Modern squamates (lizards, snakes and amphisbaenians) are the world's most diverse group of tetrapods along with birds and have a long evolutionary history, with the oldest known fossils dating from the Middle Jurassic period-168 million years ago. The evolutionary origin of squamates is contentious because of several issues: (1) a fossil gap of approximately 70 million years exists between the oldest known fossils and their estimated origin; (2) limited sampling of squamates in reptile phylogenies; and (3) conflicts between morphological and molecular hypotheses regarding the origin of crown squamates. Here we shed light on these problems by using high-resolution microfocus X-ray computed tomography data from the articulated fossil reptile Megachirella wachtleri (Middle Triassic period, Italian Alps ). We also present a phylogenetic dataset, combining fossils and extant taxa, and morphological and molecular data. We analysed this dataset under different optimality criteria to assess diapsid reptile relationships and the origins of squamates. Our results re-shape the diapsid phylogeny and present evidence that M. wachtleri is the oldest known stem squamate. Megachirella is 75 million years older than the previously known oldest squamate fossils, partially filling the fossil gap in the origin of lizards, and indicates a more gradual acquisition of squamatan features in diapsid evolution than previously thought. For the first time, to our knowledge, morphological and molecular data are in agreement regarding early squamate evolution, with geckoes-and not iguanians-as the earliest crown clade squamates. Divergence time estimates using relaxed combined morphological and molecular clocks show that lepidosaurs and most other diapsids originated before the Permian/Triassic extinction event, indicating that the Triassic was a period of radiation, not origin, for several diapsid lineages.

The Carnian Pluvial Episode (Late Triassic) was a time of global environmental changes and possibly substantial coeval volcanism. The extent of the biological turnover in marine and terrestrial ecosystems is not well understood. Here, we present a meta-analysis of fossil data that suggests a substantial reduction in generic and species richness and the disappearance of 33% of marine genera. This crisis triggered major radiations. In the sea, the rise of the first scleractinian reefs and rock-forming calcareous nannofossils points to substantial changes in ocean chemistry. On land, there were major diversifications and originations of conifers, insects, dinosaurs, crocodiles, lizards, turtles, and mammals. Although there is uncertainty on the precise age of some of the recorded biological changes, these observations indicate that the Carnian Pluvial Episode was linked to a major extinction event and might have been the trigger of the spectacular radiation of many key groups that dominate modern ecosystems.

Dinosaurs diversified in two steps during the Triassic. They originated about 245 Ma, during the recovery from the Permian-Triassic mass extinction, and then remained insignificant until they exploded in diversity and ecological importance during the Late Triassic. Hitherto, this Late Triassic explosion was poorly constrained and poorly dated. Here we provide evidence that it followed the Carnian Pluvial Episode (CPE), dated to 234–232 Ma, a time when climates switched from arid to humid and back to arid again. Our evidence comes from a combined analysis of skeletal evidence and footprint occurrences, and especially from the exquisitely dated ichnofaunas of the Italian Dolomites. These provide evidence of tetrapod faunal compositions through the Carnian and Norian, and show that dinosaur footprints appear exactly at the time of the CPE. We argue then that dinosaurs diversified explosively in the mid Carnian, at a time of major climate and floral change and the extinction of key herbivores, which the dinosaurs opportunistically replaced.

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We present a holistic approach to the study of early archosauriform evolution by integrating body and track records. The ichnological record supports a Late Permian–Early Triassic radiation of archosauriforms not well documented by skeletal material, and new footprints from the Upper Permian of the southern Alps (Italy) provide evidence for a diversity not yet sampled by body fossils. The integrative study of body fossil and footprint data supports the hypothesis that archosauriforms had already undergone substantial taxonomic diversification by the Late Permian and that by the Early Triassic archosauromorphs attained a broad geographical distribution over most parts of Pangea. Analysis of body size, as deduced from track size, suggests that archosauriform average body size did not change significantly from the Late Permian to the Early Triassic. A survey of facies yielding both skeletal and track record indicate an ecological preference for inland fluvial (lacustrine) environments for early archosauromorphs. Finally, although more data is needed, Late Permian chirotheriid imprints suggest a shift from sprawling to erect posture in archosauriforms before the end-Permian mass extinction event. We highlight the importance of approaching palaeobiological questions by using all available sources of data, specifically through integrating the body and track fossil record.

The Permian–Triassic mass extinction (PTME) had an enormous impact on life in three ways: by substantially reducing diversity, by reshuffling the composition of ecosystems and by expelling life from the tropics following episodes of intense global warming. But was there really an ‘equatorial tetrapod gap', and how long did it last? Here, we consider both skeletal and footprint data, and find a more complex pattern: (i) tetrapods were distributed both at high and low latitudes during this time; (ii) there was a clear geographic disjunction through the PTME, with tetrapod distribution shifting 10–15° poleward; and (iii) there was a rapid expansion phase across the whole of Pangea following the PTME. These changes are consistent with a model of generalized migration of tetrapods to higher latitudinal, cooler regions, to escape from the superhot equatorial climate in the earliest Triassic, but the effect was shorter in time scale, and not as pronounced as had been proposed. In the recovery phase following the PTME, this episode of forced range expansion also appears to have promoted the emergence and radiation of entirely new groups, such as the archosaurs, including the dinosaurs.

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