Knuckle-walking in Sahelanthropus? Locomotor inferences from the ulnae of fossil hominins and other hominoids.

ABSTRACT

Because the ulna supports and transmits forces during movement, its morphology can signal aspects of functional adaptation. To test whether, like extant apes, some hominins habitually recruit the forelimb in locomotion, we separate the ulna shaft and ulna proximal complex for independent shape analyses via elliptical Fourier methods to identify functional signals. We examine the relative influence of locomotion, taxonomy, and body mass on ulna contours in Homo sapiens (n = 22), five species of extant apes (n = 33), two Miocene apes (Hispanopithecus and Danuvius), and 17 fossil hominin specimens including Sahelanthropus, Ardipithecus, Australopithecus, Paranthropus, and early Homo. Ulna proximal complex contours correlate with body mass but not locomotor patterns, while ulna shafts significantly correlate with locomotion. African apes' ulna shafts are more robust and curved than Asian apes and are unlike other terrestrial mammals (including other primates), curving ventrally rather than dorsally. Because this distinctive curvature is absent in orangutans and hylobatids, it is likely a function of powerful flexors engaged in wrist and hand stabilization during knuckle-walking, and not an adaptation to climbing or suspensory behavior. The OH 36 (purported Paranthropus boisei) and TM 266 (assigned to Sahelanthropus tchadensis) fossils differ from other hominins by falling within the knuckle-walking morphospace, and thus appear to show forelimb morphology consistent with terrestrial locomotion. Discriminant function analysis classifies both OH 36 and TM 266 with Pan and Gorilla with high posterior probability. Along with its associated femur, the TM 266 ulna shaft contours and its deep, keeled trochlear notch comprise a suite of traits signaling African ape-like quadrupedalism. While implications for the phylogenetic position and hominin status of S. tchadensis remain equivocal, this study supports the growing body of evidence indicating that S. tchadensis was not an obligate biped, but instead represents a late Miocene hominid with knuckle-walking adaptations.