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Abstract Some of the earliest members of the archosaur-lineage (i.e., non-archosauriform archosauromorphs) are characterised by an extremely elongated neck. Recent fossil discoveries from the Guanling Formation (Middle Triassic) of southern China have revealed a dramatic increase in the known ecomorphological diversity of these extremely long-necked archosauromorphs, including the fully marine and viviparous Dinocephalosaurus orientalis. These recent discoveries merit a reinvestigation of enigmatic Triassic diapsid fossils from contemporaneous European deposits housed in historical collections. Here, we provide a redescription of Trachelosaurus fischeri, represented by a single, disarticulated specimen first described in 1918. Due to its unique morphology, which includes short, bifurcating cervical ribs, and a high presacral vertebral count, this taxon has been referred to either as a “protorosaurian” archosauromorph or a sauropterygian. Our revision clearly shows that Trachelosaurus represents the first unambiguous Dinocephalosaurus-like archosauromorph known from outside the Guanling Formation. Our finding has important systematic implications. Trachelosauridae Abel, 1919 represents the senior synonym for the recently identified Dinocephalosauridae Spiekman, Fraser and Scheyer, 2021. Based on our phylogenetic analyses, which employ two extensive datasets, we also corroborate previous findings that tanystropheids and trachelosaurids represent two families within a larger monophyletic group among non-crocopodan archosauromorphs, which is here named Tanysauria (clade nov.). Trachelosauridae is minimally composed of Trachelosaurus fischeri, Dinocephalosaurus orientalis, Pectodens zhenyuensis, and Austronaga minuta, but one of our analyses also found a probably taxonomically broader clade that may also include Gracilicollum latens and Fuyuansaurus acutirostris. Trachelosaurus fischeri considerably expands the known spatial and temporal range of Trachelosauridae to the earliest Anisian and the Central European Basin. Our findings add to the growing evidence for the presence of a diverse group of fully marine reptiles during the Middle Triassic among Tanysauria. These trachelosaurids possess flipper-like limbs, high vertebral counts, and elongate necks, thus superficially resembling long-necked Jurassic and Cretaceous plesiosaurs in some regards.

Some of the earliest members of the archosaur-lineage (i.e., non-archosauriform archosauromorphs) are characterised by an extremely elongated neck. Recent fossil discoveries from the Guanling Formation (Middle Triassic) of southern China have revealed a dramatic increase in the known ecomorphological diversity of these extremely long-necked archosauromorphs, including the fully marine and viviparous Dinocephalosaurus orientalis. These recent discoveries merit a reinvestigation of enigmatic Triassic diapsid fossils from contemporaneous European deposits housed in historical collections. Here, we provide a redescription of Trachelosaurus fischeri, represented by a single, disarticulated specimen first described in 1918. Due to its unique morphology, which includes short, bifurcating cervical ribs, and a high presacral vertebral count, this taxon has been referred to either as a "protorosaurian" archosauromorph or a sauropterygian. Our revision clearly shows that Trachelosaurus represents the first unambiguous Dinocephalosaurus-like archosauromorph known from outside the Guanling Formation. Our finding has important systematic implications. Trachelosauridae Abel, 1919 represents the senior synonym for the recently identified Dinocephalosauridae Spiekman, Fraser and Scheyer, 2021. Based on our phylogenetic analyses, which employ two extensive datasets, we also corroborate previous findings that tanystropheids and trachelosaurids represent two families within a larger monophyletic group among non-crocopodan archosauromorphs, which is here named Tanysauria (clade nov.). Trachelosauridae is minimally composed of Trachelosaurus fischeri, Dinocephalosaurus orientalis, Pectodens zhenyuensis, and Austronaga minuta, but one of our analyses also found a probably taxonomically broader clade that may also include Gracilicollum latens and Fuyuansaurus acutirostris. Trachelosaurus fischeri considerably expands the known spatial and temporal range of Trachelosauridae to the earliest Anisian and the Central European Basin. Our findings add to the growing evidence for the presence of a diverse group of fully marine reptiles during the Middle Triassic among Tanysauria. These trachelosaurids possess flipper-like limbs, high vertebral counts, and elongate necks, thus superficially resembling long-necked Jurassic and Cretaceous plesiosaurs in some regards. [ABSTRACT FROM AUTHOR]

The Triassic radiation of vertebrates saw the emergence of the modern vertebrate groups, as well as numerous extinct animals exhibiting conspicuous, unique anatomical characteristics. Among these, members of Tanystropheidae (Reptilia: Archosauromorpha) displayed cervical vertebral elongation to an extent unparalleled in any other vertebrate. Tanystropheids were exceptionally ecologically diverse and had a wide spatial and temporal distribution. This may have been related to their neck anatomy, yet its evolution and functional properties remain poorly understood. We used geometric morphometrics to capture the intraspecific variation between the vertebrae comprising the cervical column among early archosauromorphs, to trace the evolutionary history of neck elongation in these animals. Our results show that the cervical series of these reptiles can be divided into modules corresponding to those of extant animals. Tanystropheids achieved neck elongation through somite elongation and a shift between cervical and thoracic regions, without presacral vertebrae count increase—contrary to crown archosaurs. This suggests a peculiar developmental constraint that strongly affected the evolution of tanystropheids. The data obtained just at the base of the archosauromorph phylogenetic tree are crucial for further studies on the modularity of vertebral columns of not only Triassic reptile groups but extant and other extinct animals as well.

Prolacerta broomi is one of the most important of fossil reptiles. First considered as one of the earliest members of squamates, this basal archosauromorph has been used as a model for diapsid morphological evolution ever since its discovery, playing a pivotal role in hypotheses on the origin of diapsid reptiles. The holotype of Prolacerta broomi (UCMZ 2003.41R) is known from a mostly complete skull, but the original description is limited to the superficial features of the skull roof and palate. Since then, many other specimens of Prolacerta broomi have been recovered that potentially account for this limited access to anatomical information, but it remains unclear whether these aspects correspond well to the known material of the holotype. Here, the skull morphology of the holotype of Prolacerta broomi is revisited through the use of µCT scans. The identifications of some cranial elements have been corrected, such as the left prefrontal and lacrimal, and several new elements are revealed, including the epi- and ectopterygoids, prearticular, coronoid, and braincase bones. The orbitonasal region is described in detail and significantly shows a contribution of the lacrimal to the dorsal alveolar canal. Finally, the addition of the holotype as an independent OTU in recently published analyses indicate conflicts with the current knowledge on Prolacerta broomi phylogenetic affinity and taxonomy. First, it points to potential taxonomic inconsistency since the holotype does not form a monophyletic group with other Prolacerta broomi OTUs in any of the analyses and, second, it suggests a more basal position for the holotype than that recovered in some studies, more basal than rhynchosaurs and close to the origin of Crocopoda. Together, these findings indicate areas of future research interest in the study of early evolving archosauromorphs.

