CT-based description and phyletic evaluation of the archaic human calvarium from Ceprano, Italy

The discovery in 1994, of a fossilized human calvarium near Ceprano, Italy, dated about 800-900 thousand years before present, opened a new page for the study of human evolution in Europe. It extended the continental fossil record over the boundary between Early and Middle Pleistocene for the first time and revealed the cranial morphology of humans that where probably ancestral to both Neanderthals and modern Homo sapiens. A tomographic analysis of the Italian specimen is reported here in order to describe size and shape, vascular traces, and other features of the endocranium, as well as some relevant ectocranial traits (particularly of the frontal region). Our results show that the Ceprano calvarium displays plesiomorphies shared by early Homo taxa, involving a general archaic phenotype. At the same time, the presence of some derived features suggests a phylogenetic relationship with the populations referred to the subsequent polymorphic species H. heidelbergensis. The morphology of the supraorbital structures is different from the double-arched browridge of the African H. ergaster, while its superior shape shows similarities with African Middle Pleistocene specimens (Bodo, Kabwe). In contrast, the relationship between supraorbital torus and frontal squama points to an archaic pattern of the relationship between face and vault, associated to moderately narrow frontal lobes and limited development of the upper parietal areas. Despite a nonderived endocranial shape, the increase of cranial capacity (related to a general endocranial widening) and the probable absence of a clear occipital projection also suggest an evolutionary independence from the Asian H. erectus lineage. This analysis therefore supports the conclusion that the Ceprano calvarium represents the best available candidate for the ancestral phenotype of the cranial variation observed among Middle Pleistocene fossil samples in Africa and Europe. Nevertheless, a proper taxonomic interpretation of this crucial specimen remains puzzling.