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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.

Abstract Variations in the shape and size of teeth have been associated with changes in enamel ultrastructure across odontocetes. Characterizing these features in extinct taxa can elucidate their functional morphology and feeding strategy, while also shedding light into macroevolutionary patterns during the evolutionary history of cetaceans. This study aimed to (1) describe the enamel and dentine ultrastructure of the Early Miocene odontocetes Notocetus vanbenedeni and Phoberodon arctirostris from Patagonia (Argentina) and (2) quantify tooth and enamel ultrastructure morphological disparity among odontocetes. Enamel was predominantly prismatic, thin in the anterior tooth of N. vanbenedeni and P. arctirostris; whilst thick on the posterior tooth of N. vanbenedeni. Together with skull morphology, data suggests a raptorial feeding strategy for P. arctirostris and a combination suction feeding method for N. vanbenedeni. Statistical analyses supported these inferences, indicating that enamel characters are useful for paleoecological research. Morphological disparity analyses showed that extant odontocetes occupy a larger morphospace and have more disparate morphologies, whilst extinct odontocetes were more similar among each other than with the extant group. There was no clear phylogenetic-based grouping, suggesting that tooth and enamel ultrastructure disparity were mainly driven by ecological pressures. These results highlight enamel ultrastructure as a source for broader-scale paleoecological studies in cetaceans.

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.

Abstract Sauropodomorph dinosaurs were the dominant medium to large-sized herbivores of most Mesozoic continental ecosystems, being characterized by their long necks and reaching a size unparalleled by other terrestrial animals (> 60 tonnes). Our study of morphological disparity across the entire skeleton shows that during the Late Triassic the oldest known sauropodomorphs occupied a small region of morphospace, subsequently diversifying both taxonomically and ecologically, and shifting to a different and broader region of the morphospace. After the Triassic–Jurassic boundary event, there are no substancial changes in sauropodomorph morphospace occupation. Almost all Jurassic sauropodomorph clades stem from ghost lineages that cross the Triassic–Jurassic boundary, indicating that variations after the extinction were more related to changes of pre-existing lineages (massospondylids, non-gravisaurian sauropodiforms) rather than the emergence of distinct clades or body plans. Modifications in the locomotion (bipedal to quadrupedal) and the successive increase in body mass seem to be the main attributes driving sauropodomorph morphospace distribution during the Late Triassic and earliest Jurassic. The extinction of all non-sauropod sauropodomorphs by the Toarcian and the subsequent diversification of gravisaurian sauropods represent a second expansion of the sauropodomorph morphospace, representing the onset of the flourishing of these megaherbivores that subsequently dominated in Middle and Late Jurassic terrestrial assemblages.

Erythrosuchidae is a clade of early archosauriform reptiles that were large-bodied, hypercarnivorous, possibly apex pred-ators in late Early and Middle Triassic ecosystems following the Permo-Triassic mass extinction. Chalishevia cothurnatafrom the late Middle Triassic (Ladinian) of Russia, is the stratigraphically youngest known erythrosuchid species, but the holotype and referred material of this taxon has received little study. Here, we provide the first detailed anatomical description of C. cothurnata, including comparisons to other erythrosuchids. Although known from relatively fragmen-tary material, the anatomy of C. cothurnata is distinctive, including an autapomorphic strongly slanted ventral border of the antorbital fossa. The presence of a large accessory opening (the “accessory antorbital fenestra”) in the skull between the premaxilla, nasal and maxilla, together with the inferred presence of a narrow postnarial process of the premaxilla that articulated with a slot on the nasal, provides strong evidence for a sister taxon relationship between C. cothurnata and the erythrosuchid Shansisuchus shansisuchus from the early Middle Triassic (Anisian) of China. The inferred basal skull length of C. cothurnata was approximately 80 cm, making it one of the largest erythrosuchids known.

We describe a new small-bodied coelophysoid theropod dinosaur, Pendraig milnerae gen. 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 recovers P. milnerae as 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 for P. 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 for P. 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.

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.

