Mammals, birds, and squamates (lizards, snakes, and relatives) are key living vertebrates, and thus understanding their evolution underpins important questions in biodiversity science. Whereas the origins of mammals and birds are relatively well understood, the roots of squamates have been obscure. Here, we report a modern-type lizard from the Late Triassic of England [202 million years (Ma)], comprising a partial skeleton, skull, and mandibles. It displays at least 15 unique squamate traits and further shares unidentatan and anguimorph apomorphies. The new discovery fixes the origin of crown Squamata as much older than had been thought, and the revised dating shows substantial diversification of modern-type squamates following the Carnian Pluvial Episode, 232 Ma ago.

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Clevosaurus was an early sphenodontian, known from many specimens and species from the Late Triassic and Early Jurassic. Here we describe and compare the cranial morphology of Clevosaurus hudsoni, the type species, and Clevosaurus cambrica, both from the UK, based on digital reconstructions from segmented CT scan data of two skulls. C. hudsoni has been described in some detail, but the CT data reveal new information on cranial anatomy, and C. cambrica is a newly described species for which the CT scans enable us to identify many previously undescribed bones, re-identify bones whose identity had previously been uncertain, and refine certain cranial elements. The near complete preservation of the left side of the skull of C. cambrica, and the preservation of much of the left side of the skull of C. hudsoni, provided us with the opportunity to fully reconstruct both species' skulls in three dimensions. For the first time, coronoid bones are identified in Clevosaurus species from the British Isles. These two species show the diversity of morphology in the genus Clevosaurus. We provide evidence for two morphotypes of C. hudsoni based on study of the dentary dentition of the syntypes and other attributed specimens.

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Rhynchocephalians were a successful, globally distributed group of diapsid reptiles that thrived in the Mesozoic. Multiple species of Clevosaurus existed worldwide in the Late Triassic and Early Jurassic, characterized by shearing bladelike teeth perhaps functionally analogous to the carnassial teeth of mammals. Morphometric analysis shows that the dentary morphospace of clevosaurs differs significantly from that of other rhynchocephalians. Five Clevosaurus species occupied islands in the Bristol Channel archipelago of the UK, but generally not those occupied by mammaliaforms, suggesting dietary character displacement. Identifying the diet of such ancient, small tetrapods has been difficult. To identify the nature of their feeding mechanics and ecology, we apply finite element analysis to two near complete three-dimensional skulls of the species Clevosaurus hudsoni and Clevosaurus cambrica to estimate bite force, resistance to bending and torsion and the distribution of stresses in the jaws during biting. Both species had bite forces and tooth pressures sufficient to break apart chitin indicating that, like early Mesozoic mammaliaforms, clevosaurs could feed on tough-shelled beetles and possibly small vertebrates. In addition, the mechanical advantage of the jaws falls within the range of early mammaliaforms, so though we cannot demonstrate niche partitioning between members of the two clades, it raises the prospect that they may have been functionally similar.