Ever since Darwin, biologists have debated the relative roles of external and internal drivers of large-scale evolution. The distributions and ecology of living crocodilians are controlled by environmental factors such as temperature. Crocodilians have a rich history, including amphibious, marine and terrestrial forms spanning the past 247 Myr. It is uncertain whether their evolution has been driven by extrinsic factors, such as climate change and mass extinctions, or intrinsic factors like sexual selection and competition. Using a new phylogeny of crocodilians and their relatives, we model evolutionary rates using phylogenetic comparative methods. We find that body size evolution follows a punctuated, variable rate model of evolution, consistent with environmental drivers of evolution, with periods of stability interrupted by periods of change. Regression analyses show warmer environmental temperatures are associated with high evolutionary rates and large body sizes. We confirm that environmental factors played a significant role in the evolution of crocodiles.

Locomotor mode is an important component of an animal's ecology, relating to both habitat and substrate choice (e.g., arboreal versus terrestrial) and in the case of carnivores, to mode of predation (e.g., ambush versus pursuit). Here, we examine how the morphology of the calcaneum, the 'heel bone' in the tarsus, correlates with locomotion in extant carnivores. Other studies have confirmed the correlation of calcaneal morphology with locomotion behaviour and habitat. The robust nature of the calcaneum means that it is frequently preserved in the fossil record. Here, we employ linear measurements and 2D-geometric morphometrics on a sample of calcanea from eighty-seven extant carnivorans and demonstrate a signal of correlation between calcaneal morphology and locomotor mode that overrides phylogeny. We used this correlation to determine the locomotor mode, and hence aspects of the palaeobiology of, 47 extinct carnivorous mammal taxa, including both Carnivora and Creodonta. We found ursids (bears), clustered together, separate from the other carnivorans. Our results support greater locomotor diversity for nimravids (the extinct 'false sabertooths', usually considered to be more arboreal), than previously expected. However, there are limitations to interpretation of extinct taxa because their robust morphology is not fully captured in the range of modern carnivoran morphology.

Despite reports of sexual dimorphism in extinct taxa, such claims in non-avian dinosaurs have been rare over the last decade and have often been criticized. Since dimorphism is widespread in sexually reproducing organisms today, under-reporting in the literature might suggest either methodological shortcomings or that this diverse group exhibited highly unusual reproductive biology. Univariate significance testing, especially for bimodality, is ineffective and prone to false negatives. Species recognition and mutual sexual selection hypotheses, therefore, may not be required to explain supposed absence of sexual dimorphism across the grade (a type II error). Instead, multiple lines of evidence support sexual selection and variation of structures consistent with secondary sexual characteristics, strongly suggesting sexual dimorphism in non-avian dinosaurs. We propose a framework for studying sexual dimorphism in fossils, focusing on likely secondary sexual traits and testing against all alternate hypotheses for variation in them using multiple lines of evidence. We use effect size statistics appropriate for low sample sizes, rather than significance testing, to analyse potential divergence of growth curves in traits and constrain estimates for dimorphism magnitude. In many cases, estimates of sexual variation can be reasonably accurate, and further developments in methods to improve sex assignments and account for intrasexual variation (e.g. mixture modelling) will improve accuracy. It is better to compare estimates for the magnitude of and support for dimorphism between datasets than to dichotomously reject or fail to reject monomorphism in a single species, enabling the study of sexual selection across phylogenies and time. We defend our approach with simulated and empirical data, including dinosaur data, showing that even simple approaches can yield fairly accurate estimates of sexual variation in many cases, allowing for comparison of species with high and low support for sexual variation.

The crocodiles and their close relatives, the alligators and gharials, have a compelling evolutionary history. They are a clade of great antiquity, with their most recent common ancestor emerging within the Mesozoic. However, unlike many groups of such a great age, the crocodilians have an extensive crown-group, with around two dozen extant examples. They have a limited ecomorphology, which has varied little since their inception, and their biogeography has been shown to interact closely with climate. The biogeography of crocodilians in deep time remains an outstanding question, which is complicated further by the limitations of the fossil record. The fossil record is fundamentally incomplete yet represents the most common method used to infer biogeography of organisms. The scarcity of fossil remains makes apparent absences difficult to confirm. Preservation bias will promote fossil occurrences in areas with a high sedimentation rate, which may not be the true ecological niche for a given taxon. This study uses species distribution models of extant crocodilians to infer the ecological niche of related taxa in the Maastrichtian and Danian. Models indicate a much wider latitudinal range than is observed among extant examples, and the invasion of new ecospace following the end-Cretaceous mass extinction. In addition, we find that while temperature is of significance to crocodilian biogeography, it is precipitation that is the most influential climatic variable.

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