Sexual dimorphism in the Atapuerca-SH hominids: the evidence from the mandibles.
The pattern of sexual dimorphism in 15 mandibles from the Atapuerca-SH Middle Pleistocene site, attributed to Homo heidelbergensis, is explored. Two modern human samples of known sex are used as a baseline for establishing sexing criteria. The mandible was divided for analysis into seven study regions and differential expression of sexual dimorphism in these regions is analysed. A total of 40 continuous and 32 discrete variables were scored on the mandibles. The means method given in Regh & Leigh (Am. J. phys. Anthrop.110, 95-104, 1999) was followed for evaluating the potential of correct sex attribution for each variable. On average, the mandibles from the Atapuerca-SH site present a degree of sexual dimorphism about eight points higher than in H. sapiens samples. However, mandibular anatomy of the European Middle Pleistocene hominid records sexual dimorphism differentially. Different areas of the Atapuerca-SH mandibles exhibit quite distinct degrees of sexual dimorphism. For instance, variables of the alveolar arcade present very low or practically no sexual dimorphism. Variables related to overall size of the mandible and symphysis region present a medium degree of sex differences. Finally, ramus height, and gonion and coronoid process present a high degree of sexual dimorphism (indexes of sexual dimorphism are all above 130%). Whether this marked sexual dimorphism in specific anatomical systems affects sexual differences in body size is not completely clear and further studies are needed. Sexual differences detected in the mandible of modern humans have at least two components: differences related to musculo-skeletal development and differences related to a different growth trajectory in males and females (relative development of some of the basal border features). The Atapuerca-SH mandibles display little variation in the basal border, however. The limited variation of this mandibular region may indicate that the pattern of sexual variation in H. heidelbergensis is different enough to that of H. sapiens to caution against simple extrapolation of criteria from one pattern to the other.
Rosas A
，Bastir M
，Martínez-Maza C
，Bermúdez de Castro JM
... -
《journal of human evolution》
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.
Bermúdez de Castro JM
，Sarmiento S
，Cunha E
，Rosas A
，Bastir M
... -
《journal of human evolution》
The Neandertal nature of the Atapuerca Sima de los Huesos mandibles.
The recovery of additional mandibular fossils from the Atapuerca Sima de los Huesos (SH) site provides new insights into the evolutionary significance of this sample. In particular, morphological descriptions of the new adult specimens are provided, along with standardized metric data and phylogenetically relevant morphological features for the expanded adult sample. The new and more complete specimens extend the known range of variation in the Atapuerca (SH) mandibles in some metric and morphological details. In other aspects, the addition of new specimens has made it possible to confirm previous observations based on more limited evidence. Pairwise comparisons of individual metric variables revealed the only significant difference between the Atapuerca (SH) hominins and Neandertals was a more vertical symphysis in the latter. Similarly, principal components analysis of size-adjusted variables showed a strong similarity between the Atapuerca (SH) hominins and Neandertals. Morphologically, the Atapuerca (SH) mandibles show nearly the full complement of Neandertal-derived features. Nevertheless, the Neandertals differ from the Atapuerca (SH) mandibles in showing a high frequency of the H/O mandibular foramen, a truncated, thinned and inverted gonial margin, a high placement of the mylohyoid line at the level of the M3, a more vertical symphysis and somewhat more pronounced expression of the chin structures. Size-related morphological variation in the SH hominins includes larger retromolar spaces, more posterior placement of the lateral corpus structures, and stronger markings associated with the muscles of mastication in larger specimens. However, phylogenetically relevant features in the SH sample are fairly stable and do not vary with the overall size of the mandible. Direct comparison of the enlarged mandibular sample from Atapuerca (SH) with the Mauer mandible, the type specimen of H. heidelbergensis, reveals important differences from the SH hominins, and there is no morphological counterpart of Mauer within the SH sample, suggesting the SH fossils should not be assigned to this taxon. The Atapuerca (SH) mandibles show a greater number of derived Neandertal features, particularly those related to midfacial prognathism and in the configuration of the superior ramus, than other European middle Pleistocene specimens. This suggests that more than one evolutionary lineage co-existed in the middle Pleistocene, and, broadly speaking, it appears possible to separate the European middle Pleistocene mandibular remains into two distinct groupings. One group shows a suite of derived Neandertal features and includes specimens from the sites of Atapuerca (SH), Payre, l'Aubesier and Ehringsdorf. The other group includes specimens that generally lack derived Neandertal features and includes the mandibles from the sites of Mauer, Mala Balanica, Montmaurin and (probably) Visogliano. The two published Arago mandibles differ strongly from one another, with Arago 2 probably belonging to this former group, and Neandertal affinities being more difficult to identify in Arago 13. Outside of the SH sample, derived Neandertal features in the mandible only become more common during the second half of the middle Pleistocene. Acceptance of a cladogenetic pattern of evolution during the European middle Pleistocene has the potential to reconcile the predictions of the accretion model and the two phases model for the appearance of Neandertal morphology. The precise taxonomic classification of the SH hominins must contemplate features from the dentition, cranium, mandible and postcranial skeleton, all of which are preserved at the SH site. Nevertheless, the origin of the Neandertal clade may be tied to a speciation event reflected in the appearance of a suite of derived Neandertal features in the face, dentition and mandible, all of which are present in the Atapuerca (SH) hominins. This same suite of features also provides a useful anatomical basis to include other European middle Pleistocene mandibles and crania within the Neandertal clade.
Quam R
，Martínez I
，Rak Y
，Hylander B
，Pantoja A
，Lorenzo C
，Conde-Valverde M
，Keeling B
，Ortega Martínez MC
，Arsuaga JL
... -
《-》
Occurrence of neanderthal features in mandibles from the Atapuerca-SH site.
Analysis of variation and distribution of evolutionary novelties is meaningful in understanding evolutionary processes. The mandible, as a morphological complex, comprises a large number of derived Neanderthal features. The present study investigates whether the features usually considered as European lineage apomorphies evolved independently; the occurrence of these features is studied in the mandibles from the Sima de los Huesos (SH) site (Atapuerca, Spain). For comparative purposes, a large sample of Neanderthal mandibles as well as older fossil Homo specimens have been used for the study. Chi-square tests were employed to test for independence. The SH mandibles present a set of features that clearly show the basic architecture of the Neanderthal mandible. A highly significant association is detected in the variation of the position of the mental foramen, the lateral prominence, and the anterior marginal tubercle, as well as in the development of retromolar space. However, a much weaker association is detected in the features of the internal aspect of the mandible, with a few exceptions. Features of the external aspect of the mandible occur chronologically earlier than those observed in the internal aspect. The hypothesis that two distinct and consecutive morphological processes have driven the emergence of the European lineage throughout the Middle Pleistocene is proposed. A first transformation affects the mandible by means of backwards displacement of the structures located at the external aspect, as well as the position of the condyle. A second process would modify the features of the internal aspect of the mandible, in which the relief of the masseteric and pterygoid fossae are affected, in association with a spatial rearrangement of the corpus and ramus. Analyzed individually, some of the considered features may be questioned as Neanderthal apomorphies (Trinkaus,1993; Franciscus and Trinkaus, 995); however, the joint occurrence of many of them suggests that the complex is an evolutionary novelty.
Rosas A
《AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY》
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