We describe a new small-bodied coelophysoid theropod dinosaur,Pendraig milneraegen. et sp. nov, from the Late Triassic fissure fill deposits of Pant-y-ffynnon in southern Wales. The species is represented by the holotype, consisting of an articulated pelvic girdle, sacrum and posterior dorsal vertebrae, and an associated left femur, and by two referred specimens, comprising an isolated dorsal vertebra and a partial left ischium. Our phylogenetic analysis recoversP. milneraeas a non-coelophysid coelophysoid theropod, representing the first-named unambiguous theropod from the Triassic of the UK. Recently, it has been suggested that Pant-y-ffynnon and other nearby Late Triassic to Early Jurassic fissure fill faunas might have been subjected to insular dwarfism. To test this hypothesis forP. milnerae, we performed an ancestral state reconstruction analysis of body size in early neotheropods. Although our results indicate that a reduced body size is autapomorphic forP. milnerae, some other coelophysoid taxa show a similar size reduction, and there is, therefore, ambiguous evidence to indicate that this species was subjected to dwarfism. Our analyses further indicate that, in contrast with averostran-line neotheropods, which increased in body size during the Triassic, coelophysoids underwent a small body size decrease early in their evolution.

The historical clade “Protorosauria” represents an important group of archosauromorph reptiles that had a wide geographic distribution between the Late Permian and Late Triassic. “Protorosaurs” are characterized by their long necks, which are epitomized in the genusTanystropheusand inDinocephalosaurus orientalis. Recent phylogenetic analyses have indicated that “Protorosauria” is a polyphyletic clade, but the exact relationships of the various “protorosaur” taxa within the archosauromorph lineage is currently uncertain. Several taxa, although represented by relatively complete material, have previously not been assessed phylogenetically. We present a new phylogenetic hypothesis that comprises a wide range of archosauromorphs, including the most exhaustive sample of “protorosaurs” to date and several “protorosaur” taxa from the eastern Tethys margin that have not been included in any previous analysis. The polyphyly of “Protorosauria” is confirmed and therefore we suggest the usage of this term should be abandoned. Tanystropheidae is recovered as a monophyletic group and the Chinese taxaDinocephalosaurus orientalisandPectodens zhenyuensisform a new archosauromorph clade, Dinocephalosauridae, which is closely related to Tanystropheidae. The well-known crocopod and former “protorosaur”Prolacerta broomiis considerably less closely related to Archosauriformes than was previously considered.

A new specimen of the protorosaur Macrocnemus fuyuanensis, from the Zhuganpo Member of the Falang Formation at Fuyuan (Yunnan Province, southwestern China), is described and compared with Macrocnemus bassanii. The new specimen is the first in the genus Macrocnemus that clearly shows details of the preorbital region of the skull. A large, plate-like lacrimal is located in front of the tall, columnar prefrontal that defines the anterior margin of the orbit. A longitudinally oriented nasal groove extends along the anterior two-thirds of the snout, accommodating the external naris at its anterior part. A similar preorbital depression has previously been described for Dinocephalosaurus and it is also reported here for the first time in Tanystropheus. The new specimen confirms the status of Macrocnemus fuyuanensis as a species distinct from Macrocnemus bassanii on the basis of a humerus that significantly exceeds the radius in length. The occurrence of both Macrocnemus and Tanystropheus in southwestern China further underscores the close faunal affinities of the eastern and western Tethyan realms during the Middle and early Late Triassic.

The postcranial morphology of the extremely long-neckedTanystropheus hydroidesis well-known, but observations of skull morphology were previously limited due to compression of the known specimens. Here we provide a detailed description of the skull of PIMUZ T 2790, including a partial endocast and endosseous labyrinth, based on synchrotron microtomographic data, and compare its morphology to that of other early Archosauromorpha. In many features, such as the wide and flattened snout and the configuration of the temporal and palatal regions,Tanystropheus hydroidesdiffers strongly from other early archosauromorphs. The braincase possesses a combination of derived archosaur traits, such as the presence of a laterosphenoid and the ossification of the lateral wall of the braincase, but also differs from archosauriforms in the morphology of the ventral ramus of the opisthotic, the horizontal orientation of the parabasisphenoid, and the absence of a clearly defined crista prootica.Tanystropheus hydroideswas a ram-feeder that likely caught its prey through a laterally directed snapping bite. Although the cranial morphology of other archosauromorph lineages is relatively well-represented, the skulls of most tanystropheid taxa remain poorly understood due to compressed and often fragmentary specimens. The recent descriptions of the skulls ofMacrocnemus bassaniiand nowTanystropheus hydroidesreveal a large cranial disparity in the clade, reflecting wide ecological diversity, and highlighting the importance of non-archosauriform Archosauromorpha to both terrestrial and aquatic ecosystems during the Triassic.