Proterosuchidae represents the oldest substantial diversification of Archosauromorpha and plays a key role in understanding the biotic recovery after the end-Permian mass extinction. Proterosuchidae was long treated as a wastebasket taxon, but recent revisions have reduced its taxonomic content to five valid species from the latest Permian of Russia and the earliest Triassic (Induan) of South Africa and China. In addition to these occurrences, several isolated proterosuchid bones have been reported from the Induan Panchet Formation of India for over 150 years. Following the re-study of historical specimens and newly collected material from this unit, we erect the new proterosuchid species Samsarasuchus pamelae, which is represented by most of the presacral vertebral column. We also describe cf. proterosuchid and proterosuchid cranial, girdle and limb bones that are not referred to Samsarasuchus pamelae. Phylogenetic analyses recovered Samsarasuchus pamelae within the new proterosuchid clade Chasmatosuchinae. The taxonomic diversity of Proterosuchidae is substantially expanded here, with at least 11 nominal species and several currently unnamed specimens, and a biogeographical range encompassing present-day South Africa, China, Russia, India, Brazil, Uruguay and Australia. This indicates a broader taxonomic, phylogenetic and biogeographic diversification of Proterosuchidae than previously thought in the aftermath of the end-Permian mass extinction.

Prolacerta broomi is one of the most important of fossil reptiles. First considered as one of the earliest members of squamates, this basal archosauromorph has been used as a model for diapsid morphological evolution ever since its discovery, playing a pivotal role in hypotheses on the origin of diapsid reptiles. The holotype of Prolacerta broomi (UCMZ 2003.41R) is known from a mostly complete skull, but the original description is limited to the superficial features of the skull roof and palate. Since then, many other specimens of Prolacerta broomi have been recovered that potentially account for this limited access to anatomical information, but it remains unclear whether these aspects correspond well to the known material of the holotype. Here, the skull morphology of the holotype of Prolacerta broomi is revisited through the use of µCT scans. The identifications of some cranial elements have been corrected, such as the left prefrontal and lacrimal, and several new elements are revealed, including the epi- and ectopterygoids, prearticular, coronoid, and braincase bones. The orbitonasal region is described in detail and significantly shows a contribution of the lacrimal to the dorsal alveolar canal. Finally, the addition of the holotype as an independent OTU in recently published analyses indicate conflicts with the current knowledge on Prolacerta broomi phylogenetic affinity and taxonomy. First, it points to potential taxonomic inconsistency since the holotype does not form a monophyletic group with other Prolacerta broomi OTUs in any of the analyses and, second, it suggests a more basal position for the holotype than that recovered in some studies, more basal than rhynchosaurs and close to the origin of Crocopoda. Together, these findings indicate areas of future research interest in the study of early evolving archosauromorphs. [ABSTRACT FROM AUTHOR]

The Upper Triassic Dockum Group of Garza County, Texas (lower, middle, and upper Cooper Canyon Formation) captures the radiation of Triassic non-marine tetrapods by preserving a variety of Late Triassic taxa from the southwestern United States. Our understanding of the vertebrate assemblage from these strata largely comes from a single site, the Post Quarry (lower Cooper Canyon Formation), with previous research documenting a variety of temnospondyls, sphenodontians, non-archosauriform archosauromorphs, and archosauriforms including a phytosaur, three species of aetosaurs, a poposauroid, a rauisuchid, a crocodylomorph, and several dinosauromorphs. To more completely reconstruct the vertebrate assemblage of the Dockum Group of Garza County we use an apomorphy-based approach to identify morphologically similar disarticulated and fragmentary elements from a variety of localities that span the entire Cooper Canyon Formation (Norian-Rhaetian), allowing assignments from the large clade level to the species level. Many skeletal elements are incomplete yet diagnostic and are assigned to the least inclusive clade if discrete character states do not allow for an unambiguous species-level identification. We identify new specimens referable to numerous clades including Tanystropheidae, Allokotosauria + Prolacerta + Archosauriformes, Vancleavea + Litorosuchus, Phytosauria, Paracrocodylomorpha, Dinosauriformes, and Saurischia, in addition to additional species identifications of the aetosaur Scutarx deltatlyus, and the dinosauromorph Dromomeron gregorii. Our study of this material demonstrates the utility of an apomorphy-based approach in making testable and repeatable observations for identifying small, isolated fragmentary fossil tetrapod material to reconstruct a more accurate faunal hypothesis for a portion of the Late Triassic of Texas. Previous claims of the earliest dinosaurs from near the base of the Dockum Group do not pass the apomorphy-based identification test, and the question of whether the oldest known North American dinosaurs are present in the Chinle Formation or Dockum Group can be resolved by utilizing vertebrate biostratigraphic correlation. Our revision of these fossil assemblages supports the hypothesis that early diapsids, early archosauromorphs, and non-dinosaurian dinosauromorphs were more common, diverse, and widespread in low latitudes during this time than previously thought.

