Borja Figueirido

Cenozoic climate change influences mammalian evolutionary dynamics

Global climate change is having profound impacts on the natural world. However, climate influence on faunal dynamics at macroevolutionary scales remains poorly understood. In this paper we investigate the influence of climate over deep time on the diversity patterns of Cenozoic North American mammals. We use factor analysis to identify temporally correlated assemblages of taxa, or major evolutionary faunas that we can then study in relation to climatic change over the past 65 million years. These taxa can be grouped into six consecutive faunal associations that show some correspondence with the qualitative mammalian chronofaunas of previous workers. We also show that the diversity pattern of most of these chronofaunas can be correlated with the stacked deep-sea benthic foraminiferal oxygen isotope (δ18O) curve, which strongly suggests climatic forcing of faunal dynamics over a large macroevolutionary timescale. This study demonstrates the profound influence of climate on the diversity patterns of North American terrestrial mammals over the Cenozoic.

Shape at the cross-roads: homoplasy and history in the evolution of the carnivoran skull towards herbivory

Patterns of skull shape in Carnivora provide examples of parallel and convergent evolution for similar ecomorphological adaptations. However, although most researchers report on skull homoplasies among hypercarnivorous taxa, evolutionary trends towards herbivory remain largely unexplored. In this study, we analyse the skull of the living herbivorous carnivorans to evaluate the importance of natural selection and phylogenetic legacy in shaping the skulls of these peculiar species. We quantitatively estimated shape variability using geometric morphometrics. A principal components analysis of skull shape incorporating all families of arctoid carnivorans recognized several common adaptations towards herbivory. Ancestral state reconstructions of skull shape and the reconstructed phylogenetic history of morphospace occupation more explicitly reveal the true patterns of homoplasy among the herbivorous carnivorans. Our results indicate that both historical constraints and adaptation have interplayed in the evolution towards herbivory of the carnivoran skull, which has resulted in repeated patterns of biomechanical homoplasy.

SKULL SHAPE EVOLUTION IN DUROPHAGOUS CARNIVORANS

In this article, we investigate convergent evolution toward durophagy in carnivoran skull shape using geometric morphometrics in a sample of living and extinct species. Principal components analysis indicate that, in spite of the different dietary resources consumed by durophages—that is, bone‐crackers and bamboo‐feeders—both groups of carnivorans share portions of skull phenotypic spaces. We identify by discriminant analyses a shared set of adaptations toward durophagy in the skull of carnivores. However, ancestral states indicate that although durophages reached similar phenotypes, the evolutionary pathways that they followed are different depending upon the family to which they belong. Furthermore, while the carnivoran cranium more closely reflects the nature of the resources consumed—that is, soft or hard and tough items—the mandible shows particular feeding adaptations—that is, bamboo or bone. This finding supports the interpretation that the mandible has more evolutionary plasticity than the cranium, which is more limited to evolve toward a particular feeding adaptation. However, we find that the shapes of the cranium and the mandible are highly integrated for the whole order Carnivora. Published studies of teratological cats and dogs indicate that the role of internal constraints in shaping this pattern of integration is absent or weak and malleable by selection.

Biomechanical Consequences of Rapid Evolution in the Polar Bear Lineage

The polar bear is the only living ursid with a fully carnivorous diet. Despite a number of well-documented craniodental adaptations for a diet of seal flesh and blubber, molecular and paleontological data indicate that this morphologically distinct species evolved less than a million years ago from the omnivorous brown bear. To better understand the evolution of this dietary specialization, we used phylogenetic tests to estimate the rate of morphological specialization in polar bears. We then used finite element analysis (FEA) to compare the limits of feeding performance in the polar bear skull to that of the phylogenetically and geographically close brown bear. Results indicate that extremely rapid evolution of semi-aquatic adaptations and dietary specialization in the polar bear lineage produced a cranial morphology that is weaker than that of brown bears and less suited to processing tough omnivorous or herbivorous diets. Our results suggest that continuation of current climate trends could affect polar bears by not only eliminating their primary food source, but also through competition with northward advancing, generalized brown populations for resources that they are ill-equipped to utilize.

