The transition to full-time terrestrial bipedality is a hallmark of human evolution. A key correlate of human bipedalism is the development of longitudinal and transverse arches of the foot that provide a rigid propulsive lever and critical shock absorption during striding bipedal gait. Evidence for arches in the earliest well-known Australopithecus species, A. afarensis, has long been debated. A complete fourth metatarsal of A. afarensis was recently discovered at Hadar, Ethiopia. It exhibits torsion of the head relative to the base, a direct correlate of a transverse arch in humans. The orientation of the proximal and distal ends of the bone reflects a longitudinal arch. Further, the deep, flat base and tarsal facets imply that its midfoot had no ape-like midtarsal break. These features show that the A. afarensis foot was functionally like that of modern humans and support the hypothesis that this species was a committed terrestrial biped.