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Synopsis Analyses of form–function relationships are widely used to understand links between morphology, ecology, and adaptation across macroevolutionary scales. However, few have investigated functional trade-offs and covariance within and between the skull, limbs, and vertebral column simultaneously. In this study, we investigated the adaptive landscape of skeletal form and function in carnivorans to test how functional trade-offs among these skeletal regions contribute to ecological adaptations and the topology of the landscape. We found that morphological proxies of function derived from carnivoran skeletal regions exhibit trade-offs and covariation across their performance surfaces, particularly in the appendicular and axial skeletons. These functional trade-offs and covariation correspond as adaptations to different adaptive landscapes when optimized by various factors including phylogeny, dietary ecology, and, in particular, locomotor mode. Lastly, we found that the topologies of the optimized adaptive landscapes and underlying performance surfaces are largely characterized as a single gradual gradient rather than as rugged, multipeak landscapes with distinct zones. Our results suggest that carnivorans may already occupy a broad adaptive zone as part of a larger mammalian adaptive landscape that masks the form and function relationships of skeletal traits. 

Abstract The timing and nature of evolutionary shifts in the relative brain size of Primates have been extensively studied. Less is known, however, about the scaling of the brain-to-body size in their closest living relatives, i.e., among other members of Euarchontoglires (Dermoptera, Scandentia, Lagomorpha, Rodentia). Ordinary least squares (OLS), reduced major axis (RMA), and phylogenetic generalized least squares (PGLS) regressions were fitted to the largest euarchontogliran data set of brain and body mass, comprising 715 species. Contrary to previous inferences, lagomorph brain sizes (PGLS slope = 0.465; OLS slope = 0.593) scale relative to body mass similarly to rodents (PGLS = 0.526; OLS = 0.638), and differently than primates (PGLS = 0.607; OLS = 0.794). There is a shift in the pattern of the scaling of the brain in Primates, with Strepsirrhini occupying an intermediate stage similar to Scandentia but different from Rodentia and Lagomorpha, while Haplorhini differ from all other groups in the OLS and RMA analyses. The unique brain–body scaling relationship of Primates among Euarchontoglires illustrates the need for clade-specific metrics for relative brain size (i.e., encephalization quotients; EQs) for more restricted taxonomic entities than Mammalia. We created clade-specific regular and phylogenetically adjusted EQ equations at superordinal, ordinal, and subordinal levels. When using fossils as test cases, our results show that generalized mammalian equations underestimate the encephalization of the stem lagomorph Megalagus turgidus in the context of lagomorphs, overestimate the encephalization of the stem primate Microsyops annectens and the early euprimate Necrolemur antiquus, but provide similar EQ values as our new strepsirrhine-specific EQ when applied to the early euprimate Adapis parisiensis. 

Abstract The canine of saber‐toothed predators represents one of the most specialized dental structures known. Hypotheses about the function of hypertrophied canines range from display and conspecific interaction, soft food processing, to active prey acquisition. Recent research on the ontogenetic timing of skull traits indicates the adult canine can take years to fully erupt, but the consequences of prolonged eruption on inferences of canine functional morphology are missing from current discourse and have not been quantified. Here I evaluate hypotheses about adult canine bending strength and stiffness, respectively, during eruption in the felidSmilodon fatalis. Simulated eruption sequences of three adult canines were generated from specimen models to assess shifting cross‐sectional geometry properties, and bending strength and stiffness under laterally directed loads were estimated using finite element analysis. Consistent with beam theory expectations,S. fataliscanine cross‐sectional geometry is optimized for increased bending strength with increased erupted height. However, canine cross‐sectional geometry changes through eruption exaggerate rather than minimize lateral deflection. Spatial constraint for maximum root length from adjacent sensory structures in the maxilla and the recently identified universal power law are hypothesized to limit the growth capacity of canine anteroposterior length and, consequently, maintenance of bending stiffness through eruption. Instead, the joint presence of the deciduous and adult canines for >50% of the adult canine eruption period effectively increases canine mediolateral width and brings bending strength and stiffness estimates closer to theoretical optima. Similarly prolonged retention of deciduous canines in other sabertooths suggests dual‐canine buttressing is a convergently evolved strategy to maximize bending strength and stiffness. 