Constraint and adaptation in the evolution of carnivoran skull shape

Abstract The evolutionary history of the Order Carnivora is marked by episodes of iterative evolution. Although this pattern is widely reported in different carnivoran families, the mechanisms driving the evolution of carnivoran skull morphology remain largely unexplored. In this study we use coordinate-point extended eigenshape analysis (CP-EES) to summarize aspects of skull shape in large fissiped carnivores. Results of these comparisons enable the evaluation of the role of different factors constraining the evolution of carnivoran skull design. Empirical morphospaces derived from mandible anatomy show that all hypercarnivores (i.e., those species with a diet that consists almost entirely of vertebrate flesh) share a set of traits involved in a functional compromise between bite force and gape angle, which is reflected in a strong pattern of morphological convergence. Although the paths followed by different taxa to reach this “hypercarnivore shape-space” differ because of phylogenetic constraints, the morphological signature of hypercarnivory in the mandible is remarkably narrow and well constrained. In contrast, CP-EES of cranial morphology does not reveal a similar pattern of shape convergence among hypercarnivores. This suggests a lesser degree of morphological plasticity in the cranium compared to the mandible, which probably results from a compromise between different functional demands in the cranium (e.g., feeding, vision, olfactory sense, and brain processing) whereas the mandible is only involved in food acquisition and processing. Combined analysis of theoretical and empirical morphospaces for these skull data also show the lower anatomical disparity of felids and hyaenids compared to canids and ursids. This indicates that increasing specialization within the hypercarnivorous niche may constrain subsequent morphological and ecological flexibility. During the Cenozoic, similar skull traits appeared in different carnivoran lineages, generated by similar selection pressures (e.g., toward hypercarnivory) and shared developmental pathways. These pathways were likely the proximate source of constraints on the degree of variation associated with carnivoran skull evolution and on its direction.

Biogeochemical and Ecomorphological Inferences On Prey Selection and Resource Partitioning Among Mammalian Carnivores In An Early Pleistocene Community

Abstract Biogeochemical (δ13C, δ15N, and δ18O values) and ecomorphological analyses of the early Pleistocene fauna of Venta Micena (Orce, Guadix-Baza basin, SE Spain) provide interesting clues on the physiology, dietary regimes, habitat preferences, and ecological interactions of large mammals. Such inferences are useful in deciphering aspects of paleocommunity structure and predator-prey relationships. Specifically, the hypsodonty index combined with δ13C values allows classifying the ungulates among grazers from open habitat (Equus altidens, Bison sp., Praeovibos sp., Hemitragus albus, Hippopotamus antiquus, and Mammuthus meridionalis), mixed feeders (Soergelia minor and Pseudodama sp.), and browsers from canopy areas (Stephanorhinus sp. and Praemegaceros cf. verticornis). Given that δ13C values indicate that all these herbivores fed exclusively on C3 plants, significant differences in isotopic values between perissodactyls (monogastric, hindgut fermenters) and ruminants (foregut fermenters) reflect differences in digestive efficiency. Values of δ18O indicate the dietary water source of ungulates, revealing that Pseudodama sp., Hemitragus albus, and Soergelia minor obtained a significant fraction of their metabolic water from vegetation. Carnivores show higher δ15N values than herbivores, which records the isotopic enrichment expected with an increase in trophic level. Hippopotamus antiquus and Praeovibos sp. have unexpectedly high δ15N values, suggesting that they predominantly consumed aquatic plants and lichens, respectively. Inferences on predator-prey relationships, derived from the use of linear mixing models, indicate resource partitioning among sympatric predators; saber-tooth Megantereon whitei and jaguar Panthera cf. gombaszoegensis were ambushers in closed habitat while saber-tooth Homotherium latidens and wild dog Lycaon lycaonoides were coursing predators in open plains. The giant hyena Pachycrocuta brevirostris scavenged the prey of these hypercarnivores.

A Three-Dimensional Analysis of Morphological Evolution and Locomotor Performance of the Carnivoran Forelimb

In this study, three-dimensional landmark-based methods of geometric morphometrics are used for estimating the influence of phylogeny, allometry and locomotor performance on forelimb shape in living and extinct carnivorans (Mammalia, Carnivora). The main objective is to investigate morphological convergences towards similar locomotor strategies in the shape of the major forelimb bones. Results indicate that both size and phylogeny have strong effects on the anatomy of all forelimb bones. In contrast, bone shape does not correlate in the living taxa with maximum running speed or daily movement distance, two proxies closely related to locomotor performance. A phylomorphospace approach showed that shape variation in forelimb bones mainly relates to changes in bone robustness. This indicates the presence of biomechanical constraints resulting from opposite demands for energetic efficiency in locomotion –which would require a slender forelimb– and resistance to stress –which would be satisfied by a robust forelimb–. Thus, we interpret that the need of maintaining a trade-off between both functional demands would limit shape variability in forelimb bones. Given that different situations can lead to one or another morphological solution, depending on the specific ecology of taxa, the evolution of forelimb morphology represents a remarkable “one-to-many mapping” case between anatomy and ecology.