Erythrosuchidae were large-bodied, quadrupedal, predatory archosauriforms that dominated the hypercarnivorous niche in the aftermath of the Permo-Triassic mass extinction. Garjainia, one of the oldest members of the clade, is known from the late Olenekian of European Russia. The holotype of Garjainia prima comprises a well-preserved skull, but highly incomplete postcranium. Recent taxonomic reappraisal demonstrates that material from a bone bed found close to the type locality, previously referred to as ‘Vjushkovia triplicostata', is referable to G. prima. At least, seven individuals comprising cranial remains and virtually the entire postcranium are represented, and we describe this material in detail for the first time. An updated phylogenetic analysis confirms previous results that a monophyletic Garjainia is the sister taxon to a clade containing Erythrosuchus, Shansisuchus and Chalishevia. Muscle scars on many limb elements are clear, allowing reconstruction of the proximal locomotor musculature. We calculate the body mass of G. prima to have been 147–248 kg, similar to that of an adult male lion. Large body size in erythrosuchids may have been attained as part of a trend of increasing body size after the Permo-Triassic mass extinction and allowed erythrosuchids to become the dominant carnivores of the Early and Middle Triassic.

Correctly identifying taxa at the root of major clades or the oldest clade-representatives is critical for meaningful interpretations of evolution. A small, partially crushed skull from the Late Triassic (Norian) of Connecticut, USA, originally described as an indeterminate rhynchocephalian saurian, was recently named Colobops noviportensis and reinterpreted as sister to all remaining Rhynchosauria, one of the earliest and globally distributed groups of herbivorous reptiles. It was also interpreted as having an exceptionally reinforced snout and powerful bite based on an especially large supratemporal fenestra. Here, after a re-analysis of the original scan data, we show that the skull was strongly dorsoventrally compressed post-mortem, with most bones out of life position. The cranial anatomy is consistent with that of other rhynchocephalian lepidosauromorphs, not rhynchosaurs. The ‘reinforced snout' region and the ‘exceptionally enlarged temporal region’ are preservational artefacts and not exceptional among clevosaurid rhynchocephalians. Colobops is thus not a key taxon for understanding diapsid feeding apparatus evolution.

Mass extinctions are thought to produce ‘disaster faunas’, communities dominated by a small number of widespread species. Here, Button et al. develop a phylogenetic network approach to test this hypothesis and find that mass extinctions did increase faunal cosmopolitanism across Pangaea during the late Palaeozoic and early Mesozoic.

Abstract The early evolution of archosauromorphs (bird- and crocodile-line archosaurs and stem-archosaurs) represents an important case of adaptive radiation that occurred in the aftermath of the Permo-Triassic mass extinction. Here we enrich the early archosauromorph record with the description of a moderately large (3–4 m in total length), herbivorous new allokotosaurian, Shringasaurus indicus, from the early Middle Triassic of India. The most striking feature of Shringasaurus indicus is the presence of a pair of large supraorbital horns that resemble those of some ceratopsid dinosaurs. The presence of horns in the new species is dimorphic and, as occurs in horned extant bovid mammals, these structures were probably sexually selected and used as weapons in intraspecific combats. The relatively large size and unusual anatomy of Shringasaurus indicus broadens the morphological diversity of Early–Middle Triassic tetrapods and complements the understanding of the evolutionary mechanisms involved in the early archosauromorph diversification.

Erythrosuchidae are a globally distributed and important group of apex predators that occupied Early and Middle Triassic terrestrial ecosystems following the Permo-Triassic mass extinction. The stratigraphically oldest known genus of Erythrosuchidae is Garjainia Ochev, 1958, which is known from the late Early Triassic (late Olenekian) of European Russia and South Africa. Two species of Garjainia have been reported from Russia: the type species, Garjainia prima Ochev, 1958, and ‘Vjushkovia triplicostata’ von Huene, 1960, which has been referred to Garjainia as either congeneric (Garjainia triplicostata) or conspecific (G. prima). The holotype of G. prima has received relatively extensive study, but little work has been conducted on type or referred material attributed to ‘V. triplicostata’. However, this material includes well-preserved fossils representing all parts of the skeleton and comprises seven individuals. Here, we provide a comprehensive description and review of the cranial anatomy of material attributed to ‘V. triplicostata’, and draw comparisons with G. prima. We conclude that the two Russian taxa are indeed conspecific, and that minor differences between them result from a combination of preservation or intraspecific variation. Our reassessment therefore provides additional information on the cranial anatomy of G. prima. Moreover, we quantify relative head size in erythrosuchids and other early archosauromorphs in an explicit phylogenetic context for the first time. Our results show that erythrosuchids do indeed appear to have disproportionately large skulls, but that this is also true for other early archosauriforms (i.e. proterosuchids), and may reflect the invasion of hypercarnivorous niches by these groups following the Permo-Triassic extinction.

