Knuckle-walking anteater: a convergence test of adaptation for purported knuckle-walking features of African Hominidae

Appeals to synapomorphic features of the wrist and hand in African apes, early hominins, and modern humans as evidence of knuckle-walking ancestry for the hominin lineage rely on accurate interpretations of those features as adaptations to knuckle-walking locomotion. Because Gorilla, Pan, and Homo share a relatively close common ancestor, the interpretation of such features is confounded somewhat by phylogeny. The study presented here examines the evolution of a similar locomotor regime in New World anteaters (order Xenarthra, family Myrmecophagidae) and uses the terrestrial giant anteater (Myrmecophaga tridactyla) as a convergence test of adaptation for purported knuckle-walking features of the Hominidae. During the stance phase of locomotion, Myrmecophaga transmits loads through flexed digits and a vertical manus, with hyperextension occurring at the metacarpophalangeal joints of the weight-bearing rays. This differs from the locomotion of smaller, arboreal anteaters of outgroup genera Tamandua and Cyclopes that employ extended wrist postures during above-branch quadrupedality. A number of features shared by Myrmecophaga and Pan and Gorilla facilitate load transmission or limit extension, thereby stabilizing the wrist and hand during knuckle-walking, and distinguish these taxa from their respective outgroups. These traits are a distally extended dorsal ridge of the distal radius, proximal expansion of the nonarticular surface of the dorsal capitate, a pronounced articular ridge on the dorsal aspects of the load-bearing metacarpal heads, and metacarpal heads that are wider dorsally than volarly. Only the proximal expansion of the nonarticular area of the dorsal capitate distinguishes knuckle-walkers from digitigrade cercopithecids, but features shared with digitigrade primates might be adaptive to the use of a vertical manus of some sort in the stance phase of terrestrial locomotion. The appearance of capitate nonarticular expansion and the dorsal ridge of the distal radius in the hominin lineage might be indicative of a knuckle-walking ancestry for bipedal hominins if interpreted within the biomechanical and phylogenetic context of hominid locomotor evolution.