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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.

Recent studies have revealed interesting differences in upper first molar morphology across the hominin fossil record, particularly significant between H. sapiens and H. neanderthalensis. Usually these analyses have been performed by means of classic morphometric methods, including the measurement of relative cusp areas or the angles defined between cusps. Although these studies have provided valuable information for the morphological characterization of some hominin species, we believe that the analysis of this particular tooth could be more conclusive for taxonomic assignment. In this study, we have applied geometric morphometric methods to explore the morphological variability of the upper first molar (M(1)) across the human fossil record. Our emphasis focuses on the study of the phenetic relationships among the European middle Pleistocene populations (designated as H. heidelbergensis) with H. neanderthalensis and H. sapiens, but the inclusion of Australopithecus and early Homo specimens has helped us to assess the polarity of the observed traits. H. neanderthalensis presents a unique morphology characterized by a relatively distal displacement of the lingual cusps and protrusion in the external outline of a large and bulging hypocone. This morphology can be found in a less pronounced degree in the European early and middle Pleistocene populations, and reaches its maximum expression with the H. neanderthalensis lineage. In contrast, modern humans retain the primitive morphology with a square occlusal polygon associated with a round external outline.

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.

Mandibular premolars are increasingly used in taxon-specific diagnostic analyses of hominins. Among the principal difficulties in these evaluations is the absence of discrete, discernible, and comparable anatomical structures for rigorous quantitative assessment. Previous research has addressed either internal crown surface features (such as cusps and fossae) or the morphology of the crown outline. In the present paper, we integrate both types of information in the examination of morphological variation of lower P4s (n = 96) among various fossil hominin species with an emphasis on genus Homo. We use a set of 34 2D landmarks combining coordinate data from four classical dental landmarks on the occlusal surface and 30 sliding semilandmarks of the crown outline. Our results indicate that external shape variation is closely related to the configuration of the occlusal morphological features and influenced by dental size. The external and internal shapes of P4 are polymorphic but still useful in depicting a primitive-derived gradient. The primitive pattern seems to have been an asymmetrical contour with a mesially displaced metaconid, development of a bulging talonid, and a broad occlusal polygon. The trend toward dental reduction during the Pleistocene produced different morphological variants with a reduced occlusal polygon and decreased lingual occlusal surface in later Homo species. Homo heidelbergensis/neanderthalensis have fixed plesiomorphic traits in high percentages, whereas in modern humans a symmetrical outline with a centered metaconid and talonid reduction evolved.

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.

This study presents an analysis of linear enamel hypoplasias (LEH) and plane-form defects (PFD) in the hominine dental sample from the Sima de los Huesos (SH) Middle Pleistocene site in Atapuerca (Spain). The SH sample comprises 475 teeth, 467 permanent and 8 deciduous, belonging to a minimum of 28 individuals. The method for recording PFD and LEH is discussed, as well as the definition of LEH. The prevalence of LEH and PFD in SH permanent dentition (unilateral total count) is 4.6% (13/280). Only one deciduous tooth (lower dc) showed an enamel disruption. Prevalence by individual ranges from 18.7-30%. The most likely explanation for the relatively low LEH and PFD prevalence in the SH sample suggests that the SH population exhibited a low level of developmental stress. The age at occurrence of LEH and PFD was determined by counting the number of perikymata between each lesion and the cervix of the tooth. Assuming a periodicity of nine days for the incremental lines, the majority of LEH in the SH sample occurred during the third year of life and may be related to the metabolic stress associated with weaning.

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 (I1 and I2) of 11 individuals, whose age at death can be ascertained from the mineralization stage and tooth eruption. Results provide a range of 0.276-0.348 and 0.288-0.360 mm per year for the mean wear rate of the mandibular I1 and I2, respectively, in individuals approximately 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.

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.)

Parameters of a Middle Pleistocene human population such as the expected length of the female reproductive period (E(Y)), the expected interbirth interval (E(X)), the survival rate (tau) for females after the expected reproductive period, the rate (phi(2)) of women who, given that they reach first birth, do not survive to the end of the expected reproductive period, and the female infant plus juvenile mortality rate (phi(1)) have been assessed from a probabilistic standpoint provided that such a population were stationary. The hominid sample studied, the Sima de los Huesos (SH) cave site, Sierra de Atapuerca (Spain), is the most exhaustive human fossil sample currently available. Results suggest that the Atapuerca (SH) sample can derive from a stationary population. Further, in the case that the expected reproductive period ends between 37 and 40 yr of age, then 24 less, similarE(Y) less, similar27 yr, E(X)=3 yr, 0.224

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.)

Hominid remains found in 1994 from the stratified Gran Dolina karst-filling at the Atapuerca site in NE Spain were dated to somewhat greater than 780 ka based on palaeomagnetic measurements, making these the oldest known hominids in Europe (sensu stricto). We report new ESR and U-series results on teeth from four levels of the Gran Dolina deposit which confirm the palaeomagnetic evidence, and indicate that TD6 (from which the human remains have been recovered) dates to the end of the Early Pleistocene. The results for the other levels are consistent with estimates based mainly on microfaunal evidence, and suggest that TD8, TD10 and TD11 date to the Middle Pleistocene., (Copyright 1999 Academic Press.)

