J.M. Bermúdez de Castro

Lower Pleistocene hominids and artifacts from Atapuerca-TD6 (Spain)

Human remains dating to more than 780,000 years ago are associated with a rich faunal and lithic assemblage in the Pleistocene cave site of Gran Dolina (TD), Sierra de Atapuerca, Burgos, Spain. The micromammal species represent the late Biharian (Mimomys savini zone), and the lithic objects represent pre-Acheulean technology (Mode 1) and comes from the TD6 level below the Matuyama-Brunhes boundary. The Gran Dolina hominid fossils cannot be comfortably accommodated in any of the defined Homo species. They could be considered a primitive form of Homo heidelbergensis, but a new species might be named in the future if the sample is enlarged. The new human fossil evidence demonstrates that Western Europe was settled at least since the late early Pleistocene.

Luminescence chronology of cave sediments at the Atapuerca paleoanthropological site, Spain

Ascertaining the timing of the peopling of Europe, after the first out-of-Africa demographic expansion at the end of the Pliocene, is of great interest to paleoanthropologists. One of the earliest direct evidences for fossil hominins in western Europe comes from an infilled karstic cave site called Gran Dolina at Atapuerca, in a stratum approximately 1.5m below the Brunhes-Matuyama (B-M) geomagnetic boundary (780ka) within lithostratigraphic unit TD6. However, most of the meters of fossil- and tool-bearing strata at Gran Dolina have been difficult to date. Therefore, we applied both thermoluminescence (TL) and infrared-stimulated-luminescence (IRSL) multi-aliquot dating methods to fine-silt fractions from sediment samples within Gran Dolina and the nearby Galería cave site. We also applied these methods to samples from the present-day surface soils on the surrounding limestone hill slopes to test the luminescence-clock-zeroing-by-daylight assumption. Within the uppermost 4m of the cave deposits at Gran Dolina, TL and paired TL and IRSL ages range stratigraphically from 198+/-19ka to 244+/-26ka. Throughout Gran Dolina, all luminescence results are stratigraphically self-consistent and, excepting results from two stratigraphic units, are consistent with prior ESR-U-series ages from progressively deeper strata. Thermoluminescence ages culminate at 960+/-120ka approximately 1m below the 780ka B-M boundary. At Galería, with one exception, TL and IRSL ages range stratigraphically downward from 185+/-26ka to 503+/-95ka at the base of the lowermost surface-inwash facies. These results indicate that TL and (sometimes) IRSL are useful dating tools for karstic inwash sediments older than ca. 100ka, and that a more accurate chronostratigraphic correlation is now possible among the main Atapuerca sites (Gran Dolina, Galería, Sima de los Huesos). Furthermore, the oldest TL age of ca. 960ka from Gran Dolina, consistent with biostratigraphic and paleomagnetic evidence, implies a probable numeric age of 900-950ka for the oldest hominin remains ( approximately 0.8m below the TL sample). This age window suggests a correspondence to Marine Isotope Stage (MIS) 25, a relatively warm and humid interglaciation.

Neandertal roots: Cranial and chronological evidence from Sima de los Huesos

Neandertal ancestors from Pleistocene Spain The Sima de los Huesos site in Atapuerca, northern Spain, is a rich source of fossil hominin specimens. The site has now yielded further skull specimens that illuminate patterns of human evolution in Europe nearly half a million years ago. Arsuaga et al. studied 17 crania, including 7 that are new specimens and 6 that are more complete than before (see the Perspective by Hublin). This assemblage of specimens reveals the cranial, facial, and dental features of the Atapuerca hominins, which allows more precise evolutionary positioning of these Neandertal ancestors. Science, this issue p. 1358; see also p. 1338 Seventeen skulls from at least 430 thousand years ago illuminate hominin evolutionary patterns in Pleistocene Europe. [Also see Perspective by Hublin] Seventeen Middle Pleistocene crania from the Sima de los Huesos site (Atapuerca, Spain) are analyzed, including seven new specimens. This sample makes it possible to thoroughly characterize a Middle Pleistocene hominin paleodeme and to address hypotheses about the origin and evolution of the Neandertals. Using a variety of techniques, the hominin-bearing layer could be reassigned to a period around 430,000 years ago. The sample shows a consistent morphological pattern with derived Neandertal features present in the face and anterior vault, many of which are related to the masticatory apparatus. This suggests that facial modification was the first step in the evolution of the Neandertal lineage, pointing to a mosaic pattern of evolution, with different anatomical and functional modules evolving at different rates.

