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Rhynchosauria is a group of extinct, exclusively Triassic, terrestrial, and herbivorous archosauromorphs, characterized by a peculiar maxillary-mandibular apparatus. They reached global distribution during the Carnian, with the Hyperodapedontinae clade. The rhynchosaurian record from South America is included in the Ladinian-?earliest Carnian Dinodontosaurus Assemblage Zone (Pinheiros-Chiniquá Sequence, Brazil) and Tarjadia Assemblage Zone (Chañares Formation, Argentina), and for strictly Carnian Hyperodapedon Assemblage Zone (Lower Candelária Sequence, Brazil) and Ischigualasto Formation (Argentina). Here, we present the first record of Hyperodapedontinae from the Brazilian Early Carnian Santacruzodon Assemblage Zone (Santa Cruz Sequence). The most diagnostic elements belong to a nearly complete left pes, which presents a combination of hyperodapedontine traits. The presence of a second individual of cf. Hyperodapedontinae is based on a fragmentary, isolated left metatarsal IV. An isolated fragment of premaxilla could belong to the second specimen (based on preservation mode) or to a third individual. This new report fills a gap within the South American rhynchosaurian distribution, strengthening biostratigraphic correlation with other regions from Gondwana (i.e., Madagascar), where similar and coeval faunas are known. The inclusion of these specimens in a phylogenetic dataset resulted in low resolution results, due to missing data because postcranial characters for rhynchosaurs are still poorly explored., (© 2024 American Association for Anatomy.)

The Triassic was a time of ecological upheaval as life recovered from the Permian-Triassic mass extinction. Archosauromorphs were a key component of the recovery, diversifying substantially during the Triassic and encompassing the origins of dinosaurs, pterosaurs and crocodylomorphs. Here, we explore the evolution of locomotion in Archosauromorpha to test whether dinosaurs show any distinctive locomotory features that might explain their success. We implement geometric morphometrics on limb bone shapes and use limb ratios to calculate bipedality and cursoriality metrics. We find that the Avemetatarsalia (dinosaurs, pterosaurs and relatives) exhibit more variable limb form and limb ratios than any other group, indicating a wider range of locomotory modes. The earliest avemetatarsalians were bipedal and cursorial, and their range of form increased through the Triassic with notable diversification shifts following extinction events. This is especially true of dinosaurs, even though these changes cannot be discriminated from a stochastic process. By contrast, the Pseudosuchia (crocodilians and relatives) were more restricted in limb form and locomotor mode with disparity decreasing through time, suggesting more limited locomotor adaptation and vulnerability to extinction. Perhaps the greater locomotor plasticity of dinosaurs gave them a competitive advantage in the changing climates of the Late Triassic.

Teyujagua paradoxa is a remarkable early archosauromorph from the Lower Triassic Sanga do Cabral Formation, Brazil. The species was originally described from an almost complete skull and a few associated cervical vertebrae, and no further postcranial elements were known at that time. Additional fieldwork in the Sanga do Cabral Formation, however, was successful in recovering a fairly complete postcranial skeleton attributable to the holotype. Here, we describe this new postcranial material, which is composed of cervical, dorsal, sacral and caudal vertebrae, limbs, pectoral and pelvic girdles, ribs, and gastralia. The description of its postcranial skeleton makes T. paradoxa one of the best-known early-diverging archosauromorphs. The cladistic analysis performed after the scoring of postcranial data recovered T. paradoxa in the same position initially described, close to the node that defines the Archosauriformes. Teyujagua paradoxa shares morphological features with representatives of early-diverging archosauromorphs and archosauriforms, with certain traits demonstrating a mosaic of plesiomorphic and apomorphic character states. We also performed partitioned morphospace and disparity analysis to elucidate the morphological disparity and evolutionary patterns among archosauromorphs. Teyujagua paradoxa occupies a notable position, suggesting an intermediate morphology between early archosauromorphs and proterosuchids. Disparity estimates highlighted Pseudosuchia and Avemetatarsalia as having the highest median disparity, reflecting their diverse cranial and postcranial morphologies, respectively. These findings offer valuable insights into archosauromorph macroevolution and adaptation., (© 2024 American Association for Anatomy.)

