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Gran Dolina is a cavity infilled by at least 25 m of Pleistocene sediments. This sequence contains the TD6 stratigraphic unit, whose records include around 170 hominin bones that have allowed the definition of a new species, Homo antecessor. This fossil accumulation was studied as a single assemblage and interpreted as a succession of several human home bases. We propose a complete stratigraphic context and sedimentological interpretation for TD6, analyzing the relationships between the sedimentary facies, the clasts and archaeo-palaeontological remains. The TD6 unit has been divided into three sub-units and 13 layers. Nine sedimentary facies have been defined. Hominin remains appear related to three different sedimentary facies: debris flow facies, channel facies and floodplain facies. They show three kinds of distribution: first a group of scattered fossils, then a group with layers of fossils in fluvial facies, and third a group with a layer of fossils in mixed fluvial and gravity flow facies. The results of this work suggest that some of these hominin remains accumulated in the cave by geological processes, coming from the adjacent slope above the cave or the cave entry, as the palaeogeography and sedimentary characteristics of these allochthonous facies suggest.

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., (Copyright © 2014, American Association for the Advancement of Science.)

The aim of this study is to describe the morphology of the roots and root canals of permanent lower second premolars (LP4s) with fully developed roots of five hominin groups: Homo sp. (ATE9-1 specimen) from Atapuerca-Sima del Elefante locality, H. antecessor (ATD6-4 and ATD6-125) from Atapuerca-Gran Dolina TD6 locality, H. heidelbergensis from Atapuerca-Sima de los Huesos locality, H. neanderthalensis from Krapina, Regourdou, and Abri Bourgeois-Delaunay localities, and two contemporary H. sapiens groups. The teeth were scanned by means of microtomography. The roots were divided into three virtual segments by three planes: cemento-enamel junction (CEJ), mid-root (MR), and mid-apex (MA). Volumetric and planar direct measurements of the whole teeth and each segment were taken. Descriptive statistical analyses and nonparametric Mann-Whiney test were performed to test for significant differences (P < 0.025) between groups. ATE9-1 and Gran Dolina-TD6 fossils present intricate radicular complexes that might be transitional between the morphologies of Australopithecus robustus and African early Homo and the derived conditions typically found in later Homo. In H. neanderthalensis and H. heidelbergensis, the root canals are wide, with small apical convergence. This trait is particularly pronounced in the Sima de los Huesos sample which may reflect a particularity of this population. Our study demonstrates the potential of hominin roots and root canals as untapped sources of taxonomic information when the tooth crown is fragmented. Future studies, including more fossil specimens and species will shed light in the polarity of the morphologies observed., (Copyright © 2012 Wiley Periodicals, Inc.)

The dispersal of hominins may have been favored by the opening of the landscape during the Early-Middle Pleistocene transition (EMP) in Western Europe. The structure of the small-vertebrate assemblages of the archaeo-paleontological karstic site of Gran Dolina in Atapuerca (Burgos, Spain) shows important environmental and climatic changes in the faunal succession, across the Matuyama-Brunhes boundary at 780 ka. These changes are interpreted to indicate impoverishment of the forests, along with an increase in dry meadows, and open lands in general that entailed a tendency towards the loss of diversity in small-vertebrate communities above the EMP. We evaluate variation in diversity of the faunal succession of Gran Dolina using Shannon's Second Theorem as an index of ecosystem structure. The long cultural-stratigraphic sequence of Gran Dolina during the EMP is somewhat similar in its completeness and continuity to that in the locality of Gesher Benot Ya'aqov in the Upper Jordan Valley. We also evaluate related data including faunal and floral (pollen) succession. Both localities present cold, dry and humid, warm fluctuations at the transition between the Early and the Middle Pleistocene. Comparisons between these sites present opportunities to understand large-scale climatic changes., (Copyright © 2010 Elsevier Ltd. All rights reserved.)

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.

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.