Little is known about the large‐scale evolutionary patterns of skull size relative to body size, and the possible drivers behind these patterns, in Archosauromorpha. For example, the large skulls of erythrosuchids, a group of non‐archosaurian archosauromorphs from the Early and Middle Triassic, and of theropod dinosaurs are regarded as convergent adaptations for hypercarnivory. However, few investigations have explicitly tested whether erythrosuchid and theropod skulls are indeed disproportionately large for their body size, and whether this trend is driven by hypercarnivory. Here, we investigate archosauromorph relative skull size evolution, examining the scaling relationships between skull and body size of Palaeozoic and Mesozoic archosauromorphs using a robust phylogenetic framework and assessing the influence of potential drivers, such as taxonomy, diet, locomotory mode and inhabited biotope. Our results show that archosauromorph relative skull sizes are largely determined by phylogeny and that the other drivers have much weaker levels of influence. We find negative allometric scaling of skull size with respect to body size when all studied archosauromorphs are analysed. Within specific groups, skull size scales with positive allometry in non‐archosaurian archosauromorphs and, interestingly, scales isometrically in theropods. Ancestral reconstructions of skull–femur size ratio reveal a disproportionately large skull at the base of Erythrosuchidae and proportionately sized skulls at the bases of Theropoda, Carnosauria and Tyrannosauroidea. Relative skull sizes of erythrosuchids and theropods are therefore distinct from each other, indicating that disproportionately large skulls are not a prerequisite for hypercarnivory in archosauromorphs, and that erythrosuchids exhibit a bauplan unique among terrestrial Mesozoic carnivores. [ABSTRACT FROM AUTHOR]

The Chañares Formation exhibits one of the most important archosauriform records of early Carnian ecosystems. Here we present new data on the palaeohistology of Chañares archosauriforms and provide new insights into their paleobiology, as well as possible phylogenetically informative traits. Bone microstructure of Lagerpeton chanarensis and Tropidosuchus romeri is dominated by fibro-lamellar tissue and dense vascularization. On the other hand, Chanaresuchus bonapartei is more densely vascularized, but with cyclical growth characterized by alternate fibro-lamellar, parallel-fibered and lamellar-zonal tissues. Dense vascularization and fibro-lamellar tissue imply fast growth and high metabolic rates for all these taxa. These histological traits may be tentatively interpreted as a possible adaptative advantage in front of Chañares Formation environmental conditions.

Adaptive radiations have played a major role in generating modern and deep-time biodiversity. The Triassic radiation of the Archosauromorpha was one of the most spectacular vertebrate radiations, giving rise to many highly ecomorphologically varied lineages—including the dinosaurs, pterosaurs, and stem-crocodylians—that dominated the larger-bodied land fauna for the following 150 Ma, and ultimately gave rise to today’s > 10,000 species of birds and crocodylians. This radiation provides an outstanding testbed for hypotheses relating to adaptive radiations more broadly. Recent studies have started to characterize the tempo and mode of the archosauromorph early adaptive radiation, indicating very high initial rates of evolution, non-competitive niche-filling processes, and previously unrecognized morphological disparity even among non-crown taxa. However, these analyses rested primarily either on discrete characters or on geometric morphometrics of the cranium only, or even failed to fully include phylogenetic information. Here we expand previous 2D geometric morphometric cranial datasets to include new taxa and reconstructions, and create an analogous dataset of the pelvis, thereby allowing comparison of anatomical regions and the transition from “sprawling” to “upright” posture to be examined. We estimated morphological disparity and evolutionary rates through time. All sampled clades showed a delayed disparity peak for sum of variances and average nearest neighbor distances in both the cranium and pelvis, with disparity likely not saturated by the end of the studied time span (Late Jurassic); this contrasts with smaller radiations, but lends weight to similar results for large, ecomorphologically-varied groups. We find lower variations in pelvic than cranial disparity among Triassic-Jurassic archosaurs, which may be related to greater morphofunctional constraints on the pelvis. Contrasting with some previous work, but also confirming some previous findings during adaptive radiations, we find relatively widespread evidence of correlation between sampled diversity and disparity, especially at the largest phylogenetic scales and using average displacement rather than sum of variances as disparity metric; this also demonstrates the importance of comparing disparity metrics, and the importance of phylogenetic scale. Stem and crown archosauromorphs show a morphological diversification of both the cranium and pelvis with higher initial rates (Permian–Middle Triassic and at the base of major clades) followed by lower rates once diversification into niches has occurred (Late Triassic–Jurassic), indicating an “early burst” pattern sensu lato. Our results provide a more detailed and comprehensive picture of the early archosauromorph radiation and have significant bearing on the understanding of deep-time adaptive radiations more broadly, indicating widespread patterns of delayed disparity peaks, initial correlation of diversity and disparity, and evolutionary early bursts.

The radiation of archosauromorph reptiles in the Triassic Period produced an unprecedented collection of diverse and disparate forms with a mix of varied ecologies and body sizes. Some of these forms were completely unique to the Triassic, whereas others were converged on by later members of Archosauromorpha. One of the most striking examples of this is with Triopticus primus, the early dome‐headed form later mimicked by pachycephalosaurid dinosaurs. Here we fully describe the cranial anatomy of Triopticus primus, but also recognize a second dome‐headed form from a Upper Triassic deposit in present‐day India. The new taxon, Kranosaura kuttyi gen. et sp. nov., is likely the sister taxon of Triopticus primus based on the presence of a greatly expanded skull roof with a deep dorsal opening (possibly the pineal opening) through the dome, similar cranial sculpturing, and a skull table that is expanded more posterior than the posterior extent of the basioccipital. However, the dome of Kranosaura kuttyi gen. et sp. nov. extends anterodorsally, unlike that of any other archosauromorph. Histological sections and computed tomographic reconstructions through the skull of Kranosaura kuttyi gen. et sp. nov. further reveal the uniqueness of the dome of these early archosauromorphs. Moreover, our integrated analysis further demonstrates that there are many ways to create a dome in Amniota. The presence of 'dome‐headed' archosauromorphs at two localities on the western and eastern portions of Pangea suggests that these archosauromorphs were widespread and are likely part of more assemblages than currently recognized. [ABSTRACT FROM AUTHOR]

Tanystropheidae is a clade of early archosauromorphs with a reported distribution ranging from the Early to the Late Triassic of Asia, Europe, and North America. Although some specimens with possible tanystropheid affinities from the Lower Triassic beds of Brazil have been previously attributed to “Protorosauria”, little is known about the tanystropheid record in Gondwana. Here, two new and one previously reported specimen from the Sanga do Cabral Formation (Induan–Olenekian) of Brazil are described and interpreted as ?Tanystropheidae. These records, together with other tetrapods previously reported for the Sanga do Cabral Formation, increase the knowledge of the biotic diversification during the beginning of the Triassic. This contribution reinforces that the archosauromorph diversification occurred shortly after the Permo-Triassic extinction, making the Sanga do Cabral Formation an important unit for the study of early Mesozoic faunas.

