Gabriela Sobral

The Arnstadt Formation of Saxony-Anhalt, Germany has yielded some of Germany’s most substantial finds of Late Triassic tetrapods, including the sauropodomorph Plateosaurus and the stem-turtle Proganochelys quenstedti. Here, we describe an almost complete skull of a new sphenodontian taxon from this formation (Norian, 227–208 Ma), making it the oldest known articulated sphenodontian skull from Europe and one of the oldest in the world. The material is represented by the dermal skull roof and by the complete maxilla and temporal region, as well as parts of the palate, braincase, and lower jaw. A phylogenetic assessment recovers it as a basal sphenodontian closely related to Planocephalosaurus robinsonae and to Eusphenodontia, making it the earliest-diverging sphenodontian known with an articulated skull. Its cranial anatomy is generally similar to the well-known Diphydontosaurus avonis from the Rhaetian of England, showing that this successful phenotype was already established in the clade around 10 myr earlier than assumed. An analysis of evolutionary change rates recovers high rates of evolution in basal sphenodontians, with decreasing rates throughout the evolution of the group. However, contrary to previous studies, reversals in this trend were identified, indicating additional peaks of evolutionary change. These results improve our understanding of the early sphenodontian diversity in Europe, providing critical information on evolutionary trends throughout the history of the clade and sparking renewed interest in its evolution.

The early evolution of diapsid reptiles is marked by a deep contrast between our knowledge of the origin and early evolution of archosauromorphs (crocodiles, avian and non-avian dinosaurs) to that of lepidosauromorphs (squamates (lizards, snakes) and sphenodontians (tuataras)). Whereas the former include hundreds of fossil species across various lineages during the Triassic period¹, the latter are represented by an extremely patchy early fossil record comprising only a handful of fragmentary fossils, most of which have uncertain phylogenetic affinities and are confined to Europe^1,2,3. Here we report the discovery of a three-dimensionally preserved reptile skull, assigned as Taytalura alcoberi gen. et sp. nov., from the Late Triassic epoch of Argentina that is robustly inferred phylogenetically as the earliest evolving lepidosauromorph, using various data types and optimality criteria. Micro-computed tomography scans of this skull reveal details about the origin of the lepidosaurian skull from early diapsids, suggesting that several traits traditionally associated with sphenodontians in fact originated much earlier in lepidosauromorph evolution. Taytalura suggests that the strongly evolutionarily conserved skull architecture of sphenodontians represents the plesiomorphic condition for all lepidosaurs, that stem and crown lepidosaurs were contemporaries for at least ten million years during the Triassic, and that early lepidosauromorphs had a much broader geographical distribution than has previously been thought.

The late Paleozoic temnospondyl Sclerocephalus formed an aquatic top predator in various central European lakes of the late Carboniferous and early Permian. Despite hundreds of specimens spanning a wide range of sizes, knowledge of the endocranium (braincase and palatoquadrate) remained very insufficient in Sclerocephalus and other stereospondylomorphs because even large skulls had unossified endocrania. A new specimen from a stratigraphically ancient deposit at St. Wendel in southwestern Germany is recognized as representing a new taxon, S. concordiae new species, and reveals a completely ossified endocranium. The sphenethmoid was completely ossified from the basisphenoid to the anterior ethmoid region, co-ossified with the parasphenoid, and the basipterygoid joint was fully established. The pterygoid bears a slender, S-shaped epipterygoid, which formed a robust pillar lateral to the braincase. The massive stapes was firmly sutured to the parasphenoid. In the temnospondyl endocranium, character evolution involved various changes in the epipterygoid region, which evolved distinct morphologies in each of the major clades.UUID: http://zoobank.org/5e6d2078-eacf-4467-84cf-a12efcae7c0b