Abstract The evolution of organisms can be studied through the lens of developmental systems, as the timing of development of morphological features is an important aspect to consider when studying a phenotype. Such data can be challenging to obtain in fossil amniotes owing to the scarcity of their fossil record. However, the numerous remains of Rancho La Brea allow a detailed study of the postnatal changes in an extinct sabertoothed felid:Smilodon fatalis. Despite numerous previous studies on the ontogeny ofSmilodon, an important question remained open: how did the cubs ofSmilodonacquire and process food? By applying 3D geometric morphometrics and finite element analyses to 49 mandibles at various developmental stages (22 ofS. fatalis, 23 ofPanthera leo, and 4 of early diverging felids), we assess the changes in mandibular shape and performance during growth. Both lions and sabertooths exhibit a shift in mandibular shape, aligning with eruption of the lower carnassial. This marks the end of weaning in lions and suggests a prolonged weaning period inS. fatalisowing to its delayed eruption sequence. We also highlight distinct ontogenetic trajectories, withS. fatalisundergoing more postnatal mandibular shape changes. Finally, althoughS. fatalisappears more efficient thanP. leoat performing an anchor bite, this efficiency is acquired through ontogeny and at a quite late age. The delayed shape change compared withP. leoand the low biting efficiency during the growth inSmilodoncould indicate an extended duration of the parental care compared withP. leo. 

Synopsis Mammals exhibit a diverse range of limb morphologies that are associated with different locomotor ecologies and structural mechanics. Much remains to be investigated, however, about the combined effects of locomotor modes and scaling on the external shape and structural properties of limb bones. Here, we used squirrels (Sciuridae) as a model clade to examine the effects of locomotor mode and scaling on the external shape and structure of the two major limb bones, the humerus and femur. We quantified humeral and femoral morphologies using 3D geometric morphometrics and bone structure analyses on a sample of 76 squirrel species across their four major ecotypes. We then used phylogenetic generalized linear models to test how locomotor ecology, size, and their interaction influenced morphological traits. We found that size and locomotor mode exhibit different relationships with the external shape and structure of the limb bones, and that these relationships differ between the humerus and femur. External shapes of the humerus and, to a lesser extent, the femur are best explained by locomotor ecology rather than by size, whereas structures of both bones are best explained by interactions between locomotor ecology and scaling. Interestingly, the statistical relationships between limb morphologies and ecotype were lost when accounting for phylogenetic relationships among species under Brownian motion. That assuming Brownian motion confounded these relationships is not surprising considering squirrel ecotypes are phylogenetically clustered; our results suggest that humeral and femoral variation partitioned early between clades and their ecomorphologies were maintained to the present. Overall, our results show how mechanical constraints, locomotor ecology, and evolutionary history may enact different pressures on the shape and structure of limb bones in mammals. 

Madagascar is one of the world’s foremost biodiversity hotspots with more than 90% of its species endemic to the island. Malagasy carnivorans are one of only four extant terrestrial mammalian clades endemic to Madagascar. Although there are only eight extant species, these carnivorans exhibit remarkable phenotypic and ecological diversity that is often hypothesized to have diversified through an adaptive radiation. Here, we investigated the evolution of skull diversity in Malagasy carnivorans and tested if they exhibited characteristics of convergence and an adaptive radiation. We found that their skull disparity exceeds that of any other feliform family, as their skulls vary widely and strikingly capture a large amount of the morphological variation found across all feliforms. We also found evidence of shared adaptive zones in cranial shape between euplerid subclades and felids, herpestids and viverrids. Lastly, contrary to predictions of adaptive radiation, we found that Malagasy carnivorans do not exhibit rapid lineage diversification and only marginally faster rates of mandibular shape evolution and to a lesser extent cranial shape evolution, compared to other feliforms. These results reveal that exceptional diversification rates are not necessary to generate the striking phenotypic diversity that evolved in carnivorans after their dispersal to and isolation on Madagascar. 