Two isolated teeth, a dorsal vertebra, fragments of a humerus and femur, a fragmentary pubic “boot” and part of an ischium shaft, identified here as belonging to a large predatory archosaur were discovered in the Upper Triassic site at Marciszów near Zawiercie (southern Poland). Comparisons of the new fossils from Marciszów with the dorsal vertebrae, pubic “boot”, ischium and femur of the theropod-like Smok wawelski from Lisowice (Silesia) reveal that the two taxa are very similar. Nevertheless, due to the lack of more diagnostic elements (e.g., braincase or cranial elements), we prefer to consider all described specimens from Marciszów as Smok sp. Smok sp. shares a low mound-like, anterior trochanter with trochanteric shelf on the femur, a massive pubic “boot” with a distinct depression (= bevelled area), and a transversely lenticular ischium shaft in cross-section with S. wawelski. Some observed characters of the dorsal vertebra (e.g., lack of some lamina, shape and position of zygapophyses), however, are different from S. wawelski and may also suggest that the new findings represent a second species of the genus in the Upper Triassic of Poland. The discovery of Smok sp. at Marciszów is significant because it is the second example of the co-occurrence of this genus with: (i) bones of a large dicynodont; and (ii) record of gnawed tetrapod bones. The discovery of Smok sp. and the lack of significant morphologic divergence from S. wawelski suggest that this taxon is the only large-bodied predator currently known from the Upper Triassic of Poland. This new evidence expands the record of the genus and contributes, in some measure, to our knowledge of the stratigraphical distribution of large predatory archosaurs from the Polish Upper Triassic bone-bearing levels.

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 genus Tanystropheus and in Dinocephalosaurus 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 taxa Dinocephalosaurus orientalis and Pectodens zhenyuensis form a new archosauromorph clade, Dinocephalosauridae, which is closely related to Tanystropheidae. The well-known crocopod and former “protorosaur” Prolacerta broomi is considerably less closely related to Archosauriformes than was previously considered.

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 genus Tanystropheus and in Dinocephalosaurus 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 taxa Dinocephalosaurus orientalis and Pectodens zhenyuensis form a new archosauromorph clade, Dinocephalosauridae, which is closely related to Tanystropheidae. The well-known crocopod and former "protorosaur" Prolacerta broomi is considerably less closely related to Archosauriformes than was previously considered. [ABSTRACT FROM AUTHOR]

The Triassic radiation of vertebrates saw the emergence of the modern vertebrate groups, as well as numerous extinct animals exhibiting conspicuous, unique anatomical characteristics. Among these, members of Tanystropheidae (Reptilia: Archosauromorpha) displayed cervical vertebral elongation to an extent unparalleled in any other vertebrate. Tanystropheids were exceptionally ecologically diverse and had a wide spatial and temporal distribution. This may have been related to their neck anatomy, yet its evolution and functional properties remain poorly understood. We used geometric morphometrics to capture the intraspecific variation between the vertebrae comprising the cervical column among early archosauromorphs, to trace the evolutionary history of neck elongation in these animals. Our results show that the cervical series of these reptiles can be divided into modules corresponding to those of extant animals. Tanystropheids achieved neck elongation through somite elongation and a shift between cervical and thoracic regions, without presacral vertebrae count increase-contrary to crown archosaurs. This suggests a peculiar developmental constraint that strongly affected the evolution of tanystropheids. The data obtained just at the base of the archosauromorph phylogenetic tree are crucial for further studies on the modularity of vertebral columns of not only Triassic reptile groups but extant and other extinct animals as well., Competing Interests: We declare we have no competing interests., (© 2024 The Authors.)

The postcranial morphology of the extremely long-necked Tanystropheus hydroides is 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 hydroides differs 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 hydroides was 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 of Macrocnemus bassanii and now Tanystropheus hydroides reveal 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.

Tetrapod remains are extremely rare in the early Carnian (Late Triassic) Calcare del Predil Formation on the Italian side of the Julian Alps (Friuli Venezia Giulia Autonomous Region), which yielded the Raibl fossil assemblage including the famous "Raibl ichthyofauna". A new tanystropheid archosauromorph, Raibliania calligarisi, is here named based on a partial skeleton found in this formation along the Prasnig Brook. The new taxon is similar to Tanystropheus, sharing with it the very elongated cervical vertebrae, but is characterized by the presence of relatively large button-like teeth; apex of the neural spines not transversely thickened; articular facets on the upper corners of the neural spines of the dorsal vertebrae; short and distally pointed pleurapophyses of the second and last 'lumbar' vertebra; ilium with a long preacetabular process bearing a robust and transversely thick tuberosity and a medial ridge bordering the entire ventral part of the iliac blade; and pubis with a cranial process. Tanystropheids lived in the present day Alps and Prealps of Friuli Venezia Giulia Region from the late Anisian to the mid-late Norian and were an important component of the coastal reptile faunas, although they were less common than eusauropterygians and placodonts during the early Carnian. Tanystropheids with extremely elongated cervical vertebrae are reported in this area from the upper Anisian to the lower Carnian.

Tanystropheus and other possible archosauromorph fossils have been discovered from Middle to Upper Triassic shallow marine sedimentary sections in Villány (Villány Hills, southern Hungary). Four fragmentary cervical vertebrae can be assigned to Tanystropheus sp. based on characteristic features including the strongly elongate and hollow vertebral body with extremely reduced neural spine. Besides the cervicals, various teeth, classified into four different morphotypes including longitudinally striated, carinated and ziphodont ones, are thought to belong to archosauromorphs, since they markedly differ from the frequently found teeth of fish and sauropterygians. In addition, three enigmatic cranial bones that might represent some circumorbital elements, have been found as well, and are referred to here as Sauropsida indet. These fossils, originated from the same tectonic unit as those from the Anisian of Bihor (Romania), are of great importance for a better understanding of the poorly known semi-aquatic to terrestrial vertebrate fauna of the Middle to Late Triassic of central Europe. [ABSTRACT FROM AUTHOR]