We present a mandible recovered in 2003 from the Aurora Stratum of the TD6 level of the Gran Dolina site (Sierra de Atapuerca, northern Spain). The specimen, catalogued as ATD6-96, adds to the hominin sample recovered from this site in 1994-1996, and assigned to Homo antecessor. ATD6-96 is the left half of a gracile mandible belonging to a probably female adult individual with premolars and molars in place. This mandible shows a primitive structural pattern shared with all African and Asian Homo species. However, it is small and exhibits a remarkable gracility, a trait shared only with the Early and Middle Pleistocene Chinese hominins. Furthermore, none of the mandibular features considered apomorphic in the European Middle and Early Upper Pleistocene hominin lineage are present in ATD6-96. This evidence reinforces the taxonomic identity of H. antecessor and is consistent with the hypothesis of a close relationship between this species and Homo sapiens.

Human hearing differs from that of chimpanzees and most other anthropoids in maintaining a relatively high sensitivity from 2 kHz up to 4 kHz, a region that contains relevant acoustic information in spoken language. Knowledge of the auditory capacities in human fossil ancestors could greatly enhance the understanding of when this human pattern emerged during the course of our evolutionary history. Here we use a comprehensive physical model to analyze the influence of skeletal structures on the acoustic filtering of the outer and middle ears in five fossil human specimens from the Middle Pleistocene site of the Sima de los Huesos in the Sierra de Atapuerca of Spain. Our results show that the skeletal anatomy in these hominids is compatible with a human-like pattern of sound power transmission through the outer and middle ear at frequencies up to 5 kHz, suggesting that they already had auditory capacities similar to those of living humans in this frequency range.

The Middle Pleistocene Atapuerca-Sima de los Huesos (SH) site in Spain has yielded the largest sample of fossil hominids so far found from a single site and belonging to the same biological population. The SH dental sample includes a total of 452 permanent and deciduous teeth, representing a minimum of 27 individuals. We present a study of the dental size variation in these hominids, based on the analysis of the mandibular permanent dentition: lateral incisors, n=29; canines, n=27; third premolars, n=30; fourth premolars, n=34; first molars, n=38; second molars, n=38. We have obtained the buccolingual diameter and the crown area (measured on occlusal photographs) of these teeth, and used the bootstrap method to assess the amount of variation in the SH sample compared with the variation of a modern human sample from the Museu Antropologico of the Universidade of Coimbra (Portugal). The SH hominids have, in general terms, a dental size variation higher than that of the modern human sample. The analysis is especially conclusive for the canines. Furthermore, we have estimated the degree of sexual dimorphism of the SH sample by obtaining male and female dental subsamples by means of sexing the large sample of SH mandibular specimens. We obtained the index of sexual dimorphism (ISD=male mean/female mean) and the values were compared with those obtained from the sexed modern human sample from Coimbra, and with data found in the literature concerning several recent human populations. In all tooth classes the ISD of the SH hominids was higher than that of modern humans, but the differences were generally modest, except for the canines, thus suggesting that canine size sexual dimorphism in Homo heidelbergensis was probably greater than that of modern humans. Since the approach of sexing fossil specimens has some obvious limitations, these results should be assessed with caution. Additional data from SH and other European Middle Pleistocene sites would be necessary to test this hypothesis., (Copyright 2001 Academic Press.)

. We describe the pattern of dental development of Hominid XVIII from the Middle Pleistocene Sima de los Huesos (SH) site of the Sierra de Atapuerca (Burgos, Spain). As expected, this pattern is similar to that of modern humans. A delay of development of the lower and upper canines was observed. In contrast, the relative advanced development of the lower second molars and, especially, the upper and lower third molars is noteworthy. This latter feature seems to be common in Pleistocene hominids, and suggests that the pattern of dental development evolved in the genus Homo during the Pleistocene. In European Middle Pleistocene hominids, this pattern probably was facilitated by the extra space available in the mandible and maxilla for developing teeth., (Copyright 2001 Wiley-Liss, Inc.)

