The relative size of the calcaneal tuber reflects heel strike plantigrady in African apes and humans

Objectives: The positional repertoire of the human-chimpanzee last common ancestor is critical for reconstructing the evolution of bipedalism. African apes and humans share a heel strike plantigrade foot posture associated with terrestriality. Previous research has established that modern humans have a relatively large and intrinsically robust calcaneal tuber equipped to withstand heel strike forces associated with bipedal walking and running. However, it is unclear whether African apes have a relatively larger calcaneal tuber than non-heel-striking primates, and how this trait might have evolved among anthropoids. Here, I test the hypothesis that heel-striking primates have a relatively larger calcaneal tuber than non-heel-striking primates. Methods: The comparative sample includes 331 individuals and 53 taxa representing hominoids, cercopithecoids, and platyrrhines. Evolutionary modeling was used to test for the effect of foot posture on the relative size of the calcaneal tuber in a phylogenetic framework that accounts for adaptation and inertia. Bayesian evolutionary modeling was used to identify selective regime shifts in the relative size of the calcaneal tuber among anthropoids. Results: The best fitting evolutionary model was a Brownian motion model with regime-dependent trends characterized by relatively large calcaneal tubers among African apes and humans. Evolutionary modeling provided support for an evolutionary shift toward a larger calcaneal tuber at the base of the African ape and human clade. Conclusions: The results of this study support the view that African apes and humans share derived traits related to heel strike plantigrady, which implies that humans evolved from a semi-terrestrial quadrupedal ancestor.