The development of distinct regions in the amniote vertebral column results from somite formation andHoxgene expression, with the adult morphology displaying remarkable variation among lineages. Mammalian regionalization is reportedly very conservative or even constrained, but there has been no study investigating vertebral count variation across Amniota as a whole, undermining attempts to understand the phylogenetic, ecological, and developmental factors affecting vertebral column variation. Here, we show that the mammalian (synapsid) and reptilian lineages show early in their evolutionary histories clear divergences in axial developmental plasticity, in terms of both regionalization and meristic change, with basal synapsids sharing the conserved axial configuration of crown mammals, and basal reptiles demonstrating the plasticity of extant taxa. We conducted a comprehensive survey of presacral vertebral counts across 436 recent and extinct amniote taxa. Vertebral counts were mapped onto a generalized amniote phylogeny as well as individual ingroup trees, and ancestral states were reconstructed by using squared-change parsimony. We also calculated the relationship between presacral and cervical numbers to infer the relative influence of homeotic effects and meristic changes and found no correlation between somitogenesis andHox-mediated regionalization. Although conservatism in presacral numbers characterized early synapsid lineages, in some cases reptiles and synapsids exhibit the same developmental innovations in response to similar selective pressures. Conversely, increases in body mass are not coupled with meristic or homeotic changes, but mostly occur in concert with postembryonic somatic growth. Our study highlights the importance of fossils in large-scale investigations of evolutionary developmental processes.

The multiple skeletal components of the skull originate asynchronously and their developmental schedule varies across amniotes. Here we present the embryonic ossification sequence of 134 species, covering all major groups of mammals and their close relatives. This comprehensive data set allows reconstruction of the heterochronic and modular evolution of the skull and the condition of the last common ancestor of mammals. We show that the mode of ossification (dermal or endochondral) unites bones into integrated evolutionary modules of heterochronic changes and imposes evolutionary constraints on cranial heterochrony. However, some skull-roof bones, such as the supraoccipital, exhibit evolutionary degrees of freedom in these constraints. Ossification timing of the neurocranium was considerably accelerated during the origin of mammals. Furthermore, association between developmental timing of the supraoccipital and brain size was identified among amniotes. We argue that cranial heterochrony in mammals has occurred in concert with encephalization but within a conserved modular organization.

Sea turtles (Chelonoidea) are a charismatic group of marine reptiles that occupy a range of important ecological roles. However, the diversity and evolution of their feeding anatomy remain incompletely known. Using computed tomography and classical comparative anatomy we describe the cranial anatomy in two sea turtles, the loggerhead (Caretta caretta) and Kemp’s ridley (Lepidochelys kempii), for a better understanding of sea turtle functional anatomy and morphological variation. In both taxa the temporal region of the skull is enclosed by bone and the jaw joint structure and muscle arrangement indicate that palinal jaw movement is possible. The tongue is relatively small, and the hyoid apparatus is not as conspicuous as in some freshwater aquatic turtles. We find several similarities between the muscles of C. caretta and L. kempii, but comparison with other turtles suggests only one of these characters may be derived: connection of the m. adductor mandibulae internus into the Pars intramandibularis via the Zwischensehne. The large fleshy origin of the m. adductor mandibulae externus Pars superficialis from the jugal seems to be a characteristic feature of sea turtles. In C. caretta and L. kempii the ability to suction feed does not seem to be as well developed as that found in some freshwater aquatic turtles. Instead both have skulls suited to forceful biting. This is consistent with the observation that both taxa tend to feed on relatively slow moving but sometimes armoured prey. The broad fleshy origin of the m. adductor mandibulae externus Pars superficialis may be linked to thecheek region being almost fully enclosed in bone but the relationship is complex.

The unique ability of modern turtles to retract their head and neck into the shell through a side-necked (pleurodiran) or hidden-necked (cryptodiran) motion is thought to have evolved independently in crown turtles. The anatomical changes that led to the vertebral shapes of modern turtles, however, are still poorly understood. Here we present comprehensive geometric morphometric analyses that trace turtle vertebral evolution and reconstruct disparity across phylogeny. Disparity of vertebral shape was high at the dawn of turtle evolution and decreased after the modern groups evolved, reflecting a stabilization of morphotypes that correspond to the two retraction modes. Stem turtles, which had a very simple mode of retraction, the lateral head tuck, show increasing flexibility of the neck through evolution towards a pleurodiran-like morphotype. The latter was the precondition for evolving pleurodiran and cryptodiran vertebrae. There is no correlation between the construction of formed articulations in the cervical centra and neck mobility. An increasing mobility between vertebrae, associated with changes in vertebral shape, resulted in a more advanced ability to retract the neck. In this regard, we hypothesize that the lateral tucking retraction of stem turtles was not only the precondition for pleurodiran but also of cryptodiran retraction. For the former, a kink in the middle third of the neck needed to be acquired, whereas for the latter modification was necessary between the eighth cervical vertebra and first thoracic vertebra. Our paper highlights the utility of 3D shape data, analyzed in a phylogenetic framework, to examine the magnitude and mode of evolutionary modifications to vertebral morphology. By reconstructing and visualizing ancestral anatomical shapes we provide insight into the anatomical features underlying neck retraction mode, which is a salient component of extant turtle classification.