Among amniotes, numerous lineages are subject to an evolutionary trend toward body mass and size increases. Large terrestrial species may face important constraints linked to weight bearing, and the limb segments are particularly affected by such constraints due to their role in body support and locomotion. Such groups showing important limb modifications related to high body mass have been called “graviportal.” Often considered graviportal, rhinoceroses are among the heaviest terrestrial mammals and are thus of particular interest to understand the limb modifications related to body mass and size increase. Here, we present a morphofunctional study of the shape variation of the limb long bones among the five living rhinos to understand how the shape may vary between these species in relation with body size, body mass and phylogeny. We used three dimensional geometric morphometrics and comparative analyses to quantify the shape variation. Our results indicate that the five species display important morphological differences depending on the considered bones. The humerus and the femur exhibit noticeable interspecific differences between African and Asiatic rhinos, associated with a significant effect of body mass. The radius and ulna are more strongly correlated with body mass. While the tibia exhibits shape variation both linked with phylogeny and body mass, the fibula displays the greatest intraspecific variation. We highlight three distinct morphotypes of bone shape, which appear in accordance with the phylogeny. The influence of body mass also appears unequally expressed on the different bones. Body mass increase among the five extant species is marked by an increase of the general robustness, more pronounced attachments for muscles and a development of medial parts of the bones. Our study underlines that the morphological features linked to body mass increase are not similar between rhinos and other heavy mammals such as elephants and hippos, suggesting that the weight bearing constraint can lead to different morphological responses.

Many tetrapod lineages show extreme increases in body mass in their evolutionary history, associated with important osteological changes. The ankle joint, essential for foot movement, is assumed to be particularly affected in this regard. We investigated the morphological adaptations of the astragalus and the calcaneus in Rhinocerotidae, and analysed them in light of a comparative analysis with other Perissodactyla. We performed 3D geometric morphometrics and correlated shape with centroid size of the bone and body mass of the species. Our results show that mass has an influence on bone shape in Rhinocerotidae and in Perissodactyla, but this is not as strong as expected. In heavy animals the astragalus has a flatter trochlea, orientated more proximally, associated with a more upright posture of the limb. The calcaneus is more robust, possibly to sustain the greater tension force exerted by the muscles during plantarflexion. Both bones show wider articular facets, providing greater cohesion and better dissipation of the loading forces. The body plan of the animals also has an influence. Short-legged Teleoceratina have a flatter astragalus than the other rhinocerotids. Paraceratherium has a thinner calcaneus than expected. This study clarifies adaptations to high body weight among Rhinocerotidae and calls for similar investigations in other groups with massive forms.

Weight support is a strong functional constraint modelling limb bones in heavy quadrupeds. However, the complex relations between bone shape, mass, size and body proportions have been poorly explored. Rhinocerotoidea is one of the groups showing the highest body mass reached by terrestrial mammals through time. Here, we explore the evolutionary variation of shape in hindlimb stylopod and zeugopod bones and its relationship with mass, size and gracility in this superfamily. Our results show that bones undergo a general increase in robustness towards high masses, associated with reinforcements of the main muscle insertions. The shape of the femur, carrying a marked phylogenetic signal, varies conjointly with mass, size and gracility, whereas that of the tibia appears related to gracility and mass only. The shape of the fibula does not vary according to that of the tibia. Moreover, congruent variation of shape between the distal part of the femur and the complete tibia underlines the potentially strong covariation of the elements constituting the knee joint. These results, coupled with those previously obtained from forelimb study, allow a better comprehension of the relationship between bone shape and mass among Rhinocerotoidea, and a refining of the concept of ‘graviportality’ in this superfamily.

To date there are few quantitative studies investigating the evolution of tooth shape and function in aglyphous snakes in relation to diet. A considerable evolutionary decrease in body size is observed in whip snakes of the genus Dolichophis and their sister-group Eirenis. This was coupled with a considerable shift in diet from a regime consisting mainly of prey with endoskeleton to prey bearing a hard exoskeleton. Three-dimensional (3D) geometric morphometrics revealed that the maxillary and palatine teeth of E. persicus are blunt and conical in shape, while the same teeth are sharp and elongated in E. punctatolineatus and D. schmidti. Blunt and conically shaped teeth, as observed in E. persicus, seem to be more adapted for biting hard-bodied, arthropod prey. In contrast, the sharp and elongated teeth in Dolichophis and E. punctatolineatus, are likely specialized for puncturing prey with an endoskeleton. The results of a finite element analysis confirms that during the biting of a hard-bodied prey, the generated stresses in E. persicus teeth are well below the von Mises yield criterion, while in D. schmidti the value is roughly two to three times higher, indicating that E. persicus teeth are better suited for biting hard-bodied prey such as arthropods.

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.