The Morphology of the Bovid Calcaneus: Function, Phylogenetic Signal, and Allometric Scaling

Despite its clear functional role in hock (ankle) plantarflexion, the bovid calcaneus has been understudied with respect to the functional constraints imposed by locomotion in differing habitats, the allometric influence of inter-specific body size differences, and phylogenetic signal. This study uses a comparative sample of extant bovid species to shed light on the evolution of bovid calcaneal morphology. I measured eight linear measurements on 204 calcaneus specimens representing 41 extant bovid species. Using a morphological body size proxy validated against published species-mean body mass estimates, I performed Ordinary Least Squares regression to examine the allometric relationships of each measurement with body size. I classified each bovid species to a preferred habitat type based on published literature, and performed Phylogenetic Generalized Least Squares (PGLS) to test for differences in morphology between bovid taxa with different preferred habitats while considering evolutionary relatedness. I visualized morphological differences between taxa using Principal Components Analysis plotted in a phylomorphospace. Results demonstrate that several measurements of the bovid calcaneus have an allometric relationship to body size. The functional length of the calcaneus scales with negative allometry, which likely maintains a comparable safety factor within the calcaneal tuber at larger body sizes. While open-habitat bovids have relatively shorter calcaneal tubers, this difference is not significant when controlling for the influence of body size and phylogenetic signal using PGLS. Among bovid tribes that have a deep evolutionary history of adaptation to open habitats, Antilopini have relatively longer calcaneal tubers than Alcelaphini or Hippotragini, which may reflect the unique importance of stotting behavior in predator avoidance among antelopins. Overall, the morphology of the bovid calcaneus has been shaped by a complex interaction of phylogenetic and body-size constraints as well as adaptation to modes of predator avoidance mediated by preferred habitat.