Metric and shape features of the Lower Pleistocene mandibular specimen ATD605 from the level 6 of Gran Dolina site (Atapuerca, Spain) are compared with a large sample of fossil hominid mandibles. The analysis shows that ATD6-5 displays a generalized morphology largely shared with both African and European Lower and Middle Pleistocene samples. However, distinctive African traits, such as corpus robustness and strong alveolar prominence, are absent in the Gran Dolina specimen. At the same time, none of the apomorphic features that characterize Middle and early Upper Pleistocene European hominids can be recognized in ATD6-5. Finally, the Gran Dolina specimen displays a remarkable position of the mylohyoid groove, only comparable to that found in immature specimens of Homo ergaster, and very rarely in adult H. sapiens. The morphology of ATD6-5 supports the hypothesis of an African origin for the first Europeans with subsequent phylogenetic continuity with Middle Pleistocene populations in Europe. These findings are consistent with H. antecessor being the last common ancestor of Neandertals and H. sapiens., (Copyright 1999 Academic Press.)

Gran Dolina is part of an archaeological and paleontological complex located in the Sierra de Atapuerca karstic system (Burgos, Spain). The Trinchera del Ferrocarril sites were discovered as a consequence of the construction of a railway for the transport of minerals at the end of the nineteenth century. The systematic excavation of the upper Gran Dolina levels was initiated in 1981. In 1993, a 6 m(2)biostratigraphic survey pit was started, reaching level TD6 in 1994. This level was excavated during four consecutive years, yielding human fossils, identified as Homo antecessor, in association with lithic and faunal remains, dating to more than 0.78 m.y.a., (Copyright 1999 Academic Press.)

The study of the faunal and lithic assemblage (including almost a hundred human fossil remains) recovered from the Aurora stratum-TD6 level of the Lower Pleistocene cave site of Gran Dolina (Sierra de Atapuerca, Spain) has allowed us to answer some important questions concerning the debate about the earliest evidence for human occupation of Europe. However, it has also started new discussions about some geographical, ecological, and economic aspects of this earliest occupation. The nature (definitive or ephemeral) of the first occupation, as well as the model for the arrival of the Acheulean (Mode 2) in Europe are also issues for discussion., (Copyright 1999 Academic Press.)

The Middle Pleistocene site of Sima de los Huesos in Sierra de Atapuerca, Spain, has yielded around 2,500 fossils from at least 33 different hominid individuals. These have been dated at more than 200,000 years ago and have been classified as ancestors of Neanderthals. An almost complete human male pelvis (labelled Pelvis 1) has been found, which we associate with two fragmentary femora. Pelvis 1 is robust and very broad with a very long superior pubic ramus, marked iliac flare, and a long femoral neck. This pattern is probably the primitive condition from which modern humans departed. A modern human newborn would pass through the birth canal of Pelvis 1 and this would be even larger in a female individual. We estimate the body mass of this individual at 95 kg or more. Using the cranial capacities of three specimens from Sima de los Huesos, the encephalization quotients are substantially smaller than in Neanderthals and modern humans.

The recent discovery of unexpectedly ancient human remains has fuelled interest about the first dispersion of Homo outside Africa. The Dmanisi mandible is perhaps one of the most interesting findings, as it supposedly represents one of the oldest hominids outside of Africa. Recently, different interpretations have been published about this specimen. Our comparison of the Dmanisi mandible with a large sample of mandibles and teeth has led us to a new interpretation. In our view, the Dmanisi mandible exhibits a unique combination of traits. Some of its features, taken in isolation, may be attributed to morphological extremes within the genus Homo. The architecture of the mandible as well as the morphology and dimensions of incisors, canines, and P3s are clearly primitive. However, dental traits such as the reduction of the talonid in the P4s and a distally decreasing molar series seems to be derived. Some combinations of these traits are found in specimens of Homo ergaster and differ from those generally present in later hominids. Thus, we propose that the Dmanisi mandible might be taxonomically classified as Homo sp. indet. (aff. ergaster). Furthermore, some aspects of the dentition in Dmanisi display close similarities to Asian Homo erectus. If the 1.8-1.6 Myr dating for the Dmanisi mandible is correct, the differentiation of the Asian branch of the genus Homo could be regarded as a very ancient event.

It has been suggested that European Middle Pleistocene humans, Neandertals, and prehistoric modern humans had a greater sexual dimorphism than modern humans. Analysis of body size variation and cranial capacity variation in the large sample from the Sima de los Huesos site in Spain showed instead that the sexual dimorphism is comparable in Middle Pleistocene and modern populations.

