Evolution of M1 crown size and cusp proportions in the genus Homo.

Previous research into tooth crown dimensions and cusp proportions has proved to be a useful way to identify taxonomic differences in Pliocene and Pleistocene fossil hominins. The present study has identified changes in both M(1) crown size and cusp proportions within the genus Homo, with M(1) overall crown size reduction apparently occurring in two main stages. The first stage (a reduction of ca. 17%) is associated with the emergence of Homo ergaster and Homo erectus sensu stricto. The second stage (a reduction of ca. 10%) occurs in Homo sapiens, but the reduced modern human M(1) tooth crown size was only attained in Upper Paleolithic times. The absolute sizes of the individual cusps are highly positively correlated with overall crown size and dental reduction produces a reduction in the absolute size of each of the cusps. Most of the individual cusps scale isometrically with crown size, but the paracone shows a negative allometric relationship, indicating that the reduction in paracone size is less than in the other M(1) cusps. Thus, the phylogenetically oldest cusp in the upper molars also seems to be the most stable cusp (at least in the M(1)). The most striking change in M(1) cusp proportions is a change in the relative size of the areas of the paracone and metacone. The combination of a small relative paracone and a large relative metacone generally characterizes specimens attributed to early Homo, and the presence of this character state in Australopithecus and Paranthropus suggests it may represent the primitive condition for the later part of the hominin clade. In contrast, nearly all later Homo taxa, with the exception of Homo antecessor, show the opposite condition (i.e. a relatively large paracone and a relatively small metacone). This change in the relationship between the relative sizes of the paracone and metacone is related to an isometric reduction of the absolute size of the metacone. This metacone reduction occurs in the context of relative stability in the paracone as crown size decreases. Among later Homo taxa, both Homo heidelbergensis and Homo neanderthalensis show a further reduction of the metacone and an enlargement of the hypocone. Fossil and contemporary H. sapiens samples show a trend toward increasing the relative size of the protocone and decreasing the relative size of the hypocone. In Europe, modern human M(1) cusp proportions are essentially reached during the Upper Paleolithic. Although some variation was documented among the fossil taxa, we suggest that the relative size of the M(1) paracone and metacone areas may be useful for differentiating the earliest members of our genus from subsequent Homo species.