Allokotosauria, a clade of non-archosauriform archosauromorphs with a broad diversity of body plans, plays a crucial role in better understanding the evolutionary history of early diverging stem-archosaurs. Here we provide a detailed redescription of Malerisaurus robinsonae, a malerisaurine allokotosaur from the middle Carnian-lowermost Norian lower Maleri Formation, Pranhita-Godavari Basin, India. The new anatomical information available from recently discovered and well-preserved skeletons of various allokotosaurs, such as Azendohsaurus madagaskarensis, Shringasaurus indicus, Puercosuchus traverorum, and Malerisaurus-like taxa, and their comparison with Malerisaurus robinsonae enriches our understanding of the anatomy of this species. To reassess the phylogenetic relationships of Malerisaurus robinsonae, we revised its scorings and included eight additional allokotosaurian species to the already most comprehensive phylogenetic dataset focused on Permo-Triassic archosauromorphs. We modified 70 scorings for Malerisaurus robinsonae and the new analysis recovered this species at the base of Malerisaurinae and this group as the earliest branch of Azendohsauridae. Pamelaria dolichotrachela is found as the earliest diverging non-malerisaurine azendohsaurid and sister taxon to the Shringasaurus indicus + Azendohsaurus spp. clade. Trilophosaurid interrelationships are well resolved, with Teraterpeton hrynewichorum, Coelodontognathus ricovi, and Rutiotomodon tytthos as their successive earliest-branching species. The position of Anisodontosaurus greeri as a sister taxon to Variodens inopinatus bolsters long ghost lineages in the Late Triassic trilophosaurid record. A disparity analysis of tooth crown morphology shows that Allokotosauria is the most disparate Permo-Triassic archosauromorph clade, exploring the almost complete range of basic crown morphologies. Trilophosaurids occupy an area of the dental morphospace unique among archosauromorphs., (© 2024 American Association for Anatomy.)

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 Triassic was a time of ecological upheaval as life recovered from the Permian-Triassic mass extinction. Archosauromorphs were a key component of the recovery, diversifying substantially during the Triassic and encompassing the origins of dinosaurs, pterosaurs and crocodylomorphs. Here, we explore the evolution of locomotion in Archosauromorpha to test whether dinosaurs show any distinctive locomotory features that might explain their success. We implement geometric morphometrics on limb bone shapes and use limb ratios to calculate bipedality and cursoriality metrics. We find that the Avemetatarsalia (dinosaurs, pterosaurs and relatives) exhibit more variable limb form and limb ratios than any other group, indicating a wider range of locomotory modes. The earliest avemetatarsalians were bipedal and cursorial, and their range of form increased through the Triassic with notable diversification shifts following extinction events. This is especially true of dinosaurs, even though these changes cannot be discriminated from a stochastic process. By contrast, the Pseudosuchia (crocodilians and relatives) were more restricted in limb form and locomotor mode with disparity decreasing through time, suggesting more limited locomotor adaptation and vulnerability to extinction. Perhaps the greater locomotor plasticity of dinosaurs gave them a competitive advantage in the changing climates of the Late Triassic., 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.

Heteropelta boboi is a new archosauriform reptile from the upper Anisian of northeastern Italy represented by a fragment of dorsal armor with a row of neural arches of the dorsal vertebrae. The dorsal armor of the new taxon is composed of two columns of paramedian osteoderms and at least six columns of lateral osteoderms. Unlike other armored archosaurs, the osteoderms are imbricated with the posterior osteoderm overlapping the anterior one. The low neural arches bear small neural spines and long postzygapophyses. The osteoderms of the lateral columns increase in size and change in shape from the most medial to the most lateral columns. Among the Archosauriformes, only the non-archosaur proterochampsians Vancleavea campi, Litorosuchus somnii, and the doswelliids have dorsal armor comprised of more than two columns of osteoderms per side, but the morphology and arrangement of their osteoderms is unlike those of the new Italian taxon. A cladistic analysis of Archosauromorpha positions Heteropelta boboi as either a basal phytosaur or a basal suchian. However, a second cladistic analysis focused on armored archosaurs alternatively positions the new taxon as a basal archosauriform, basal suchian, basal loricatan or crocodylomorph. Better resolution of the phylogenetic relationships of Heteropelta boboi will likely be obtained only with the discovery of cranial and postcranial remains associated with its diagnostic armor elements.