Demythologizing Arctodus simus, the ‘Short-Faced’ Long-Legged and Predaceous Bear that Never Was

ABSTRACT In this study, we review the previous evidence on the paleobiology of the giant, ‘short-faced’ bear Arctodus simus (Mammalia: Carnivora: Ursidae) and contribute new ecomorphological inferences on the paleobiology of this enigmatic species. Craniodental variables are used in a comparative morphometric study across the families Felidae, Hyaenidae, Canidae, and Ursidae. Principal components analyses (PCAs) do not show an ecomorphological adaptation towards bonecracking or hypercarnivory in the ‘short-faced’ bear. In contrast, PCAs and discriminant analyses restricted to the craniodental data set of ursids suggest close morphological resemblance between A. simus and the extant omnivorous bears. In addition, the scaling of snout length on neurocranial length in bears indicates that the face of A. simus was not particularly short. Body mass estimates obtained from major limb bone measurements reveal that A. simus specimens of around 1000 kilograms were more common than previously suspected. Scaling relationships in extant bears of limb lengths on the least width of the femoral shaft (the variable best correlated with body mass) indicate that A. simus was not as relatively long-legged as previously thought. For these reasons, although the isotopic signature of A. simus has been interpreted as evidencing that it consumed large amounts of flesh relative to some contemporary populations of Ursus arctos, our results do not support the previous views of A. simus as a fast-running super-predator or as a specialized scavenger. In contrast, the picture that emerges from this study is one of a colossal omnivorous bear whose diet probably varied according to resource availability.

Patterns of morphological integration in the appendicular skeleton of mammalian carnivores

We investigated patterns of evolutionary integration in the appendicular skeleton of mammalian carnivores. The findings are discussed in relation to performance selection in terms of organismal function as a potential mechanism underlying integration. Interspecific shape covariation was quantified by two‐block partial least‐squares (2B‐PLS) analysis of 3D landmark data within a phylogenetic context. Specifically, we compared pairs of anatomically connected bones (within‐limbs) and pairs of both serially homologous and functional equivalent bones (between‐limbs). The statistical results of all the comparisons suggest that the carnivoran appendicular skeleton is highly integrated. Strikingly, the main shape covariation relates to bone robustness in all cases. A bootstrap test was used to compare the degree of integration between specialized cursorial taxa (i.e., those whose forelimbs are primarily involved in locomotion) and noncursorial species (i.e., those whose forelimbs are involved in more functions than their hindlimb) showed that cursors have a more integrated appendicular skeleton than noncursors. The findings demonstrate that natural selection can influence the pattern and degree of morphological integration by increasing the degree of bone shape covariation in parallel to ecological specialization.

A three-dimensional analysis of the morphological evolution and locomotor behaviour of the carnivoran hind limb

BackgroundThe shape of the appendicular bones in mammals usually reflects adaptations towards different locomotor abilities. However, other aspects such as body size and phylogeny also play an important role in shaping bone design.We used 3D landmark-based geometric morphometrics to analyse the shape of the hind limb bones (i.e., femur, tibia, and pelvic girdle bones) of living and extinct terrestrial carnivorans (Mammalia, Carnivora) to quantitatively investigate the influence of body size, phylogeny, and locomotor behaviour in shaping the morphology of these bones. We also investigated the main patterns of morphological variation within a phylogenetic context.ResultsSize and phylogeny strongly influence the shape of the hind limb bones. In contrast, adaptations towards different modes of locomotion seem to have little influence. Principal Components Analysis and the study of phylomorphospaces suggest that the main source of variation in bone shape is a gradient of slenderness-robustness.ConclusionThe shape of the hind limb bones is strongly influenced by body size and phylogeny, but not to a similar degree by locomotor behaviour. The slender-robust “morphological bipolarity” found in bone shape variability is probably related to a trade-off between maintaining energetic efficiency and withstanding resistance to stresses. The balance involved in this trade-off impedes the evolution of high phenotypic variability. In fact, both morphological extremes (slender/robust) are adaptive in different selective contexts and lead to a convergence in shape among taxa with extremely different ecologies but with similar biomechanical demands. Strikingly, this “one-to-many mapping” pattern of evolution between morphology and ecology in hind limb bones is in complete contrast to the “many-to-one mapping” pattern found in the evolution of carnivoran skull shape. The results suggest that there are more constraints in the evolution of the shape of the appendicular skeleton than in that of skull shape because of the strong biomechanical constraints imposed by terrestrial locomotion.