Dental evidence on the hominin dispersals during the Pleistocene

A common assumption in the evolutionary scenario of the first Eurasian hominin populations is that they all had an African origin. This assumption also seems to apply for the Early and Middle Pleistocene populations, whose presence in Europe has been largely explained by a discontinuous flow of African emigrant waves. Only recently, some voices have speculated about the possibility of Asia being a center of speciation. However, no hard evidence has been presented to support this hypothesis. We present evidence from the most complete and up-to-date analysis of the hominin permanent dentition from Africa and Eurasia. The results show important morphological differences between the hominins found in both continents during the Pleistocene, suggesting that their evolutionary courses were relatively independent. We propose that the genetic impact of Asia in the colonization of Europe during the Early and Middle Pleistocene was stronger than that of Africa.

A new early Pleistocene hominin mandible from Atapuerca-TD6, Spain

We present the description of a new mandibular specimen, ATD6-113, recovered in 2006 from the TD6 level of the Gran Dolina cave site in Sierra de Atapuerca, northern Spain. A detailed study of the lithostratigraphy of the top sequence of this level, the section from where all human remains have been recovered so far, is also presented. We have observed that the hominin stratum, previously defined as Aurora Stratum, represents a condensed deposit of at least six layers, which could not be distinguished in the test pit made in 1994-95. Therefore, the human fossil remains were probably deposited during a discrete and undetermined time period. The new mandibular fragment exhibits a very similar morphology to that of the most complete specimen, ATD6-96, which was recovered in 2003 from a different layer. This suggests that both specimens represent the same biological population. The two mandibles, as well as the small mandibular fragment ATD6-5 (which constitutes part of the holotype of Homo antecessor), present a morphological pattern clearly derived with regard to that of the African early Homo specimens usually included in H. habilis and H. rudolfensis, the mandibles D211 and D2735 from Dmanisi, and most of the early Pleistocene mandibles from Sangiran. The TD6 mandibles also exhibit some derived features with regard to the African early Pleistocene specimens included in H. ergaster (or African H. erectus). Thus, the TD6 hominins seem to represent a lineage different from other African and Asian lineages, although some (metric in particular) similarities with Chinese middle Pleistocene mandibles are noted. Interestingly, none of the apomorphic mandibular features of the European middle and early late Pleistocene hominins are present in the TD6 mandibles.

Gran Dolina-TD6 versus Sima de los Huesos dental samples from Atapuerca: evidence of discontinuity in the European Pleistocene population?

Abstract Comparative analyses between Gran Dolina-TD6 and Sima de los Huesos (SH) dental samples from Sierra de Atapuerca (Spain) suggest that hominins represented by these samples belonged to two very different populations and even to distinct paleospecies. Gran Dolina-TD6 hominins ( Homo antecessor ) have been dated to about 0.8 million years ago (mya), whereas a new radiometric dating of the SH hominins ( Homo heidelbergensis ) suggests an interval of 0.4–0.5 mya for these hominins. Current results as well as the relative temporal closeness between the populations represented by Gran Dolina-TD6 and SH favour a replacement scenario hypothesis (or possibly a crossbreeding scenario) for the Europeans during the early Middle Pleistocene. Although the information available from the European Lower Pleistocene populations is limited (Ceprano and Gran Dolina-TD6), current data are noteworthy and their combination with archaeological evidence will stimulate future discussions on the dynamics of the first European settlements.

NONOCCLUSAL DENTAL MICROWEAR ANALYSIS OF 300,000-YEAR-OLD HOMO HEILDERBERGENSIS TEETH FROM SIMA DE LOS HUESOS (SIERRA DE ATAPUERCA, SPAIN)

Casts of nonocclusal enamel surfaces of 190 teeth from the Middle Pleistocene site of Sima de los Huesos have been micrographed by scanning electron microscopy. Microscopic analyses of striation density and length by orientation show distinct patterns of intrapopulation variability. Significant differences in the number and length of the striations by orientation are found between maxillary and mandibular teeth. This probably reflects differences in the mechanical forces involved in the process of chewing food. Significant differences are present between isolated and in situ teeth that could be caused by postdepositional processes differentially affecting the isolated teeth. In addition, a distinct and very unusual striation pattern is observed in a sample of teeth that can be explained only by a strong nondietary, most probably postmortem abrasion of the enamel surfaces. These teeth have a very high density of scratches, shorter in length than those found on other teeth, that are not indicative of dietary habits. No known depositional process may account for the presence of such postmortem wear since heavy transportation of materials within the clayish sediments has been discarded for the site. Despite this, a characteristic dietary striation pattern can be observed in most of the teeth analyzed. Most likely the diet of the Homo heidelbergensis hominids from Sima de los Huesos was highly abrasive, probably with a large dependence on hard, poorly processed plant foods, such as roots, stems, and seeds. A highly significant sex-related difference in the striation pattern can also be observed in the teeth analyzed, suggesting a differential consistency in the foods eaten by females and males.

