Bipedalism has traditionally been regarded as the fundamental adaptation that sets hominids apart from other primates. Fossil I evidence demonstrates that by 4.1 million years ago(1), and perhaps earlier(2), hominids exhibited adaptations to bipedal walking. At present, however, the fossil record offers little information about the origin of bipedalism, and despite nearly a century of research on existing fossils and comparative anatomy, there is still no consensus concerning the mode of locomotion that preceded bipedalism(3-10). Here we present evidence that fossils attributed to Australopithecus anamensis (KNM-ER 20419)(11) and A. afarensis (AL 288-1)(12) retain specialized wrist morphology associated with knuckle-walking. This distal radial morphology differs from that of later hominids and non-knuckle-walking anthropoid primates, suggesting that knuckle-walking is a derived feature of the African ape and human clads. This removes key morphological evidence for a Pan-Gorilla clade, and suggests that bipedal hominids evolved from a knuckle-walking ancestor that was already partly terrestrial.