We report here on the palaeodemographic analysis of the hominid sample recovered to date from the Sima de los Huesos (SH) Middle Pleistocene cave site in the Sierra de Atapuerca (Burgos, Spain). The analysis of the mandibular, maxillary, and dental remains has made it possible to estimate that a minimum of 32 individuals, who probably belonged to the same biological population, are represented in the current SH human hypodigm. The remains of nine-individuals are assigned to males, and nine to females, suggesting that a 1:1 sex ratio characterizes this hominid sample. The survivorship curve shows a low representation of infants and children, a high mortality among the adolescents and prime-age adults, and a low older adult mortality. Longevity was probably no greater than 40 years. This mortality pattern (adolescents and adults); which in some aspects resembles that observed in Neandertals, is quite different from those reported for recent foraging human groups. The adult age-at-death distribution of the SH hominid sample appears to be neither the consequence of underaging the older adults, nor of differential preservation or of the recognition of skeletal remains. Thus if we accept that they had a life history pattern similar to that of modern humans there would appear to be a clear contradiction between the demographic distribution and the demographic viability of the population represented by the SH hominid fossils. The possible representational bias of the SH hominid sample, as well as some aspects of the reproductive biology of the Pleistocene populations are also discussed.

Human fossil remains recovered from the TD6 level (Aurora stratum) of the lower Pleistocene cave site of Gran Dolina, Sierra de Atapuerca, Spain, exhibit a unique combination of cranial, mandibular, and dental traits and are suggested as a new species of Homo-H. antecessor sp. nov. The fully modern midfacial morphology of the fossils antedates other evidence of this feature by about 650, 000 years. The midfacial and subnasal morphology of modern humans may be a retention of a juvenile pattern that was not yet present in H. ergaster. Homo antecessor may represent the last common ancestor for Neandertals and modern humans.

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.

We present the palaeopathological analysis of a right fourth metatarsal (ATD6-124) recovered from the Atapuerca-Gran Dolina site (Spain). This fossil, ca. 1 Ma, belongs to Homo antecessor, the earliest known European hominin species. The metatarsal exhibits a proliferative lesion on its medial periosteal surface. Periosteal reaction can be the bone response to a wide number of injurious processes. We describe a lesion on the basis of macroscopic and microscopic analyses, including microtomography and scanning electron microscopy. Externally, the osteoblastic lesion presents a highly porotic and disorganised morphology. Internally, we observe a series of micro-fractures on the compact bone that do not affect the medullary canal. We provide a differential diagnosis and suggest that the ATD6-124 lesion could correspond to a pedal stress fracture, also known as fatigue or march fracture. Stress fractures have been related to a load increase and muscular fatigue. This type of fracture has been widely reported in the foot of soldiers and athletes, which are usually engaged in strenuous, excessive or prolonged locomotive activities. Despite its high frequency in these groups, stress fractures have not been reported as such in fossil collections, with the exception of a metatarsal belonging to the Sima de los Huesos site (Atapuerca). Copyright © 2013 John Wiley & Sons, Ltd. [ABSTRACT FROM AUTHOR]

This study presents quantitative data on the rates of anterior tooth wear in a Pleistocene human population. The data were obtained for the hominin sample of the Sima de los Huesos site in Atapuerca, Spain. The fossil record belongs to a minimum of 28 individuals of the same biological population, assigned to the species Homo heidelbergensis. We have estimated the original and the preserved crown height of the mandibular incisors (11 and 12) of 11 individuals, whose age at death can be ascertained from the mineralization stage and tooth eruption. Results provide a range of 0.2760.348 and 0.288-0.360 mm per year for the mean wear rate of the mandibular I1 and 12, respectively, in individuals &ap;16-18 years old. These data suggest that incisors' crowns would be totally worn out toward the fifth decade of life. Thus, we expect the life expectancy of this population to be seriously limited. These data, which could be contrasted with results obtained on hominins at other sites, could be of interest for estimating the death age of adult individuals. [ABSTRACT FROM AUTHOR]