The Late Triassic archosauromorph Trilophosaurus buettneri is known from many sites in the Dockum Group and Chinle Formation of Texas and Arizona, spanning nearly 10 Ma from the Otischalkian to the Adamanian estimated holochrons. The holotype specimen, UMMP 2338, was described as a small fragment of a lower jaw in 1928 by E.C. Case. However, at the time no other specimens of the taxon were known. The present availability of several nearly complete skulls, mandibles, and postcranial skeletons referred to T. buettneri allows for reexamination of the anatomical context of the holotype. Here, we provide high-resolution photographs and 3D models of UMMP 2338 to justify our reidentification of the holotypic specimen of T. buettneri as an incomplete maxilla, which may have important implications for future work on trilophosaurid archosauromorphs. [ABSTRACT FROM AUTHOR]

Erythrosuchids are Early–Middle Triassic bulky, quadrupedal archosauromorphs with proportionally large heads, and they were among the first archosauromorphs to occupy the role of terrestrial hypercarnivores. Recent work has expanded anatomical and phylogenetic knowledge of erythrosuchids, but a thorough understanding is still lacking, and Erythrosuchus africanus is the only species for which a thorough anatomical description of most of the skeleton has been published. Garjainia prima, from the upper Olenekian of Russia, represents one of the oldest known erythrosuchids and includes the best-preserved skull of an Early Triassic erythrosuchid. Here, we expand understanding of erythrosuchid anatomy by providing the first detailed and comprehensive description of the holotype of Garjainia prima. We consider ' Vjushkovia triplicostata ' to be a junior synonym of Garjainia prima, because of their extremely similar morphology, with the two character states that differ between them being variable intraspecifically in some other diapsids. A new phylogenetic analysis recovered a monophyletic Garjainia as sister to Middle Triassic erythrosuchids and recovered Fugusuchus hejiapanensis as sister to all other erythrosuchids. Branch support for Erythrosuchidae is low, but the Middle Triassic erythrosuchid clade is well supported by characters with low homoplasy, indicating that their morphology was specialized in ways rarely explored by other Triassic archosauromorphs. [ABSTRACT FROM AUTHOR]

Tanystropheidae is a clade of early archosauromorphs with high morphological disparity and a wide geographical distribution. The origin and early radiation of Tanystropheidae are still incompletely understood. Here we report Luxisaurus terrestris gen. et sp. nov., a new archosauromorph collected from the marine Upper Member of Guanling Formation (Pelsonian substage, Anisian, Middle Triassic) in Luxi County, Yunnan Province, China, and phylogenetically recovered as a tanystropheid. The morphology of Luxisaurus is consistent with a terrestrial lifestyle. This is the first occurrence of a tanystropheid found from the Anisian Guanling Formation in SW China and also the earliest tanystropheid from the eastern Tethys. Phylogenetic analysis shows that Dinocephalosauridae forms the sister group of Tanystropheidae, Fuyuansaurus occupies the most basal position in Tanystropheidae, and Luxisaurus forms the sister group to a clade consisting of all other tanystropheids. Considering that Fuyuansaurus and Luxisaurus are found exclusively in South China, we hypothesize that Tanystropheidae originated in the Early Triassic of South China and then dispersed globally. The discovery of Early Triassic tanystropheids in South China is needed to further support this hypothesis. The discovery of Luxisaurus expands the stratigraphical and geographical distribution of Tanystropheidae, and provides new insights into the origin and early radiation of the clade. [ABSTRACT FROM AUTHOR]

Rhynchosauria is a group of archosauromorph reptiles abundant in terrestrial ecosystems of the Middle Triassic. Mesosuchus is one of the earliest and basalmost rhynchosaurs, playing an important role not only for the understanding of the evolution of the group as a whole, but also of archosauromorphs in general. The braincase of Mesosuchus has been previously described, albeit not in detail, and the middle and inner ears were missing. Here, we provide new information based on micro-computed tomography scanning of the best-preserved specimen of Mesosuchus, SAM-PK-6536. Contrary to what has been stated previously, the braincase of Mesosuchus is dorso-ventrally tall. The trigeminal foramen lies in a deep recess on the prootic whose flat ventral rim could indicate the articulation surface to the laterosphenoid, although no such element was found. The middle ear of Mesosuchus shows a small and deeply recessed fenestra ovalis, with the right stapes preserved in situ. It has a rather stout, imperforated and posteriorly directed shaft with a small footplate. These features suggest that the ear of Mesosuchus was well-suited for the detection of low-frequency sounds. The semicircular canals are slender and elongate and the floccular fossa is well-developed. This is indicative of a refined mechanism for gaze stabilization, which is usually related to non-sprawling postures. The most striking feature of the Mesosuchus braincase is, however, the presence of a pneumatic sinus in the basal tubera. The sinus is identified as originating from the pharyngotympanic system, implying ossified Eustachian tubes. Braincase pneumatization has not yet been a recognized feature of stem-archosaurs, but the potential presence of pneumatic foramina in an array of taxa, recognized here as such for the first time, suggests braincase sinuses could be present in many other archosauromorphs.