Phylogenetic Implications. It is apparent from the suite of Neandertal lineage features present in the Aubesier Middle Pleistocene human remains that they provide further evidence for the gradual emergence during the second half of the Middle Pleistocene and the early Late Pleistocene of Europe of the derived (at least in frequency) Neandertal morphological pattern. This pattern is evident in the Aubesier 4 crown morphology, the Aubesier 10 root configuration, and several features (mental foramen and anterior marginal tubercle position, retromolar space, superior medial pterygoid tubercle, and mandibular notch crest position) of the Aubesier 11 mandibular lateral corpus and ramus.
At the same time, taken in the context of the variable frequencies of all of these traits within Neandertal lineage samples and other hominids groups, plus the clear presence of this pattern by the late Middle Pleistocene and initial Late Pleistocene of western Asia (21, 25, 39), it is unlikely that the Neandertal morphological complex evolved in isolation in Europe. It is more probable that these features increased in frequency through a combination of genetic drift and isolation by distance across the northwestern Old World, as one aspect of the general regional differentiation of hominid craniofacial morphology as the genus Homo became increasingly geographically dispersed during the Middle Pleistocene (40).
Paleobiological Implications: The Teeth. The Aubesier 4 I2 occlusal beveling reinforces a pattern of considerable use of the anterior dentition for nonmasticatory purposes. This degree of beveling is well documented for more recent Neandertal lineage specimens (21, 33, 41), and labial wear striae have been documented for anterior teeth from AT-SH (42), Krapina (43), and several Neandertal specimens (12, 21, 44, 45). Sufficiently worn non-European archaic Homo anterior maxillary teeth are rare, but at least the African Broken Hill 1 I1s exhibit a degree and pattern of anterior dental wear suggesting nondietary dental use similar to that seen in the European Neandertal lineage.
The mesial interproximal wear groove on Aubesier 10 appears most likely to have been produced by some form of a tool to remove debris from between the teeth, because the large adjacent interproximal facet (7.1 mm wide) reflects close approximation of the adjacent mesial tooth and would have made stripping material between the teeth difficult. The groove serves to reinforce a pattern of dental care, despite the absence of documented dental caries in Eurasia before the last glacial.
Paleobiological Implications: The Mandible. In the Aubesier 11 mandible, the combination of apical abscesses, M3 socket enlargement and lingual fenestration, rounding of exposed trabeculae and generalized alveolar bone loss implies pervasive and active periodontal-endodontal lesions throughout the dental arcades with all of the teeth being lost antemortem or mechanically unstable at the time of death. Although Aubesier 11 was still chewing, as indicated by the C1 and P3 root wear, the individual's ability to masticate tough or hard food items must have been severely compromised. The probable associated pain would have further impeded the process.
Significant antemortem tooth loss is documented for last glacial Neandertals [e.g., La Chapelle-aux-Saints 1, Guattari 1, and Shanidar 5 (21, 46, 47)], and alveolar bone loss and apical abscesses are commonly noted among the Neandertals (21, 41, 47-49). However, no hominid specimen dating before the last glacial with significant antemortem masticatory impairment has been previously documented. Broken Hill 1 experienced dental caries and tooth loss (50), but retained sufficient occlusal surface for mastication. The AT-SH 700/888 skull (AT-SH cranium 5 and mandible) sustained maxillary and mandibular dental apical abscesses and associated maxillary lesions after dental trauma, but without apparent dental loss (4). Arago 21 lost a P4 antemortem (51), and Ehringsdorf 6 exhibits anterior alveolar lesions and resorption (52).
Aubesier 11 is therefore unique among Middle Pleistocene (or earlier) hominids in its degree of antemortem masticatory impairment. This reduction in the ability to chew food implies significant behavioral adjustment to maintain Aubesier 11 alive for some period. This behavior could have involved selective allocation of softer food items, extensive manual preparation of food, and/or extensive cooking of food. Evidence for fire in the Middle Pleistocene of Europe before 150 ka B.P. is scattered and scarce. It consists mostly of evidence of burned bones and charcoal within archeological levels, with only scarce evidence for constructed hearths (53-57). Significant evidence for the concentrated use of fire comes in particular from layer H-1 of the Bau de l'Aubesier, the "black combustion horizon" overlying the hominid layers yielding Aubesier 10 and 11. It contains an abundance of charcoal, burnt bones, heated lithics, and other archeological remains, but there is no evidence of constructed hearths. It is therefore apparent that the ability to make and maintain fire was present around the time of Aubesier 11, but the degree to which it was closely controlled and used for culinary activities, as opposed to thermoregulation and illumination, remains uncertain at present.
Paleontological cases of serious lesions are rare among Middle Pleistocene (and earlier) human remains. In the Middle Pleistocene, the Salé 1 cranium exhibits evidence of a congenital torticollis (58), the Berg Aukas 1 femur had abnormal development of the hip region (59), and the Singa 1 cranium exhibits biparietal diploic expansion and unilateral labyrinthine ossification (60). Among the Neandertals, Shanidar 1 persisted through multiple traumatic injuries and hyperostotic disease (21, 61), whereas other Neandertals experienced varying degrees of traumatic and degenerative lesions (47, 62), in addition to alveolar ones. Aubesier 11 is therefore not unique but remains unusual for a Middle Pleistocene human in its degree of impairment of a vital function.
Combined with other evidence of the survival of pathological individuals during the Middle Pleistocene, the Aubesier 11 lesions provide support for an important social component in the persistence of individuals during this time period. At the same time, the nondietary use of the anterior dentition, combined with pronounced appendicular hypertrophy (63-66), indicates that these Middle Pleistocene humans were using their bodies extensively to accomplish routine activities, albeit technologically aided in so doing.