Function, ontogeny and canalization of shape variance in the primate scapula

Primates have shoulders adapted to a wide range of locomotor functions from terrestrial pronograde quadrupedalism to highly arboreal suspensory behaviours. The shape of the scapula tightly follows these functional differences. Previous analyses of primate postcrania, including the scapula, indicate that quadrupedal monkeys are less variable than non-quadrupeds. It was previously suggested that this difference was due to a relationship between the strength of stabilizing selection and the functional demands of the upper limb. Here it is shown that intraspecific scapular shape variance is highly correlated with the degree of committed quadrupedalism. Primates that engage in frequent suspensory behaviours (e.g. apes and ateline monkeys) average twice the amount of shape variance as quadrupeds (e.g. Old World monkeys and Saimiri). Because this difference in intraspecific shape variance is apparent in infants and does not increase or decrease appreciably over ontogeny, it is not likely that differences in postnatal growth, neuromuscular control or environmental factors such as habitat structure/composition are the primary contributors to differences in adult shape variance. Instead variance in embryonic factors that affect the shape/size of the scapula or epigenetic factors associated with muscle attachments are more likely candidates. In particular, the heterogeneous functional demands of the non-quadrupedal shoulder probably reduce the stringency of stabilizing selection, resulting in the persistence into adulthood of increased amounts of embryonically generated scapular shape variance.