The Middle Triassic was an important time in the evolutionary history of reptiles because it was during this time that many modern groups originated and initially diversified. In this context, the rich fossiliferous deposits of the Ladinian-age Erfurt Formation in southern Germany play an important role in elucidating reptilian diversity during this interval. Here we present an interesting record of a reptile with an unusual dentition. The material comprises a fragment of a left dentary. The teeth are tightly spaced and have chisel-shaped crowns, with the posterolingually extending apices abutting those of the successive teeth and forming a continuous cutting edge. Micro-computed tomography scanning of the jaw fragment revealed the teeth have expanded roots and sit on an open, shallow groove. The Meckelian canal is partially open lingually and a second canal for the inferior alveolar nerve and artery extends parallel to it. Although the preserved anatomical features of the material are insufficient for precise phylogenetic placement, the structure of the robust dentary and its teeth resembles those of some lepidosaurs and certain early Mesozoic marine diapsids. The distinctive morphology of the teeth, combined with the apparent lack of tooth wear, makes it difficult to assess their functional significance. It is interesting that extinct tetrapods with similar tooth morphologies come from strata representing freshwater and brackish-water paleoenvironments, which may serve as an additional line of evidence for future Morpho-functional assessments.

Archosauria (birds, crocodilians and their extinct relatives) form a major part of terrestrial ecosystems today, with over 10 000 living species, and came to dominate the land for most of the Mesozoic (over 150 Myr) after radiating following the Permian–Triassic extinction. The archosaur skull has been essential to this diversification, itself diversified into myriad forms. The archosauriformEuparkeria capensisfrom the Middle Triassic (Anisian) of South Africa has been of great interest since its initial description in 1913, because its anatomy shed light on the origins and early evolution of crown Archosauria and potentially approached that of the archosaur common ancestor.Euparkeriahas been widely used as an outgroup in phylogenetic analyses and when investigating patterns of trait evolution among archosaurs. Although described monographically in 1965, subsequent years have seen great advances in the understanding of early archosaurs and in imaging techniques. Here, the cranium and mandible ofEuparkeriaare fully redescribed and documented using all fossil material and computed tomographic data. Details previously unclear are fully described, including vomerine dentition, the epiptergoid, number of premaxillary teeth and palatal arrangement. A new diagnosis and cranial and braincase reconstruction is provided, and an anatomical network analysis is performed on the skull ofEuparkeriaand compared with other amniotes. The modular composition of the cranium suggests a flexible skull well adapted to feeding on agile food, but with a clear tendency towards more carnivorous behaviour, placing the taxon at the interface between ancestral diapsid and crown archosaur ecomorphology, corresponding to increases in brain size, visual sensitivity, upright locomotion and metabolism around this point in archosauriform evolution. The skull ofEuparkeriaepitomizes a major evolutionary transition, and places crown archosaur morphology in an evolutionary context.

The Middle Triassic was a time of major changes in tetrapod faunas worldwide, but the fossil record for this interval is largely obscure for terrestrial faunas. This poses a severe limitation to our understanding on the earliest stages of diversification of lineages representing some of the most diverse faunas in the world today, such as lepidosauromorphs (e.g., lizards and tuataras). Here, we report a tiny new lepidosauromorph from the Middle Triassic from Vellberg (Germany), which combines a mosaic of features from both early evolving squamates and rhynchocephalians, such as the simultaneous occurrence of a splenial bone and partial development of acrodonty. Phylogenetic analyses applying different optimality criteria, and combined morphological and molecular data, consistently recover the new taxon as a stem-lepidosauromorph, implying stem-lepidosauromorph species coinhabited areas comprising today’s central Europe at the same time as the earliest known rhynchocephalians and squamates. It further demonstrates a more complex evolutionary scenario for dental evolution in early lepidosauromorphs, with independent acquisitions of acrodonty early in their evolutionary history. The small size of most terrestrial vertebrates from Vellberg is conspicuous, contrasting to younger Triassic deposits worldwide, but comparable to Early Triassic faunas, suggesting a potential long-lasting Lilliput effect in this fauna.