Exploring bone volume and skeletal weight in the Middle Pleistocene humans from the Sima de los Huesos site (Sierra de Atapuerca, Spain).

Body mass estimation in fossil human species is a crucial topic in paleoanthropology as it yields information about ecologically relevant characteristics. Nevertheless, variables crucial to body mass estimation such as bone volume and skeletal weight have never before been calculated in a fossil human species. The exceptional state of preservation of several fossil human long bones from the Sima de los Huesos (SH) Middle Pleistocene site, in the Sierra de Atapuerca, makes it possible to calculate for the first time the absolute bone volume in five complete long bones (two femora and three humeri) of a fossil human species, an approach not possible in fragmentary or poorly preserved fossils. We have relied on computed tomography scans and 3D reconstructions to calculate bone volume. A sample of 62 complete bones of robust recent humans was also used for comparative purposes. The male SH femora (weight-bearing bones) and humeri (non-weight-bearing bones) have, relative to their size, greater bone volume (volume of bone tissue over total bone volume) than the equivalent bones in our recent human sample. As mass is volume × density, and bone tissue density (as a material) is similar across mammals, we calculate bone mass, and our results show that the SH hominins had on average heavier long bones than extant humans of the same size. From the femoral weight at hand, we have estimated the total skeletal weight in two SH individuals, which is about 36% heavier than in the recent humans of the equivalent body size. Using different methods and skeletal variables, including skeletal weight, to estimate body mass in these two SH humans, we highlight the considerable differences in body mass estimates we obtained, and that the largest body mass estimate is the one based on the skeletal weight. Our results suggest that we cannot assume the same relative proportion of bone volume and bone and skeletal weight characterized the entire genus Homo. Given that skeletal weight has a significant influence on body mass, current body mass estimates of fossil Homo specimens could be systematically underestimated. Thus, the significantly larger bone volume and heavier bones, probably throughout the entire skeleton, of SH humans could have had consequences for many biological parameters in this Pleistocene population and considerable importance for studies focusing on adaptive and ecologically relevant characteristics. Although more recent human samples should be analyzed, in our view, the high skeletal robusticity of the SH sample, including larger bone volume and skeletal weight, is part of their adaptive body type selected for throughout the Pleistocene to support different mechanical and activity regimes and formed under tight genetic control, including control over bone formative and regulatory processes.