The Chañares Formation exhibits one of the most important archosauriform records of early Carnian ecosystems. Here we present new data on the palaeohistology of Chañares archosauriforms and provide new insights into their paleobiology, as well as possible phylogenetically informative traits. Bone microstructure of Lagerpeton chanarensis and Tropidosuchus romeri is dominated by fibro-lamellar tissue and dense vascularization. On the other hand, Chanaresuchus bonapartei is more densely vascularized, but with cyclical growth characterized by alternate fibro-lamellar, parallel-fibered and lamellar-zonal tissues. Dense vascularization and fibro-lamellar tissue imply fast growth and high metabolic rates for all these taxa. These histological traits may be tentatively interpreted as a possible adaptative advantage in front of Chañares Formation environmental conditions. [ABSTRACT FROM AUTHOR]

The homology and evolution of the archosaur ankle is a controversial topic that has been deeply studied using evidence from both extinct and extant taxa. In early stem archosaurs, the astragalus and calcaneum form the ancestral proximal tarsus and a single ossification composes the centrale series. In more recent stem archosaurs, the centrale is incorporated to the proximal row of tarsals laterally contacting the astragalus. This bone is subsequently lost as an independent ossification before the last common ancestor of birds and crocodilians, but the evolutionary fate of this element remains mostly unexplored. Here, we integrate embryological and palaeontological data with morphogeometric analyses to test the hypothesis of loss of the centrale or, alternatively, its incorporation into the archosaurian astragalus. Our results support the latter hypothesis, indicating that the astragalus developed ancestrally from two ossification centres in stem archosaurs and that the supposed tibiale of bird embryos represents a centrale. This conclusion agrees with previous embryological studies that concluded that the tibiale never develops in diapsids. [ABSTRACT FROM AUTHOR]

Tetrapod remains are extremely rare in the early Carnian (Late Triassic) Calcare del Predil Formation on the Italian side of the Julian Alps (Friuli Venezia Giulia Autonomous Region), which yielded the Raibl fossil assemblage including the famous "Raibl ichthyofauna". A new tanystropheid archosauromorph, Raibliania calligarisi, is here named based on a partial skeleton found in this formation along the Prasnig Brook. The new taxon is similar to Tanystropheus, sharing with it the very elongated cervical vertebrae, but is characterized by the presence of relatively large button-like teeth; apex of the neural spines not transversely thickened; articular facets on the upper corners of the neural spines of the dorsal vertebrae; short and distally pointed pleurapophyses of the second and last 'lumbar' vertebra; ilium with a long preacetabular process bearing a robust and transversely thick tuberosity and a medial ridge bordering the entire ventral part of the iliac blade; and pubis with a cranial process. Tanystropheids lived in the present day Alps and Prealps of Friuli Venezia Giulia Region from the late Anisian to the mid-late Norian and were an important component of the coastal reptile faunas, although they were less common than eusauropterygians and placodonts during the early Carnian. Tanystropheids with extremely elongated cervical vertebrae are reported in this area from the upper Anisian to the lower Carnian. [ABSTRACT FROM AUTHOR]

Several partially articulated specimens and numerous isolated bones of Ozimek volans gen. et sp. nov., from the late Carnian lacustrine deposits exposed at Krasiejów in southern Poland, enable a reconstruction of most of the skeleton. The unique character of the animal is its enlarged plate-like coracoids presumably fused with sterna. Other aspects of the skeleton seem to be comparable to those of the only known specimen of Sharovipteryx mirabilis from the latest Middle Triassic of Kyrgyzstan, which supports interpretation of both forms as protorosaurians. One may expect that the pectoral girdle of S. mirabilis, probably covered by the rock matrix in its only specimen, was similar to that of O. volans gen. et sp. nov. The Krasiejów material shows sharp teeth, low crescent scapula, three sacrals in a generalized pelvis (two of the sacrals being in contact with the ilium) and curved robust metatarsal of the fifth digit in the pes, which are unknown in Sharovipteryx. Other traits are plesiomorphic and, except for the pelvic girdle and extreme elongation of appendages, do not allow to identify any close connection of the sharovipterygids within the Triassic protorosaurians.

Erythrosuchidae is a clade of early archosauriform reptiles, which were apex predators in many late Early and Middle Triassic ecosystems, following the Permo-Triassic mass extinction. Erythrosuchids had a worldwide distribution, with well-preserved fossil material known from South Africa, European Russia, and China. We here redescribe the anatomy and revise the taxonomy of Guchengosuchus shiguaiensis, which is one of the stratigraphically oldest erythrosuchids and is known from a single partial skeleton from the lowermost Middle Triassic (lower Anisian) lower Ermaying Formation of Shaanxi Province, China. We provide a new differential diagnosis for Guchengosuchus shiguaiensis, and identify a series of autapomorphies relating to the morphologies of the skull roof and vertebrae. Incorporating updated anatomical information for Guchengosuchus into the most comprehensive morphological phylogenetic analysis available for early archosauromorphs recovers it as an early branching member of Erythrosuchidae, outside of the clade formed by Garjainia, Erythrosuchus, Chalishevia, and Shansisuchus. Fugusuchus hejiapanensis, from the uppermost Lower Triassic to lower Middle Triassic Heshanggou Formation of China, is recovered as the earliest branching member of Erythrosuchidae.

Many traits have been linked to extinction risk among modern vertebrates, including mode of life and body size. However, previous work has indicated there is little evidence that body size, or any other trait, was selective during past mass extinctions. Here, we investigate the impact of the Triassic–Jurassic mass extinction on early Archosauromorpha (basal dinosaurs, crocodylomorphs and their relatives) by focusing on body size and other life history traits. We built several new archosauromorph maximum‐likelihood supertrees, incorporating uncertainty in phylogenetic relationships. These supertrees were then employed as a framework to test whether extinction had a phylogenetic signal during the Triassic–Jurassic mass extinction, and whether species with certain traits were more or less likely to go extinct. We find evidence for phylogenetic signal in extinction, in that taxa were more likely to become extinct if a close relative also did. However, there is no correlation between extinction and body size, or any other tested trait. These conclusions add to previous findings that body size, and other traits, were not subject to selection during mass extinctions in closely‐related clades, although the phylogenetic signal in extinction indicates that selection may have acted on traits not investigated here. [ABSTRACT FROM AUTHOR]

