New interpretation of the Gran Dolina-TD6 bearing Homo antecessor deposits through sedimentological analysis.

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.

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

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.

The Early-Middle Pleistocene environmental and climatic change and the human expansion in Western Europe: A case study with small vertebrates (Gran Dolina, Atapuerca, Spain).

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.

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.

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 geometric morphometric analysis of hominin upper first molar shape.

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.

Hominin lower second premolar morphology: evolutionary inferences through geometric morphometric analysis.

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.

Matuyama-age lithic tools from the Sima del Elefante site, Atapuerca (northern Spain).

Paleomagnetic results obtained from the sedimentary fill at the Sima del Elefante site in Atapuerca, Spain, reveal a geomagnetic reversal, interpreted as the Matuyama-Brunhes boundary (0.78 Ma). The uppermost lithostratigraphic units (E17 through E19), which contain Mode II and III archaeological assemblages, display normal polarity magnetization, whereas the six lowermost units (E9 through E16) yield negative latitudinal virtual geomagnetic pole positions. Units E9 through E13, all of which display reverse magnetic polarity, contain Mode I (Oldowan) lithic tools, testifying to the presence of humans in the early Pleistocene (0.78-1.77 Ma).

An Early Pleistocene hominin mandible from Atapuerca-TD6, Spain.

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.

Enamel hypoplasias and physiological stress in the Sima de los Huesos Middle Pleistocene hominins.

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.

Auditory capacities in Middle Pleistocene humans from the Sierra de Atapuerca in Spain.

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.

Rates of anterior tooth wear in Middle Pleistocene hominins from Sima de los Huesos (Sierra de Atapuerca, Spain).

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.

Contribution of snacks and meals in the diet of French adults: a diet-diary study.

To investigate the relative contributions of meals and snacks in the daily intake of free-living humans, 54 French adults maintained food intake diaries for four 7-day periods. They recorded all food and fluid intakes mentioning whether, in their opinion, each intake event was a snack or a meal. The weekly food diaries also contained information on the circumstances of each event such as time and place, number of persons present, and affective states (hunger, satiety, etc.) before and after intake. On average, 2.7 meals and 1.3 snacks were consumed each day. Very few days included no snacking. Total daily energy and nutrient intake were not different between days with and days without snacks. Snacks differed from meals in several dimensions. Meals were about twice as large as snacks in energy and weight. Nutrient intake, in absolute values, was higher in meals. In proportions, however, snacks contained more CHO and less fat and proteins. Most foods were consumed in larger amounts in the context of meals but a few (sweets, cereal bars, biscuits, and sodas) were mostly consumed as snacks. Hunger was more intense before but less intense after meals than snacks. The satiety ratio was higher for snacks than meals. Time of day affected many intake parameters. For example, afternoon snacks exhibited a high satiety ratio for a modest intake. The present study describes the status of several potential determining factors at the time of snacks in humans, demonstrating a specific role for snacks, as opposed to meals, in the daily eating pattern of healthy adults.

Human remains from Valdegoba Cave (Huérmeces, Burgos, Spain).

Systematic excavations, begun in 1987, at the Valdegoba cave site in northern Spain have yielded the remains of five individuals associated with a Middle Paleolithic stone tool technology and Pleistocene fauna. A fragmentary mandible of an adolescent (VB1), preserving nearly a full set of teeth, exhibits a symphyseal tubercle and slight incurvatio mandibulae anterior on the external symphysis. Both the superior and inferior transverse tori are present on the internal aspect. A second individual (VB2) is represented by a set of ten deciduous teeth consistent with an age at death of 6-9 months. A proximal manual phalanx (VB3) displays a relatively broad head, a characteristic which is found in both Neandertals, as well as European Middle Pleistocene hominids. VB4 is a fourth metatarsal that lacks the distal epiphysis, indicating it comes from an adolescent individual, and has a relatively high robusticity index. Finally, VB5 is a fifth metatarsal of an adult. The VB1 mandible shows a combination of archaic characteristics as well as more specific Neandertal morphological traits. The VB2 deciduous teeth are very small, and both the metrics and morphology seem more consistent with a modern human classification. The postcranial elements are undiagnostic, U-Th dating has provided an age of >350 ka for the base of the sequence and a date of <73.2+/-5 ka for level 7, near the top. Faunal analysis and radiometric dates from other nearby Mousterian sites suggests that the Valdegoba site is correlative with oxygen isotope stages 3-6 on the Iberian peninsula, and an Upper Pleistocene age for the Valdegoba hominids seems most reasonable.

