Homo in the middle pleistocene: Hypodigms, variation, and species recognition

It is generally accepted that modern humans evolved in Africa. This consensus has emerged in the last two decades, as molecular evidence has been coupled with findings from paleontology and prehistory. Patterns of DNA variation in living populations, morphology of fossils, and archeological traces can all be read to show that our species has deep roots in Africa and began to disperse into other regions only late in the Pleistocene. There are still questions about the timing of these dispersals and the extent to which modern people replaced or exchanged genes with other, more archaic groups, but new problems are now coming more clearly into focus. These relate to the ancestry of Homo sapiens in the Middle Pleistocene. In this essay, I emphasize the evidence available from Middle Pleistocene localities in Africa and Europe, exploring variation among individuals, composition of hypodigms, species-level taxonomy, and evolutionary relationships of the hominin populations. One obvious difficulty is that the fossils are scarce. For the most part, only incomplete crania are available from African localities. The record is more comprehensive for the Sima de los Huesos in Spain, but principally crania and jaws are known from other European sites. This means that skull traits and measurements must provide the basis for sorting individuals to groups and building differential diagnoses. For much of the material, dating is still poorly controlled, although a few of the most important assemblages can now be placed more securely within a Middle Pleistocene chronological framework. Despite these constraints, it is possible to point toward tentative solutions.