Locomotory Adaptations in 3D Humerus Geometry of Xenarthra: Testing for Convergence

Three-dimensional (3D) models of fossil bones are increasingly available, thus opening a novel frontier in the study of organismal size and shape evolution. We provide an example of how photogrammetry can be combined with Geometric Morphometrics (GMM) techniques to study patterns of morphological convergence in the mammalian group of Xenarthra. Xenarthrans are currently represented by armadillos, sloths, and anteaters. However, this clade shows an incredibly diverse array of species and ecomorphotypes in the fossil record, including gigantic ground sloths and glyptodonts. Since the humerus is a weight-bearing bone in quadrupedal mammals and its morphology correlates with locomotor behavior, it provides an ideal bone to gain insight into adaptations of fossil species. A 3D sample of humerii belonging to extant and fossil Xenarthra allowed us to identify a significant phylogenetic signal and a strong allometric component in the humerus shape. Although no rate shift in the evolution of the humerus shape was recorded for any clade, fossorial and arboreal species humerii did evolve at significantly slower and faster paces, respectively, than the rest of the Xenarthran species. Significant evidence for morphological convergence found among the fossorial species and between the two tree sloth genera explains these patterns. These results suggest that the highly specialized morphologies of digging taxa and tree sloths represent major deviations from the plesiomorphic Xenarthran body plan, evolved several times during the history of the group

Ecomorphology of radii in Canidae: Application to fragmentary fossils from Plio-Pleistocene hominin assemblages

Fragmentary long bone material from fossil Carnivora is rarely considered to support palaeoenvironmental reconstructions. Here, we use morphometry of the radius in extant carnivorans of the dog family (Canidae) to reconstruct the palaeobiology of extinct canids from Olduvai Gorge, Tanzania (Bed I and II) and Koobi Fora, Kenya. We use radius morphometrics to predict adaptation to prey size and introduce a new method for quantifying canid habitat adaptations based on the geographic distributions of the extant species sampled. Linear Discriminant Function Analyses (DFA) and cluster neighbour-joining algorithms are employed to investigate radial morphometrics as described by 29 linear measurements. Results of our analyses suggest that a phylogenetic signal is present in radial morphometrics, even if it does not allow us to accurately discriminate among genera. A binary prey size categorisation of “small-medium” versus “large” prey can be more accurately predicted than a habitat categorisation scheme (Open, Mixed, Closed). The East African fossil specimens examined show morphometric affinities with the golden jackal (Canis aureus) and coyote (Canis latrans) and are likely attributable to the genus Canis. Fragmentary fossil specimens from Olduvai Gorge are predicted as habitat generalists (Open for Bed I and Mixed for Bed II) adapted for hunting small-medium prey, whereas the specimen from Koobi Fora was predicted as inhabiting mixed habitats and adapted for killing large prey. This study supports the inclusion of fossil Canidae in palaeoecological analyses attempting to clarify the palaeoenvironment of early hominin fossil sites

Ecogeographical Variation in Skull Shape of South-American Canids: Abiotic or Biotic Processes?

Species morphological changes can be mutually influenced by environmental or biotic factors, such as competition. South American canids represent a quite recent radiation of taxa that evolved forms very disparate in phenotype, ecology and behaviour. Today, in the central part of South America there is one dominant large species (the maned wolf, Chrysocyon brachyurus) that directly influence sympatric smaller taxa via interspecific killing. Further south, three species of similar sized foxes (Lycalopex spp.) share the same habitats. Such unique combination of taxa and geographic distribution makes South American dogs an ideal group to test for the simultaneous impact of climate and competition on phenotypic variation. Using geometric morphometrics, we quantified skull size and shape of 431 specimens belonging to the eight extant South American canid species: Atelocynus microtis, Cerdocyon thous, Ch. brachyurus, Lycalopex culpaeus, L. griseus, L. gymnocercus, L. vetulus and Speothos venaticus. South American canids are significantly different in both skull size and shape. The hypercarnivorous bush dog is mostly distinct in shape from all the other taxa while a degree of overlap in shape—but not size—occurs between species of the genus Lycalopex. Both climate and competition impacts interspecific morphological variation. We identified climatic adaptations as the main driving force of diversification for the South American canids. Competition has a lower degree of impact on their skull morphology although it might have played a role in the past, when canid community was richer in morphotypes

Elbow Joint Geometry in Bears (Ursidae, Carnivora): a Tool to Infer Paleobiology and Functional Adaptations of Quaternary Fossils

Bears are currently represented by eight species among Carnivora. Being all particularly large and generally plantigrade limits to certain extent their functional morphology so that inferences about their past diversification are difficult to achieve. We analyzed variation in bears’ elbow joint size and shape to reconstruct paleobiology of Quaternary fossil species. By using 2D geometric morphometrics, we were able to discriminate with high degree of accuracy species, locomotor and habitat adaptations among extant bears. The giant panda and the spectacled bear are well characterized by an enlarged medial epicondyle, while large members of the genus Ursus can be distinguished by their relatively longer and wider trochlea. Elbow joint size varies consistently among ecological categories of extant bears and is generally selected by discriminant function models providing a high degree of classification accuracy (> 80%). American genera Arctotherium and Arctodus are predicted as non-climbing species potentially adapted to open and mixed environments in agreement with their potentially opportunistic feeding behavior. They retain a wide medial epicondyle probably in relation to a high degree of forelimb dexterity. Cave bears are equally predicted as non-climbers adapted to open habitats while the middle Pleistocene Ursus deningeri and fossil Ursus arctos are generally classified as scansorial species with a preference towards mixed habitats. Our study is the first to demonstrate that fragmentary distal epiphyses also can be useful for interpreting functional morphology and ecological adaptations within the family Ursidae