Dental size variation in the Atapuerca-SH Middle Pleistocene hominids.

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.

Heritability of diurnal changes in food intake in free-living humans.

OBJECTIVES: The time of day of meal ingestion, the number of people present at the meal, the subjective state of hunger, and the estimated before-meal contents in the stomach have been established as influences on the amount eaten in a meal, and this influence has been shown to be heritable. Because these factors intercorrelate, the possibility that the calculated heritabilities for some of these variables could result indirectly from their convariation with one of the other heritable variables was assessed. METHODS: The independence of the heritability of the influence of these four factors was investigated with 110 identical and 102 fraternal same-sex and 53 fraternal mixed-sex adult twin pairs who were paid to maintain 7-d food intake diaries. From the diary reports, the meal sizes were calculated and subjected to multiple regression analysis using the estimated before-meal stomach contents, the reported number of other people present, the subjective hunger ratings, and the time of day of the meal as predictors. Linear structural modeling was applied to the beta coefficients from the multiple regression to investigate whether the heritability of the influences of these four variables was independent. RESULTS: Significant genetic effects were found for the beta coefficients for all four variables, indicating that the heritability of their relationship with intake is to some extent heritable. CONCLUSIONS: These results suggest that the influences of multiple factors on intake are influenced by the genes and become part of the total package of genetically determined physiologic, sociocultural, and psychological processes that regulate energy balance.

Palatability and intake relationships in free-living humans: the influence of heredity.

The heritability of the before and after meal self-ratings of palatability and their relationship to food intake was investigated with 86 identical and 78 fraternal same-sex and 51 fraternal mixed gender adult twin pairs who were paid to maintain 7-day food intake diaries. From the diary reports, the total and meal intakes of food energy and the amounts of the macronutrients ingested were estimated. Participants rated the meals for palatability on a 7-point (Bad - Good) scale both before and after eating. Linear structural modeling was applied to investigate the nature and degree of genetic and environmental influences and revealed significant genetic influences on subjective palatability both before and after the meals. In addition, the relationship between palatability and intake was influenced by the genes. Significant dominance genetic effects were found for the change in meal size between low-moderate and high palatability meals, indicating that palatability and it's relationship with intake are to some extent heritable. This suggests that the level of subjective experience and the individuals responses to them are influenced by the genes and become part of the total package of genetically determined physiological, socio/cultural, and psychological processes that regulate energy balance.

Pattern of dental development in Hominid XVIII from the Middle Pleistocene Atapuerca-Sima de los Huesos site (Spain).

. 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.

Eating behavior: lessons from the real world of humans.

Food intake by normal humans has been investigated both in the laboratory and under free-living conditions in the natural environment. For measurement of real-world intake, the diet-diary technique is imperfect and tends to underestimate actual intakes but it appears to be sensitive, can detect subtle influences on eating behavior, and produces reliable and valid measures. Research studies in the real world show the multivariate richness of the natural environment, which allows investigation of the complexities of intake regulation, and even causation can be investigated. Real-world research can overcome some of the weaknesses of laboratory studies, where constraints on eating are often removed or missing, facilitatory influences on eating are often controlled or eliminated, the importance of variables can be overestimated, and important influences can be missed because of the short durations of the studies. Real-world studies have shown a wide array of physiologic, psychological, and social variables that can have potent and immediate effects on intake. Compensatory mechanisms, including some that operate with a 2- to 3-d delay, adjust for prior excesses. Heredity affects all aspect of food-intake regulation, from the determination of body size to the subtleties of the individual preferences and social proclivities and the extent to which environmental factors affect the individual. Hence, real-world research teaches valuable lessons, and much more is needed to complement laboratory studies.