The Jinniushan hominin pedal skeleton from the late Middle Pleistocene of China

Bipedalism has long been recognized as the seminal adaptation of the hominin radiation and thus used to distinguish hominins from great ape fossils. Notwithstanding preconceptions and varied interpretations, the distinctive features of the modern human foot and accompanying striding gait, appear to be recent innovations that are largely absent in the earliest facultative bipeds. These distinctive features are mainly components of fixed longitudinal and transverse pedal arches, and of a uniquely derived hallucal metatarsophalangeal joint. They enhance ankle joint plantar flexor function and accommodate localized peak plantar pressures at the medial ball during terminal stance. To date, the paleontological record has yielded very little of the hominin foot, especially of the Middle Pleistocene hominins. New specimens from this time interval should help provide insights into the timing and pattern of what appears to be a mosaic pattern of evolution of the modern human foot features. Here we describe the fossil hominin foot skeleton recovered from the Jinniushan site, Liaoning Province, People's Republic of China. It affords a singular glimpse of the pedal morphology of a late Middle Pleistocene hominin (c.f. Homo heidlebergensis). Dated to 200 ka or older, this foot offers the earliest evidence for increased stability of the medial longitudinal arch, while retaining a number of primitive features apparently characteristic of robust premodern hominins, including lower arches and a less stable hallucal metatarsophalangeal joint (medial ball) than in modern humans. These features reflect different foot capabilities and suggest the bipedal stride of the Jinniushan hominin differed subtlety from that of modern humans.