645, 08028 Barcelona, Spain; eA´ rea de Prehistoria, Universitat Rovira i Virgili, Plac¸a Imperial Tarraco 1, 43005 Tarragona, Spain; fEscuela Te´cnica Superior de Ingenieros de Minas, Universidad Polite´cnica de Madrid, Calle Rı´osRosas 21, 28003 Madrid, Spain; gInstituto Jaume Almera, CSIC, Calle Lluı´sSole´i Sabarı´ss/n, 08028 Barcelona, Spain; hInstituto de Productos Naturales, CSIC, Avenida Astrofı´sicoFrancisco Sa´nchez 3, 38206 Santa Cruz de Tenerife, Spain; iDepartamento de Geologı´a, Universidad de Oviedo, Calle Jesu´s Arias de Velasco s/n, 33005 Oviedo, Spain; jDepartamento de Ciencias de la Tierra, Universidad de Alicante, Apartado de Correos 99, 03080 Alicante, Spain; kDepartamento de Geologı´a,Museo Nacional de Ciencias Naturales, CSIC, Calle Jose´Gutierrez Abascal 2, 28006 Madrid, Spain; mDepartamento de Historia, Universidad de Oviedo, Calle Teniente Alfonso Martı´nezs/n, 33011 Oviedo, Spain; cHull–York Medical School, University of York, Heslington, YO 105DD, England; and lDepartamento de Ciencias Analı´ticas,Universidad Nacional de Educacio´n Distancia, Paseo Senda del Rey 9, 28040 Madrid, Spain Communicated by Erik Trinkaus, Washington University, St. Louis, MO, November 4, 2006 (received for review September 21, 2006) Fossil evidence from the Iberian Peninsula is essential for understand- area. The large sample from El Sidro´n provides a portrait of ing Neandertal evolution and history. Since 2000, a new sample those late Neandertals living in the Cantabrian range. Ϸ43,000 years old has been systematically recovered at the El Sidro´n cave site (Asturias, Spain). Human remains almost exclusively com- El Sidro´n Site pose the bone assemblage. All of the skeletal parts are preserved, and The accidental unearthing in 1994 of an outstanding set of there is a moderate occurrence of Middle Paleolithic stone tools. A human fossils gave rise to the archeological excavation and minimum number of eight individuals are represented, and ancient multidisciplinary study of the site (11). As a result, a significant mtDNA has been extracted from dental and osteological remains. archeopaleontological record is being recovered, largely com- Paleobiology of the El Sidro´n archaic humans fits the pattern found posed of Neandertal remains. The karstic site of El Sidro´n is in other Neandertal samples: a high incidence of dental hypoplasia located in the region of Asturias, Spain, (43°23Ј01ЉN, 5°19Ј and interproximal grooves, yet no traumatic lesions are present. 44ЉW) (Fig. 1). Geologically, the site is found in the so-called Moreover, unambiguous evidence of human-induced modifications ‘‘Surco Oviedo-Infiesto’’ (11), a strip of Mesozoic and Ceno- has been found on the human remains. Morphologically, the El Sidro´n zoic sediments limited by Paleozoic relief to the north and humans show a large number of Neandertal lineage-derived features south. The cavity is a pression tube of Ϸ600 m long, with a even though certain traits place the sample at the limits of Neandertal central stretch of 200 m oriented nearly west–east (‘‘Galerı´a variation. Integrating the El Sidro´n human mandibles into the larger del Rı´o’’). This tube shows on its southern bank transverse Neandertal sample reveals a north–south geographic patterning, galleries in a NE–SW to N–S direction, generally of a restricted with southern Neandertals showing broader faces with increased nature. The fossil site is located in one of these transverse lower facial heights. The large El Sidro´n sample therefore augments galleries, the Osario Gallery, of Ϸ28 m long and 12 m at its the European evolutionary lineage fossil record and supports eco- widest part (Fig. 1). All of the remains are recovered from a geographical variability across Neandertal populations. surface Յ6m2 within stratum III of the sedimentary sequence. Sediments accumulated in the Osario Gallery constitute a dental hypoplasia ͉ geographic patterning ͉ geometric morphometrics ͉ relatively thin deposit, so far prospected at a maximum mandible ͉ Neandertal diversity thickness of 227 cm. The archeological assemblage recovered at the site is in eandertal morphology evolved in the northwestern corner secondary position, and it certainly comes from a close exterior Nof the Old World through a long evolutionary process, location. Ex hypothesis, the original deposit was located outside whose fossil evidence is present through the European Middle the cavity, possibly in a doline close to the vertical of the site. and Late Pleistocene geological record (1–6). In this