Locomotor adaptation in the hominoid clavicle through ontogeny

Reconstructions of the locomotor behavior of early hominins have been hindered by our incomplete understanding of the form-function relationship in the extant hominoid shoulder. Although extensive research has highlighted the role of the highly mobile shoulder in supporting the locomotor diversity and versatility observed in hominoids, the contribution of the clavicle and its morphological diversity to shoulder function remains significantly underexplored. In this study, we analyzed the cross-sectional geometry of the ape clavicle using a large ontogenetic sample to identify new osteological signals related to locomotor adaptation in the shoulder. We assessed the interspecific and intraspecific differences in cortical bone distribution, with ratios of cortical properties describing the relative eccentricity of the cross section (the ratio of the second moments of area about the maximum [I_MAX] and minimum [I_MIN] principal axes [I_MAX/I_MIN]), the orientation of the anatomical plane that eccentricity is occurring in (the ratio of the second moments of area relative to the craniocaudal [I_X] and dorsoventral [I_Y] axes [I_X/I_Y]), and the relative proportion of cortical bone in each section. Our analyses demonstrate that the hominoid clavicle holds strong signals of locomotor adaptation that can be identified both across taxa and through ontogeny. Gibbons and orangutans have a relatively uniform clavicular cortical geometry throughout life, with gibbon clavicles built to best withstand habitual, unidirectional bending forces and orangutan clavicles remodeled to resist unpredictable, multidirectional loading. Furthermore, we find a clear signal of increased clavicular bending in the same portion of the diaphysis through ontogeny in the cortical geometry of chimpanzees and gorillas, likely reflecting both the shifts toward terrestriality through ontogeny and bending rigidity needed for continued arboreality at a larger body mass. Ultimately, these results are promising for the identification of locomotor adaptation in the shoulder of early hominins, especially Australopithecus, and highlight the key structural role of the clavicle in ape locomotion.