BackgroundThe dentitions of extinct organisms can provide pivotal information regarding their phylogenetic position, as well as paleobiology, diet, development, and growth. Extant birds are edentulous (toothless), but their closest relatives among stem birds, the Cretaceous Hesperornithiformes and Ichthyornithiformes, retained teeth. Despite their significant phylogenetic position immediately outside the avian crown group, the dentitions of these taxa have never been studied in detail. To obtain new insight into the biology of these ‘last’ toothed birds, we use cutting-edge visualisation techniques to describe their dentitions at unprecedented levels of detail, in particular propagation phase contrast x-ray synchrotron microtomography at high-resolution.ResultsAmong other characteristics of tooth shape, growth, attachment, implantation, replacement, and dental tissue microstructures, revealed by these analyses, we find that tooth morphology and ornamentation differ greatly between the Hesperornithiformes and Ichthyornithiformes. We also highlight the first Old World, and youngest record of the major Mesozoic clade Ichthyornithiformes. Both taxa exhibit extremely thin and simple enamel. The extension rate of Hesperornis tooth dentine appears relatively high compared to non-avian dinosaurs. Root attachment is found for the first time to be fully thecodont via gomphosis in both taxa, but in Hesperornis secondary evolution led to teeth implantation in a groove, at least locally without a periodontal ligament. Dental replacement is shown to be lingual via a resorption pit in the root, in both taxa.ConclusionsOur results allow comparison with other archosaurs and also mammals, with implications regarding dental character evolution across amniotes. Some dental features of the ‘last’ toothed birds can be interpreted as functional adaptations related to diet and mode of predation, while others appear to be products of their peculiar phylogenetic heritage. The autapomorphic Hesperornis groove might have favoured firmer root attachment. These observations highlight complexity in the evolutionary history of tooth reduction in the avian lineage and also clarify alleged avian dental characteristics in the frame of a long-standing debate on bird origins. Finally, new hypotheses emerge that will possibly be tested by further analyses of avian teeth, for instance regarding dental replacement rates, or simplification and thinning of enamel throughout the course of early avian evolution.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-016-0753-6) contains supplementary material, which is available to authorized users.

A major goal of evolutionary studies is to better understand how complex morphologies are related to the different functions and behaviours in which they are involved. For example, during locomotion and hunting behaviour, the head and the eyes have to stay at an appropriate level in order to reliably judge distance as well as to provide postural information. The morphology and orientation of the orbits and cranial base will have an impact on eye orientation. Consequently, variation in orbital and cranial base morphology is expected to be correlated with aspects of an animal's lifestyle. In this study, we investigate whether the shape of the skull evolves in response to the functional demands imposed by ecology and behaviour using geometric morphometric methods. We test if locomotor habitats, diet, and activity pattern influence the shape of the skull in musteloid carnivorans using (M)ANOVAs and phylogenetic (M)ANOVAs, and explore the functional correlates of morphological features in relation to locomotor habitats, diet, and activity pattern. Our results show that phylogeny, locomotion and, diet strongly influence the shape of the skull, whereas the activity pattern seems to have a weakest influence. We also show that the locomotor environment is highly integrated with foraging and feeding, which can lead to similar selective pressures and drive the evolution of skull shape in the same direction. Finally, we show similar responses to functional demands in musteloids, a super family of close related species, as are typically observed across all mammals suggesting the pervasiveness of these functional demands.

Inner vertebral architecture is poorly known, except in human and laboratory animals. In order to document this topic at a broad comparative level, a 2D-histomorphometric study of vertebral centra was conducted in a sample of 98 therian mammal species, spanning most of the size range and representing the main locomotor adaptations known in therian taxa. Eleven variables relative to the development and geometry of trabecular networks were extracted from CT scan mid-sagittal sections. Phylogeny-informed statistical tests were used to reveal the respective influences of phylogeny, size, and locomotion adaptations on mammalian vertebral structure. The use of random taxon reshuffling and squared change parsimony reveals that 9 of the 11 characteristics (the two exceptions are total sectional area and structural polarization) contain a phylogenetic signal. Linear discriminant analyses suggest that the sampled taxa can be arranged into three categories with respect to locomotion mode: a) terrestrial + flying + digging + amphibious forms, b) coastal oscillatory aquatic taxa, and c) pelagic oscillatory aquatic forms represented by oceanic cetaceans. Pairwise comparison tests and linear regressions show that, when specific size increases, the length of trabecular network (Tt.Tb.Le), as well as trabecular proliferation in total sections (Pr.Tb.Tt), increase with positive allometry. This process occurs in all locomotion categories but is particularly pronounced in pelagic oscillators. Conversely, mean trabecular width has a lesser increase with size in pelagic oscillators. Trabecular orientation is not influenced by size. All tests were corrected for multiple testing. By using six structural variables or indices, locomotion mode can be predicted with a 97.4% success rate for terrestrial forms, 66.7% for coastal oscillatory, and 81.3% for pelagic oscillatory. The possible functional meaning of these results and their potential use for paleobiological inference of locomotion in extinct taxa are discussed.

Despite great interest and decades of research, the musculoskeletal relationships of the masticatory system in primates are still not fully understood. However, without a clear understanding of the interplay between muscles and bones it remains difficult to understand the functional significance of morphological traits of the skeleton. Here, we aim to study the impacts of the masticatory muscles on the shape of the cranium and the mandible as well as their co-variation in strepsirrhine primates. To do so, we use 3D geometric morphometric approaches to assess the shape of each bone of the skull of 20 species for which muscle data are available in the literature. Impacts of the masticatory muscles on the skull shape were assessed using non-phylogenetic regressions and phylogenetic regressions whereas co-variations were assessed using two-blocks partial least square (2B-PLS) and phylogenetic 2B-PLS. Our results show that there is a phylogenetic signal for skull shape and masticatory muscles. They also show that there is a significant impact of the masticatory muscles on cranial shape but not as much as on the mandible. The co-variations are also stronger between the masticatory muscles and cranial shape even when taking into account phylogeny. Interestingly, the results of co-variation between the masticatory muscles and mandibular shape show a more complex pattern in two different directions to get strong muscles associated with mandibular shape: a folivore way (with the bamboo lemurs and sifakas) and a hard-object eater one (with the aye-aye). Anat Rec, 301:291-310, 2018. © 2018 Wiley Periodicals, Inc.

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