Erythrosuchidae is a clade of early archosauriform reptiles, which were apex predators in many late Early andMiddle Triassic ecosystems, following the Permo-Triassic mass extinction. Erythrosuchids had a worldwide distribution, with well-preserved fossil material known from South Africa, European Russia, and China. We here redescribe the anatomy and revise the taxonomy of Guchengosuchus shiguaiensis, which is one of the stratigraphically oldest erythrosuchids and is known from a single partial skeleton from the lowermost Middle Triassic (lower Anisian) lower Ermaying Formation of Shaanxi Province, China. We provide a new differential diagnosis for Guchengosuchus shiguaiensis, and identify a series of autapomorphies relating to the morphologies of the skull roof and vertebrae. Incorporating updated anatomical information for Guchengosuchus into the most comprehensive morphological phylogenetic analysis available for early archosauromorphs recovers it as an early branching member of Erythrosuchidae, outside of the clade formed by Garjainia, Erythrosuchus, Chalishevia, and Shansisuchus. Fugusuchus hejiapanensis, from the uppermost Lower Triassic to lower Middle Triassic Heshanggou Formation of China, is recovered as the earliest branching member of Erythrosuchidae. [ABSTRACT FROM AUTHOR]

Tanystropheidae is a clade of early archosauromorphs with a reported distribution ranging from the Early to the Late Triassic of Asia, Europe, and North America. Although some specimens with possible tanystropheid affinities from the Lower Triassic beds of Brazil have been previously attributed to "Protorosauria", little is known about the tanystropheid record in Gondwana. Here, two new and one previously reported specimen from the Sanga do Cabral Formation (Induan-Olenekian) of Brazil are described and interpreted as Tanystropheidae. These records, together with other tetrapods previously reported for the Sanga do Cabral Formation, increase the knowledge of the biotic diversification during the beginning of the Triassic. This contribution reinforces that the archosauromorph diversification occurred shortly after the Permo-Triassic extinction, making the Sanga do Cabral Formation an important unit for the study of early Mesozoic faunas. [ABSTRACT FROM AUTHOR]

Two isolated teeth, a dorsal vertebra, fragments of a humerus and femur, a fragmentary pubic "boot" and part of an ischium shaft, identified here as belonging to a large predatory archosaur were discovered in the Upper Triassic site at Marciszów near Zawiercie (southern Poland). Comparisons of the new fossils from Marciszów with the dorsal vertebrae, pubic "boot", ischium and femur of the theropod-like Smok wawelski from Lisowice (Silesia) reveal that the two taxa are very similar. Nevertheless, due to the lack of more diagnostic elements (e.g., braincase or cranial elements), we prefer to consider all described specimens from Marciszów as Smok sp. Smok sp. shares a low mound-like, anterior trochanter with trochanteric shelf on the femur, a massive pubic "boot" with a distinct depression (= bevelled area), and a transversely lenticular ischium shaft in cross-section with S. wawelski. Some observed characters of the dorsal vertebra (e.g., lack of some lamina, shape and position of zygapophyses), however, are different from S. wawelski and may also suggest that the new findings represent a second species of the genus in the Upper Triassic of Poland. The discovery of Smok sp. at Marciszów is significant because it is the second example of the co-occurrence of this genus with: (i) bones of a large dicynodont; and (ii) record of gnawed tetrapod bones. The discovery of Smok sp. and the lack of significant morphologic divergence from S. wawelski suggest that this taxon is the only large-bodied predator currently known from the Upper Triassic of Poland. This new evidence expands the record of the genus and contributes, in some measure, to our knowledge of the stratigraphical distribution of large predatory archosaurs from the Polish Upper Triassic bone-bearing levels. [ABSTRACT FROM AUTHOR]

The first osteohistological study focused exclusively on rhynchosaurs (non-archosauriform archosauromorphs), based on the hyperodapedontines Teyumbaita sulcognathus and Hyperodapedon sp., from the Upper Triassic of Southern Brazil, indicates a relatively rapid growth rate in early ontogeny shown by the fibrolamellar complex, with a change to slow intermittent growth during late ontogeny represented by parallel-fibred bone with several growth marks. Contrary to previous studies, which described a typical non-archosaur reptilian bone tissue pattern for rhynchosaurs, with growth marks extending across the entire cortex, we demonstrate that, in both studied taxa, the initial growth rate was faster in comparison to the later. This suggests that the ability of rapid growth at high rates was already present in basal non-archosauriform archosauromorphs.

Proterosuchidae represents the oldest substantial diversification of Archosauromorpha and plays a key role in understanding the biotic recovery after the end-Permian mass extinction. Proterosuchidae was long treated as a wastebasket taxon, but recent revisions have reduced its taxonomic content to five valid species from the latest Permian of Russia and the earliest Triassic (Induan) of South Africa and China. In addition to these occurrences, several isolated proterosuchid bones have been reported from the Induan Panchet Formation of India for over 150 years. Following the re-study of historical specimens and newly collected material from this unit, we erect the new proterosuchid species Samsarasuchus pamelae , which is represented by most of the presacral vertebral column. We also describe cf. proterosuchid and proterosuchid cranial, girdle and limb bones that are not referred to Samsarasuchus pamelae . Phylogenetic analyses recovered Samsarasuchus pamelae within the new proterosuchid clade Chasmatosuchinae. The taxonomic diversity of Proterosuchidae is substantially expanded here, with at least 11 nominal species and several currently unnamed specimens, and a biogeographical range encompassing present-day South Africa, China, Russia, India, Brazil, Uruguay and Australia. This indicates a broader taxonomic, phylogenetic and biogeographic diversification of Proterosuchidae than previously thought in the aftermath of the end-Permian mass extinction., Competing Interests: We declare we have no competing interests., (© 2023 The Authors.)

