Reevaluating the functional implications of Australopithecus afarensis navicular morphology

The longitudinal arch is a unique characteristic of the human foot, yet the timing and pattern of its evolution remain controversial, in part due to the disagreement among researchers over which skeletal traits are the best indicators of its presence or absence. The small size of the human navicular tuberosity has previously been linked to the presence of a longitudinal arch, implying that the large tuberosity of early hominins such as Australopithecus afarensis reflects a flat foot. However, this hypothesis is at odds with other evidence of pedal form and function, such as metatarsal, tarsal, and footprint morphology, which show that a longitudinal arch was probably present in A. afarensis. This study reevaluates the morphometric affinities of the A. afarensis naviculars among other Plio-Pleistocene fossil hominins and anthropoid primates (N = 170). Multivariate cluster analyses show that all fossil hominin naviculars, including those attributed to A. afarensis, are most similar to modern humans. A measure of navicular tuberosity size quantified as the ratio of the tuberosity volume to the surface area of the talar facet shows that Ateles has the largest navicular tuberosity among the anthropoid sample and that there is no difference between highly arboreal and terrestrial taxa in this metric (e.g., Hylobates and Gorilla beringei). Instead, a relatively large navicular tuberosity may reflect the development of leg musculature associated with ankle plantarflexion. The functional inferences derived from the morphology of the A. afarensis naviculars are consistent with the morphology of the Laetoli footprints.