Geometric morphometric analyses of sexual dimorphism and allometry in two sympatric snakes: Natrix helvetica (Natricidae) and Vipera berus (Viperidae)

The non-venomous grass snake (Natrix helvetica) and the venomous adder (Vipera berus) are two native species that are often found in sympatry in Great Britain and Europe. They occupy partially overlapping ecological niches and prey on small vertebrates, but use different feeding strategies. Here, we investigated the morphologies of grass snakes and adders from Dorset (UK) using twodimensional geometric morphometrics to assess the degree of sexual dimorphism in size and shape together with the relative impact of allometry and general body dimensions on head shape. Both species showed significant sexual dimorphism in head size, but not in head shape. We found a clear allometric pattern in N. helvetica, whereas allometry in V. berus was generally less pronounced. Body dimensions were strongly correlated with head shape in the grass snake, but not in the adder. The fact that V. berus is venomous appears to explain the lack of allometric patterns and the lack of an association between body dimensions and head shape. The high degree of size dimorphism identified in both species could originate from the advantages of reduced intraspecific competition that are conveyed by a partial differentiation in feeding morphology

Ecomorphology of Carnivora challenges convergent evolution

Convergent evolution is often reported in the mammalian order Carnivora. Their adaptations to particularly demanding feeding habits such as hypercarnivory and durophagy (consumption of tough food) appear to favour morphological similarities between distantly related species, especially in the skull. However, phylogenetic effect in phenotypic data might obscure such a pattern. We first validated the hypotheses that extant hypercarnivorous and durophagous large carnivorans converge in mandibular shape and form (size and shape). Hypercarnivores generally exhibit smaller volumes of the multidimensional shape and form space than their sister taxa, but this pattern is significantly different from random expectation only when hunting behaviour categorisations are taken into account. Durophages share areas of the morphospace, but this seems to be due to factors of contingency. Carnivorans that hunt in pack exhibit incomplete convergence while even stronger similarities occur in the mandible shape of solitary hunters due to the high functional demands in killing the prey. We identified a stronger phylogenetic signal in mandibular shape than in size. The quantification of evolutionary rates of changes suggests that mandible shape of solitary hunters evolved slowly when compared with other carnivorans. These results consistently indicate that the need for a strong bite force and robust mandible override sheer phylogenetic effect in solitary hunters

Macroevolutionary ecomorphology of the Carnivora skull: adaptations and constraints in the extant species

The mammalian order Carnivora is characterized by a broad taxonomic and ecological diversity. By using a large sample of extant species, we tested the impact of ecological factors on carnivoran skull (cranium and mandible) morphology, taking advantage of a combined geometric morphometrics and comparative method approach. We implemented several evolutionary models to account for different tempo and mode of evolution in size and shape data. These models validated the association between skull morphology and diet at the interspecific scale. The functional distinction between pinniped (aquatic) and fissiped (mostly terrestrial) taxa was found valid only in mandible shape and cranial size. High levels of morphological disparity and evolutionary rates were identified in specialized dietary groups, and positive association between rates and disparity was found for skull size. Cranium and mandible showed consistent patterns of covariation that reflect constrained functional processes, which stabilize the ecomorphological evolution of Carnivora. Aquatic adaptations allowed carnivorans to invade and persist within novel regions of the mandibular morphospace. This ecological shift did not increase morphological disparity but occurred at a faster rate than in terrestrial species. Those species exhibit a stronger level of cranio-mandibular covariation due to constraints imposed by more demanding masticatory adaptations

The impact of large terrestrial carnivores on Pleistocene ecosystems

At very high densities, populations of the largest herbivores, such as elephants, have devastating effects on the environment. What prevented widespread habitat destruction in the Pleistocene, when the ecosystem sustained many species of huge herbivores? We use data on predator–prey body mass relationships to predict the prey size ranges of large extinct mammalian carnivores, which were more diverse and much larger than living species. We then compare these prey size ranges with estimates of young mammoth sizes and show that juvenile mammoths and mastodons were within predicted prey size ranges of many of the Pleistocene carnivores. From this and other fossil evidence we argue that, by limiting population sizes of megaherbivores, large carnivores had a major impact on Pleistocene ecosystems

The potential and pitfalls of using simple dental metrics to infer the diets of African antelopes (Mammalia: Bovidae)

The use of mesowear to infer diets of extinct species is fast becoming widespread in palaeoecological studies. Nevertheless, traditional mesowear analyses suffer from a specimen number limitation, in that a minimum number of specimens identified to the species level is necessary to make accurate dietary predictions. This is problematic in many fossil African antelope (Mammalia: Bovidae) assemblages, where isolated teeth cannot always be assigned to species. Hereweexplore the possibility of using simple dental metrics to predict diets on the basis of individual teeth as well as gnathic rows using linear discriminant function analyses.We find that browsers are accurately classified at both the individual and species levels, across all models and tooth positions. Mixed feeders and grazers are classified accurately only sometimes, and this is probably a reflection of the more limited sample size of larger bodied species in our study. Body size was a highly significant predictor of the inaccurate classifications obtained in our models, with larger bodied species tending to grazing classifications and smaller bodied species browsing classifications. Nevertheless, the models correctly classify the majority of specimens we examined to their correct trophic group, as determined through stable isotope analyses or as defined through the literature. The methods outlined hold some promise for determining the diets of isolated fossil specimens unassigned to species in a simple manner and, when used in conjunction with other palaeodietary and palaeoecological proxies, may help determine palaeoenvironments more accurately