Homo erectus and Middle Pleistocene hominins: brain size, skull form, and species recognition.

Hominins that differ from Homo erectus, the Neanderthals, and recent humans are known from Middle Pleistocene localities across the Old World. The taxonomic status of these populations has been clouded by controversy. Perhaps the most critical problem has been an incomplete understanding of variation in skull form. Here, both H. erectus and later mid-Pleistocene hominins are the focus of an investigation aimed at clarifying the relationships among brain volume, basicranial dimensions, neurocranial shape, and certain facial characters. Brain size in H. erectus averages about 950 cm(3), while in a series of Middle Pleistocene crania from Africa and Europe, volume is about 1230 cm(3). If encephalization is the primary mechanism operating in the mid-Pleistocene, then diverse aspects of cranial form cannot all be treated as independent variables. Correlation is utilized to examine the associations among measurements for more than 30 H. erectus crania that are reasonably well preserved. A similar approach is used with the Middle Pleistocene sample. Patterns of covariation are compared in order to assess integration. Next, factor analysis is applied to the H. erectus specimens in an attempt to identify modules, tightly integrated traits that can evolve independently. Studies of the variation within H. erectus are followed by direct comparisons with the Middle Pleistocene population. Discriminant functions facilitate the description of intergroup differences. Traits that vary independently from brain volume include anterior frontal broadening, lateral expansion of the parietal vault, elevation of the lambda-inion chord, and rounding of the sagittal contour of the occipital. This finding helps to resolve the problem of species recognition. Neurocranial proportions as well as characters from the cranial base and face can be incorporated into a differential diagnosis for the mid-Pleistocene sample. Evidence presented here supports arguments for speciation in the Middle Pleistocene.