Conversely, the covariation of the zeugopodium bones seems more associated with body mass, particularly for the radius-ulna pair. The integration of the stylopodium elements does not seem to relate to body mass in our sample, which suggests a greater effect of shared developmental factors. The forelimb integration appears higher and more related to body mass than that of the hind limb, suggesting a specialization for weight support. At the interspecific level, the shape covariation is roughly similar between all pairs of bones and mainly concerns the muscular insertions related to powerful flexion and extension movements. Our results indicate that the appendicular skeleton of modern rhinos is a strongly integrated structure. We used a 3D geometric morphometric approach to describe the shape covariation of the six bones composing the stylopodium and zeugopodium both among and within species. We investigate the five modern rhinoceros species, which display an important range of bodyweight. Here, we propose to explore the covariation patterns of the long bones in heavy animals and their link to body mass. Among these constraints, body mass is considered strongly to influence its integration but its effect on shape covariation has rarely been addressed in mammals, especially in heavy taxa. The appendicular skeleton of tetrapods is a particularly integrated structure due to the shared developmental origin or similar functional constraints exerted on its elements.
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