One of the key faunal transitions in Earth history occurred after the Permo- Triassic mass extinction (ca 252.2 Ma), when the previously obscure archosauromorphs (which include crocodylians, dinosaurs and birds) become the dominant terrestrial vertebrates. Here, we place all known middle Permian--early Late Triassic archosauromorph species into an explicit phylogenetic context, and quantify biodiversity change through this interval. Our results indicate the following sequence of diversification: a morphologically conservative and globally distributed post-extinction 'disaster fauna'; a major but cryptic and poorly sampled phylogenetic diversification with significantly elevated evolutionary rates; and a marked increase in species counts, abundance, and disparity contemporaneous with global ecosystem stabilization some 5 million years after the extinction. This multiphase event transformed global ecosystems, with far-reaching consequences for Mesozoic and modern faunas. [ABSTRACT FROM AUTHOR]

The origin of the avian hand, with its reduced and fused carpals and digits, from the five-fingered hands and complex wrists of early dinosaurs represents one of the major transformations of manus morphology among tetrapods. Much attention has been directed to the later part of this transition, from four- to three-fingered taxa. However, earlier anatomical changes may have influenced these later modifications, possibly paving the way for a later frameshift in digit identities. We investigate the five- to four-fingered transition among early dinosaurs, along with changes in carpus morphology. New three-dimensional reconstructions from computed tomography data of the manus of the Triassic and Early Jurassic theropod dinosaurs Coelophysis bauri and Megapnosaurus rhodesiensis are described and compared intra- and interspecifically. Several novel findings emerge from these reconstructions and comparisons, including the first evidence of an ossified centrale and a free intermedium in some C. bauri specimens, as well as confirmation of the presence of a vestigial fifth metacarpal in this taxon. Additionally, a specimen of C. bauri and an unnamed coelophysoid from the Upper Triassic Hayden Quarry, New Mexico, are to our knowledge the only theropods (other than alvarezsaurs and birds) in which all of the distal carpals are completely fused together into a single unit. Several differences between the manus of C. bauri and M. rhodesiensis are also identified. We review the evolution of the archosauromorph manus more broadly in light of these new data, and caution against incorporating carpal characters in phylogenetic analyses of fine-scale relationships of Archosauromorpha, in light of the high degree of observed polymorphism in taxa for which large sample sizes are available, such as the theropod Coelophysis and the sauropodomorph Plateosaurus. We also find that the reduction of the carpus and ultimate loss of the fourth and fifth digits among early dinosaurs did not proceed in a neat, stepwise fashion, but was characterized by multiple losses and possible gains of carpals, metacarpals and phalanges. Taken together, the high degree of intra- and interspecific variability in the number and identities of carpals, and the state of reduction of the fourth and fifth digits suggest the presence of a 'zone of developmental variability' in early dinosaur manus evolution, from which novel avian-like morphologies eventually emerged and became channelized among later theropod clades. [ABSTRACT FROM AUTHOR]

The vertebrate recovery after the end-Permian mass extinction can be approached through the ichnological record, which is much more abundant than body fossils. The late Olenekian (Early Triassic) tetrapod ichnoassemblage of the Catalan Pyrenean Basin is the most complete and diverse of this age from Western Tethys. This extensional basin, composed of several depocenters, was formed in the latest phases of the Variscan orogeny (Pangea breakup) and was infilled by braided and meandering fluvial systems of the red-beds Buntsandstein facies. Abundant and diverse tetrapod ichnites are recorded in these facies, including Prorotodactylus mesaxonichnus isp. nov. (tracks possibly produced by euparkeriids), cf. Rotodactylus, at least two large chirotheriid morphotypes (archosauriform trackmakers), Rhynchosauroides cf. schochardti, two other undetermined Rhynchosauroides forms, an undetermined Morphotype A (archosauromorph trackmakers) and two types of Characichnos isp. (swimming traces, here associated to archosauromorph trackmakers). The Pyrenean ichnoassemblage suggests a relatively homogeneous ichnofaunal composition through the late Early Triassic of Central Pangea, characterized by the presence of Prorotodactylus and Rotodactylus. Small archosauromorph tracks dominate and present a wide distribution through the different fluviatile facies of the Triassic Pyrenean Basin, with large archosaurian footprints being present in a lesser degree. Archosauromorphs radiated and diversified through the Triassic vertebrate recovery, which ultimately lead to the archosaur and dinosaur dominance of the Mesozoic. [ABSTRACT FROM AUTHOR]

Several partially articulated specimens and numerous isolated bones of Ozimek volans gen. et sp. nov., from the late Carnian lacustrine deposits exposed at Krasiejów in southern Poland, enable a reconstruction of most of the skeleton. The unique character of the animal is its enlarged plate-like coracoids presumably fused with sterna. Other aspects of the skeleton seem to be comparable to those of the only known specimen of Sharovipteryx mirabilis from the latest Middle Triassic of Kyrgyzstan, which supports interpretation of both forms as protorosaurians. One may expect that the pectoral girdle of S. mirabilis, probably covered by the rock matrix in its only specimen, was similar to that of O. volans gen. et sp. nov. The Krasiejów material shows sharp teeth, low crescent scapula, three sacrals in a generalized pelvis (two of the sacrals being in contact with the ilium) and curved robust metatarsal of the fifth digit in the pes, which are unknown in Sharovipteryx. Other traits are plesiomorphic and, except for the pelvic girdle and extreme elongation of appendages, do not allow to identify any close connection of the sharovipterygids within the Triassic protorosaurians. [ABSTRACT FROM AUTHOR]