process, A collapse of nearby fissures produced the sudden entry of the variation across the geographical range of Neandertals archeological material in a single event. Several taphonomic through its evolutionary history should be evident. The recov- signals help to clarify the scenario. Refitting of several bone ery of mtDNA in an increasingly number of early Late fragments and 53 stone tools indicates a limited displacement Pleistocene specimens is beginning to document an emerging as well as synchrony of the assemblage. Preservation of Neandertal phylogeographic pattern (7–9), and this pattern osteological surfaces is excellent with very limited trampling may be present as well in their skeletal morphology. and erosion. There are no toothmarks of large carnivores on The paleoanthropological collection presently retrieved at the bones, with marginal action of rodents and a small the El Sidro´n cave site (Asturias, Spain) represents the most carnivore (e.g., fox) on a few nonhuman remains. A few bones significant Neandertal sample in the Iberian Peninsula, and it allows further insight into these evolutionary processes and Author contributions: A.R. and J.F. designed research; A.R., C.M.-M., M.B., A.G.-T., C.L.-F., diversification of Middle Paleolithic populations across geo- R.H., J.E.O., R.J., V.S., T.d.T., E.M., J.C.C., S.S.-M., S.C., J.L., D.S., M.d.l.R., and J.F. performed graphic regions. At a local scale, the El Sidro´n site fulfills a research; C.M.-M., M.B., A.G.-T., C.L.-F., R.H., J.E.O., R.J., V.S., T.d.T., E.M., J.C.C., S.S.-M., S.C., significant gap in the fossil record of the Cantabrian region. A J.L., D.S., M.d.l.R., and J.F. analyzed data; and A.R. wrote the paper. long tradition of Paleolithic research in the region (10) has The authors declare no conflict of interest. yielded a rich Pleistocene cultural record (e.g., El Castillo, La Abbreviation: EMP, European Middle Pleistocene. Vin˜a, El Pendo, Morı´n), but the scarcity of human remains has bTo whom correspondence should be addressed. E-mail: [email protected]. precluded a characterization of the humans inhabiting the © 2006 by The National Academy of Sciences of the USA 19266–19271 ͉ PNAS ͉ December 19, 2006 ͉ vol. 103 ͉ no. 51 www.pnas.org͞cgi͞doi͞10.1073͞pnas.0609662104 Downloaded by guest on September 28, 2021 Fig. 1. Localization of the El Sidro´n site in Asturias (Spain). A map of the cave system with the Osario Gallery show the excavated area. have hydraulic abrasion but no weathering. In short, the data yielded a 48-bp sequence (between positions 16,230 and 16,278 of point to a limited exposure of bones outside the cavity; a mass the mtDNA reference sequence), and the second yielded an almost displacement moved the archeological deposit into the cave complete mtDNA hypervariable region 1 (302-nt sequence, be- secondarily, with very little movement after final deposition. tween positions 16,076 and 16,378). The sequences are identical in the overlapping 48-bp segment, and thus it cannot be discarded Dating. Three human specimens were 14C accelerator mass from the genetic data that they belong to the same individual. spectrometer-dated: SD-500 (tooth), 40,840 Ϯ 1,200 14C B.P. (Beta 192065); SD-599a (bone), 37,300 Ϯ 830 14C B.P. (Beta Lithics. A total of 333 lithic artifacts have been recovered, including 192066); and SD-763a (tooth), 38,240 Ϯ 890 14C B.P. (Beta side scrapers, denticulates, a hand axe, and several Levallois points. 192067). Calibrated with CalPal (www.calpal.de), the dates The raw material comes from the immediate cave environment, and become 44,310 Ϯ 978, 42,320 Ϯ 367, and 42,757 Ϯ 464 cal B.P., it is mostly chert, and quartzite in a lesser proportion. The lithic respectively, the average calibrated age being 43,129 Ϯ 129 cal assemblage is largely flake-based, with a few laminar products and B.P. (7). A similar age was obtained from preliminary of amino some Levallois supports. Also, up to 17 flakes, some of them Ϯ acid razemization (11), both from gastropods (39,000 7,000) retouched, have been refitted on a core fragment, reflecting Ϯ and human fossils 32,000 11,000). knapping at the primary location of the assemblage. Paleontological Sample. A series of Ϸ1,323 human remains has been Morphological Affinities of the Human Remains recovered at El Sidro´n (Table 1). The collection has been divided The El Sidro´n teeth are large, with crenulated enamel and acces- into the 140 specimens unprofessionally unearthed (labeled SDR) sory cusps.
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