The Atapuerca sites and the ibeas hominids

The Atapuerca railway Trench and Ibeas sites near Burgos, Spain, are cave fillings that include a series of deposits ranging from below the Matuyama/Bruhnes reversal up to the end of Middle Pleistocene. The lowest fossil-bearing bed in the Trench contains an assemblage of large and small Mammals includingMimomys savini, Pitymys gregaloides, Pliomys episcopalis, Crocuta crocuta, Dama sp. and Megacerini; the uppermost assemblage includesCanis lupus, Lynx spelaea, Panthera (Leo) fossilis, Felis sylvestris, Equus caballus steinbeimensis, E.c. germanicus, Pitymys subterraneus, Microtus arvalis agrestis, Pliomys lenki, and alsoPanthera toscana, Dicerorbinus hemitoechus, Bison schoetensacki, which are equally present in the lowest level. The biostratigraphic correlation and dates of the sites are briefly discussed, as are the paleoclimatic interpretation of the Trench sequences. Stone artifacts are found in several layers; the earliest occurrences correspond to the upper beds containingMimomys savini. A set of preserved human occupation floors has been excavated in the top fossil-bearing beds. The stone-tool assemblages of the upper levels are of upper-medial Acheulean to Charentian tradition. The rich bone breccia SH, in the Cueva Mayor-Cueva del Silo, Ibeas de Juarros, is a derived deposit, due to a mud flow that dispersed and carried the skeletons of many carnivores and humans. The taxa represented are:Ursus deningeri (largely dominant),Panthera (Leo) fossilis, Vulpes vulpes, Homo sapiens var. Several traits of both mandibular and cranial remains are summarized. Preliminary attempts at dating suggest that the Ibeas fossil man is older than the Last Interglacial, or oxygen-isotope stage 5.

The human remains from Cova Negra (Valencia, Spain) and their place in European Pleistocene human evolution

Abstract The phylogenetic and chronological positions of the Cova Negra human remains are discussed. These remains include an adult right parietal bone, a juvenile right mandibular body fragment with a deciduous second molar, and an isolated permanent upper central incisor. There are 40 levels in the Cova Negra sedimentary sequence, ranging from the Riss-Wurm to the beginning of the Wurm III. The Cova Negra A phase (Riss-Wurm) is faunally and industrially sterile. Several variants of the Mousterian (Charentian type) are known from the beginning of the Wurm (Cova Negra B phase) to the end of the sequence (Cova Negra E and F phases). All human remains are assigned to the Wurm. On the basis of the parietal bone, uni- and bivariate analyses show that the Cova Negra biparietal vault was very broad. The cluster analysis groups Cova Negra with the biggest Neandertal neurocrania (specimens generally sexed as males). Also, this parietal bone exhibits a subcircular profile in posterior view, a trait considered as a Neandertal autapomorphy. The mandibular fragment shows three foramina mentalia and a steep linea mylohyoidea. These and other traits lead us to include this specimen in the Neandertal group. The crown dimensions and the morphology of the deciduous molar are consistent with this taxonomic attribution. The comparative study of the crown and root dimensions of the permanent incisor, as well as the presence of traits such as a strong tuberculum linguale, well developed marginal ridges (shovelling), and the marked lingual inclination of the buccal face, lead us to attribute this tooth to the Neandertal group.

Gran Dolina-TD6 and Sima de los Huesos dental samples: Preliminary approach to some dental characters of interest for phylogenetic studies

The Sima de los Huesos (SH) and Gran Dolina-TD6 sites in Sierra de Atapuerca (Spain) have each yielded an impressive fossil hominin sample representing Middle Pleistocene and Late Lower Pleistocene European populations, respectively. Paleontological evidence, paleomagnetic analyses, and radiometric dates (U/Th) suggest an interval of 400 to 500 ky for the SH hominins. At Gran Dolina, radiometric dates (ESR and U-series) combined with paleomagnetic analyses and fossil evidence indicate an age range between 780 and 860 ky for the Aurora Stratum of the TD6 level where the fossil hominins were found. We have assigned the SH hominins to the Homo heidelbergensis species, whereas the TD6 hominins are representative of Homo antecessor, the species named in 1997 (Bermudez de Castro et al., 1997) to accommodate the variability observed in the TD6 fossil human assemblage. Dental collections of the SH and TD6 sites include more than five hundred deciduous and permanent teeth. The detailed description and morphological comparison of the Atapuerca dental samples will be published elsewhere in a near future, but the examination of an extensive human fossil record, has already revealed some dental characters we consider crucial for phylogenetic studies. We describe those characters and provide an overview of their distribution across the hominin fossil dental record. On the basis of these traits we explore some questions about the phylogenetic relationship between TD6 and SH hominins as well as the evolutionary scenario of these two populations.