Did knuckle walking evolve twice?

Abstract

Although African great apes share a similar quadrupedal locomotor behaviour, there are marked differences in hand morphology and size between the species. Hence, whilst all three species (two genera) of African ape frequently knuckle walk as adults, debate remains as to whether this behaviour is derived from a common ancestor or whether it evolved in parallel in chimpanzees and gorillas. This exploratory morphometric study of the sub-adult and adult wrist of these two genera aims to contribute to this debate. A total of twenty-seven dimensions of the lunate, triquetral, hamate and capitate of sub-adult and adult Pan troglodytes and Gorilla gorilla were analysed in order to determine whether carpal dimensions are generally ontogenetically scaled, and whether differences in growth trajectories, or length of growth, and adult morphologies can be explained by behavioural differences between the two species. Only 56% of all dimensions studied were ontogenetically scaled in sub-adults and some of these dimensions exhibit differing adult proportions between the two species. In general, the dimensions analysed fell into two categories: Pan and Gorilla either follow the same growth trajectories (Pattern A) or the Pan reduced major axis (RMA) regressions were significantly transposed above those of Gorilla (Pattern B). Additionally, it was found that Gorilla carpals appear to cease growing relatively earlier than those of Pan. While a small number of differences, notably those of the lunate, can be accounted for by differences in behaviour between the species, the majority of differences indicate heterochronic modifications of development during evolution, which correspond to kinematic differences in knuckle walking between the African great apes. In light of morphological, behavioural and ecological data currently available it is parsimonious to suggest that knuckle walking has evolved in parallel in the two lineages.