The early evolution of archosauromorphs during the Permo-Triassic constitutes an excellent empirical case study to shed light on evolutionary radiations in deep time and the timing and processes of recovery of terrestrial faunas after a mass extinction. However, macroevolutionary studies of early archosauromorphs are currently limited by poor knowledge of their phylogenetic relationships. In particular, one of the main early archosauromorph groups that need an exhaustive phylogenetic study is “Proterosuchia,” which as historically conceived includes members of both Proterosuchidae and Erythrosuchidae. A new data matrix composed of 96 separate taxa (several of them not included in a quantitative phylogenetic analysis before) and 600 osteological characters was assembled and analysed to generate a comprehensive higher-level phylogenetic hypothesis of basal archosauromorphs and shed light on the species-level interrelationships of taxa historically identified as proterosuchian archosauriforms. The results of the analysis using maximum parsimony include a polyphyletic “Prolacertiformes” and “Protorosauria,” in which the Permian Aenigmastropheus and Protorosaurus are the most basal archosauromorphs. The enigmatic choristoderans are either found as the sister-taxa of all other lepidosauromorphs or archosauromorphs, but consistently placed within Sauria. Prolacertids, rhynchosaurs, allokotosaurians and tanystropheids are the major successive sister clades of Archosauriformes. The Early Triassic Tasmaniosaurus is recovered as the sister-taxon of Archosauriformes. Proterosuchidae is unambiguosly restricted to five species that occur immediately after and before the Permo-Triassic boundary, thus implying that they are a short-lived “disaster” clade. Erythrosuchidae is composed of eight nominal species that occur during the Early and Middle Triassic. “Proterosuchia” is polyphyletic, in which erythrosuchids are more closely related to Euparkeria and more crownward archosauriforms than to proterosuchids, and several species are found widespread along the archosauromorph tree, some being nested within Archosauria (e.g., “Chasmatosaurus ultimus,” Youngosuchus). Doswelliids and proterochampsids are recovered as more closely related to each other than to other archosauromorphs, forming a large clade (Proterochampsia) of semi-aquatic to aquatic forms that includes the bizarre genus Vancleavea. Euparkeria is one of the sister-taxa of the clade composed of proterochampsians and archosaurs. The putative Indian archosaur Yarasuchus is recovered in a polytomy with Euparkeria and more crownward archosauriforms, and as more closely related to the Russian Dongusuchus than to other species. Phytosaurs are recovered as the sister-taxa of all other pseudosuchians, thus being nested within Archosauria.

The rhynchosaurian archosauromorphs are an important and diverse group of fossil tetrapods that first appeared during the Early Triassic and probably became extinct during the early Late Triassic (early Norian). Here, the early evolution of rhynchosaurs during the Early and early Middle Triassic (Induan−Anisian: 252.2−242 Mya) is reviewed based on new anatomical observations and the implications of these observations for the taxonomy, phylogenetic relationships and macroevolutionary history of the group. A quantitative phylogenetic analysis recovered a paraphyletic genus Rhynchosaurus, with Rhynchosaurus brodiei more closely related to hyperodapedontines than to Rhynchosaurus articeps. Therefore, a new genus is erected, resulting in the new combination Langeronyx brodiei. A body size analysis found two independent increases in size in the evolutionary history of rhynchosaurs, one among stenaulorhynchines and the other in the hyperodapedontine lineage. Maximum likelihood fitting of phenotypic evolution models to body size data found ambiguous results, with body size evolution potentially interpreted as either fitting either a non-directional Brownian motion model or a stasis model. A Dispersal−Extinction−Cladogenesis analysis reconstructed the areas that are now South Africa and Europe as the ancestral areas of Rhynchosauria and Rhynchosauridae, respectively. The reconstruction of dispersal events between geographic areas that are broadly separated paleolatitudinally implies that barriers to the dispersal of rhynchosaurs from either side of the paleo-Equator during the Middle Triassic were either absent or permeable.

The Permo-Triassic archosauromorph record is crucial to understand the impact of the Permo-Triassic mass extinction on the early evolution of the group and its subsequent dominance in Mesozoic terrestrial ecosystems. However, the Permo-Triassic archosauromorph record is still very poor in most continents and hampers the identification of global macroevolutionary patterns. Here we describe cranial and postcranial bones from the Permo-Triassic Buena Vista Formation of northeastern Uruguay that contribute to increase the meagre early archosauromorph record from South America. A basioccipital fused to both partial exoccipitals and three cervical vertebrae are assigned to Archosauromorpha based on apomorphies or a unique combination of characters. The archosauromorph remains of the Buena Vista Formation probably represent a multi-taxonomic assemblage composed of non-archosauriform archosauromorphs and a ‘proterosuchid-grade’ animal. This assemblage does not contribute in the discussion of a Late Permian or Early Triassic age for the Buena Vista Formation, but reinforces the broad palaeobiogeographic distribution of ‘proterosuchid grade’ diapsids in Permo-Triassic beds worldwide.

Background: Archosauromorpha originated in the middle-late Permian, radiated during the Triassic, and gave rise to the crown group Archosauria, a highly successful clade of reptiles in terrestrial ecosystems over the last 250 million years. However, scientific attention has mainly focused on the diversification of archosaurs, while their stem lineage (i.e. non-archosaurian archosauromorphs) has often been overlooked in discussions of the evolutionary success of Archosauria. Here, we analyse the cranial disparity of late Permian to Early Jurassic archosauromorphs and make comparisons between non-archosaurian archosauromorphs and archosaurs (including Pseudosuchia and Ornithodira) on the basis of two-dimensional geometric morphometrics. Results: Our analysis recovers previously unappreciated high morphological disparity for non-archosaurian archosauromorphs, especially during the Middle Triassic, which abruptly declined during the early Late Triassic (Carnian). By contrast, cranial disparity of archosaurs increased from the Middle Triassic into the Late Triassic, declined during the end-Triassic extinction, but re-expanded towards the end of the Early Jurassic. Conclusions: Our study indicates that non-archosaurian archosauromorphs were highly diverse components of terrestrial ecosystems prior to the major radiation of archosaurs, including dinosaurs, while disparity patterns of the Ladinian and Carnian indicate a gradual faunal replacement of stem archosaurs by the crown group, including a short interval of partial overlap in morphospace during the Ladinian. [ABSTRACT FROM AUTHOR]

The bone histology of non-archosauriform archosauromorphs is understudied but is important for tracing the evolution of growth in archosaurs and their closest relatives. Here, we describe the femoral and tibial histology of a single individual of Stenaulorhynchus stockleyi , a Middle Triassic rhynchosaur from Tanzania. Both elements are composed largely of moderately vascularized parallel-fibered bone tissue, which becomes avascular and more lamellar approaching the periosteum. The spacing between growth marks in S. stockleyi strongly attenuates with age, suggesting determinate growth for this taxon. Whereas the bone tissue suggests slower growth compared to South American rhynchosaurs, which have fibrolamellar bone tissue and exhibit more anastomoses between canals, the determinate growth pattern seems to be shared among hyperodapedontid rhynchosaurs. Although non-archosauriform archosauromorphs vary in the rate of bone deposition and growth in their first year, all taxa examined to date grew relatively slowly compared to most archosauriforms. [ABSTRACT FROM AUTHOR]