The diversity of vertebrate skeletons is often attributed to adaptations to distinct ecological factors such as diet, locomotion, and sensory environment. Although the adaptive evolution of skull, appendicular skeleton, and vertebral column is well studied in vertebrates, comprehensive investigations of all skeletal components simultaneously are rarely performed. Consequently, we know little of how modes of evolution differ among skeletal components. Here, we tested if ecological and phylogenetic effects led to distinct modes of evolution among the cranial, appendicular and vertebral regions in extant carnivoran skeletons. Using multivariate evolutionary models, we found mosaic evolution in which only the mandible, hindlimb and posterior (i.e. last thoracic and lumbar) vertebrae showed evidence of adaptation towards ecological regimes whereas the remaining skeletal components reflect clade-specific evolutionary shifts. We hypothesize that the decoupled evolution of individual skeletal components may have led to the origination of distinct adaptive zones and morphologies among extant carnivoran families that reflect phylogenetic hierarchies. Overall, our work highlights the importance of examining multiple skeletal components simultaneously in ecomorphological analyses. Ongoing work integrating the fossil and palaeoenvironmental record will further clarify deep-time drivers that govern the carnivoran diversity we see today and reveal the complexity of evolutionary processes in multicomponent systems. 

The evolutionary shift from a single-element ear, multi-element jaw to a multi-element ear, single-element jaw during the transition to crown mammals marks one of the most dramatic structural transformations in vertebrates. Research on this transformation has focused on mammalian middle-ear evolution, but a mandible comprising only the dentary is equally emblematic of this evolutionary radiation. Here, we show that the remarkably diverse jaw shapes of crown mammals are coupled with surprisingly stereotyped jaw stiffness. This strength-based morphofunctional regime has a genetic basis and allowed mammalian jaws to effectively resist deformation as they radiated into highly disparate forms with markedly distinct diets. The main functional consequences for the mandible of decoupling hearing and mastication were a trade-off between higher jaw stiffness versus decreased mechanical efficiency and speed compared with non-mammals. This fundamental and consequential shift in jaw form–function underpins the ecological and taxonomic diversification of crown mammals. This article is part of the theme issue ‘The mammalian skull: development, structure and function’. 

Body size is often hypothesized to facilitate or constrain morphological diversity in the cranial, appendicular, and axial skeletons. However, how overall body shape scales with body size ( i.e. , body shape allometry) and whether these scaling patterns differ between ecological groups remains poorly investigated. Here, we test whether and how the relationships between body shape, body size, and limb lengths differ among species with different locomotor specializations, and describe the underlying morphological components that contribute to body shape evolution among squirrel (Sciuridae) ecotypes. We quantified the body size and shape of 87 squirrel species from osteological specimens held at museum collections. Using phylogenetic comparative methods, we first found that body shape and its underlying morphological components scale allometrically with body size, but these allometric patterns differ among squirrel ecotypes: chipmunks and gliding squirrels exhibited more elongate bodies with increasing body sizes whereas ground squirrels exhibited more robust bodies with increasing body size. Second, we found that only ground squirrels exhibit a relationship between forelimb length and body shape, where more elongate species exhibit relatively shorter forelimbs. Third, we found that the relative length of the ribs and elongation or shortening of the thoracic region contributes the most to body shape evolution across squirrels. Overall, our work contributes to the growing understanding of mammalian body shape evolution and how it is influenced by body size and locomotor ecology, in this case from robust subterranean to gracile gliding squirrels.