Long bone shape variation in the forelimb of Rhinocerotoidea: relation with size, body mass and body proportions

In quadrupeds, limb bones are strongly affected by functional constraints linked to weight support, but few studies have addressed the complementary effects of mass, size and body proportions on limb bone shape. During their history, Rhinocerotoidea have displayed a great diversity of body masses and relative size and proportions of limb bones, from small tapir-like forms to giant species. Here, we explore the evolutionary variation of shapes in forelimb bones and its relationship with body mass in Rhinocerotoidea. Our results indicate a general increase in robustness and greater development of muscular insertions in heavier species, counteracting the higher weight loadings induced by an increased body mass. The shape of the humerus changes allometrically and exhibits a strong phylogenetic signal. Shapes of the radius and ulna display a stronger link with body mass repartition than with the absolute mass itself. Congruent shape variation between the humerus and the proximal part of the ulna suggests that the elbow joint is comprised of two strongly covariant structures. In addition, our work confirms the uniqueness of giant Paraceratheriidae among Rhinocerotoidea, whose shape variation is related to both a high body mass and a cursorial forelimb construction.