A descriptive and comparative study of the human dental remains recovered from level six (Aurora stratum) of the Gran Dolina (TD) site, Sierra de Atapuerca, northern Spain, is presented. The TD6 dental sample consists of two deciduous and 28 permanent teeth, belonging to a minimum of six individuals. Paleomagnetic analyses of the TD sequence suggest an age for the Aurora stratum of greater than 780 ka. The associated fossil macro- and micrommals are consistent with a date for the Atapuerca-TD6 hominids at the end of the Early Pleistocene. As a whole, the size of the TD6 teeth is large and comparable to that of the African Lower Pleistocene hominids. Most morphological dental traits are plesiomorphic for the genus Homo, whereas the shape analysis has revealed some size-related trends that differ from those seen in early Homo. Thus, the relative size increase of the maxillary and mandibular anterior teeth and buccolingual diameter of the first molars, the reduction of the third molars and lower canines, and the P(3)>P(4)size sequence are all characteristic of the population represented by the TD6 hominids. The morphological evidence suggests that the TD6 hominids are closer to African Lower and early Middle Pleistocene hominids than they are to European Middle Pleistocene hominids. However, some dental traits of the TD6 hominids, such as the expansion of the anterior teeth, the P(3)>P(4)size sequence, and the morphology of the lower canine also suggest an evolutionary continuity between the population represented by these hominids and the European Middle Pleistocene population. Furthermore, dental evidence indicates that the TD6 hominids are clearly distinct from Asian H. erectus. Dental evidence also suggests that the TD6 hominids could represent an evolutionary connection between H. ergaster and H. heidelbergensis., (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 study of life history evolution in hominids is crucial for the discernment of when and why humans have acquired our unique maturational pattern. Because the development of dentition is critically integrated into the life cycle in mammals, the determination of the time and pattern of dental development represents an appropriate method to infer changes in life history variables that occurred during hominid evolution. Here we present evidence derived from Lower Pleistocene human fossil remains recovered from the TD6 level (Aurora stratum) of the Gran Dolina site in the Sierra de Atapuerca, northern Spain. These hominids present a pattern of development similar to that of Homo sapiens, although some aspects (e.g., delayed M3 calcification) are not as derived as that of European populations and people of European origin. This evidence, taken together with the present knowledge of cranial capacity of these and other late Early Pleistocene hominids, supports the view that as early as 0.8 Ma at least one Homo species shared with modern humans a prolonged pattern of maturation.

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.

In order to reassess previous hypotheses concerning dental size reduction of the posterior teeth during Pleistocene human evolution, current fossil dental evidence is examined. This evidence includes the large sample of hominid teeth found in recent excavations (1984-1993) in the Sima de los Huesos Middle Pleistocene cave site of the Sierra de Atapuerca (Burgos, Spain). The lower fourth premolars and molars of the Atapuerca hominids, probably older than 300 Kyr, have dimensions similar to those of modern humans. Further, these hominids share the derived state of other features of the posterior teeth with modern humans, such as a similar relative molar size and frequent absence of the hypoconulid, thus suggesting a possible case of parallelism. We believe that dietary changes allowed size reduction of the posterior teeth during the Middle Pleistocene, and the present evidence suggests that the selective pressures that operated on the size variability of these teeth were less restrictive than what is assumed by previous models of dental reduction. Thus, the causal relationship between tooth size decrease and changes in food-preparation techniques during the Pleistocene should be reconsidered. Moreover, the present evidence indicates that the differential reduction of the molars cannot be explained in terms of restriction of available growth space. The molar crown area measurements of a modern human sample were also investigated. The results of this study, as well as previous similar analyses, suggest that a decrease of the rate of cell proliferation, which affected the later-forming crown regions to a greater extent, may be the biological process responsible for the general and differential dental size reduction that occurred during human evolution.

The prevalence and chronology of enamel hypoplasias were studied in a hominid dental sample from the Sima de los Huesos (SH) Middle Pleistocene site at the Sierra de Atapuerca (Burgos, northern Spain). A total of 89 permanent maxillary teeth, 143 permanent mandibular teeth, and one deciduous lower canine, belonging to a minimum of 29 individuals, were examined. Excluding the antimeres (16 maxillary and 37 mandibular cases) from the sample, the prevalence of hypoplasias in the permanent dentition is 12.8% (23/179), whereas the deciduous tooth also showed an enamel defect. No statistically significant differences were found between both arcades and between the anterior and postcanine teeth for the prevalence of hypoplasias. In both the maxilla and the mandible the highest frequency of enamel hypoplasias was recorded in the canines. Only one tooth (a permanent upper canine) showed two different enamel defects, and most of the hypoplasias were expressed as faint linear horizontal defects. Taking into account the limitations that the incompleteness of virtually all permanent dentitions imposes, we have estimated that the frequency by individual in the SH hominid sample was not greater than 40%. Most of the hypoplasias occurred between birth and 7 years (N = 18, X = 3.5, SD = 1.3). Both the prevalence and severity of the hypoplasias of the SH hominid sample are significantly less than those of a large Neandertal sample. Furthermore, prehistoric hunter-gatherers and historic agricultural and industrial populations exhibit a prevalence of hypoplasias generally higher than that of the SH hominids. Implications for the survival strategies and life quality of the SH hominids are also discussed.