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.

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.

Non-avian saurischian skulls underwent at least 165 million years of evolution and shapes varied from elongated skulls, such as in the theropod Coelophysis, to short and box-shaped skulls, such as in the sauropod Camarasaurus. A number of factors have long been considered to drive skull shape, including phylogeny, dietary preferences and functional constraints. However, heterochrony is increasingly being recognized as an important factor in dinosaur evolution. In order to quantitatively analyse the impact of heterochrony on saurischian skull shape, we analysed five ontogenetic trajectories using two-dimensional geometric morphometrics in a phylogenetic framework. This allowed for the comparative investigation of main ontogenetic shape changes and the evaluation of how heterochrony affected skull shape through both ontogenetic and phylogenetic trajectories. Using principal component analyses and multivariate regressions, it was possible to quantify different ontogenetic trajectories and evaluate them for evidence of heterochronic events allowing testing of previous hypotheses on cranial heterochrony in saurischians. We found that the skull shape of the hypothetical ancestor of Saurischia likely led to basal Sauropodomorpha through paedomorphosis, and to basal Theropoda mainly through peramorphosis. Paedomorphosis then led from Orionides to Avetheropoda, indicating that the paedomorphic trend found by previous authors in advanced coelurosaurs may extend back into the early evolution of Avetheropoda. Not only are changes in saurischian skull shape complex due to the large number of factors that affected it, but heterochrony itself is complex, with a number of possible reversals throughout non-avian saurischian evolution. In general, the sampling of complete ontogenetic trajectories including early juveniles is considerably lower than the sampling of single adult or subadult individuals, which is a major impediment to the study of heterochrony on non-avian dinosaurs. Thus, the current work represents an exploratory analysis. To better understand the cranial ontogeny and the impact of heterochrony on skull evolution in saurischians, the data set that we present here must be expanded and complemented with further sampling from future fossil discoveries, especially of juvenile individuals.

Here we describe a new theropod, Bicentenaria argentina nov. gen. et nov. sp., from the early Late Cretaceous of Patagonia. It is represented by more than a hundred bones belonging to different sized individuals, which were buried together in disarticulation after little transportation. The available association of skeletal elements suggests a gregarious behaviour for Bicentenaria, an ethological trait also recorded among other theropod clades. Increasing documentation of monospecific assemblages of different groups of theropods suggests that a gregarious behaviour may have constituted the ancestral condition for Theropoda, at least. Bicentenaria characterizes for the surangular bone with a high dorsal margin and a prominent lateral shelf, a retroarticular process that is low, wide and spoon-shaped, and quadrate bone with its lateral condyle larger than the medial one. Phylogenetic analysis found the Chinese Tugulusaurus and the Patagonian Bicentenaria as successive sister taxa of all other coelurosaurs, thus revealing the importance of the new taxon in the understanding of the early diversification of Coelurosauria. In particular, Bicentenaria amplifies the array of basal coelurosaurs that inhabited Gondwana during the Cretaceous, also including compsognathids, Aniksosaurus and Santanaraptor. Although still restricted to a handful of forms, available information indicates that Gondwana was a cradle for the evolution of different lineages of basal coelurosaurs, different from those documented in Upper Cretaceous beds in the northern landmasses. Analysis of body size distribution in averostran theropods results in the identification of two main episodes of drastic size reduction in the evolutionary history of Coelurosauria: one occurred at the initial radiation of the group (as represented by Bicentenaria, Zuolong, Tugulusaurus, compsognathids, and Aniksosaurus), and a second episode occurred at the early diverification of Paraves or avialans. Reduction in body size may have allowed adult coelurosaurians to exploit ecological niches not occupied before by larger basal averostrans.En la presente contribución se describe el nuevo terópodo Bicentenaria argentina nov. gen. et nov. sp., del Cretácico Tardío temprano de Patagonia. El nuevo taxón se encuentra representado por más de una centena de huesos pertenecientes a individuos de diferentes tamaños, los cuales fueron enterrados todos juntos en desarticulación, luego de un leve transporte. La asociación disponible de materiales sugiere hábitos gregarios para Bicentenaria, un rasgo etológico también registrado en otros clados de terópodos. La documentación recurrente de asociaciones monoespecíficas de diferentes grupos de terópodos sugiere que los hábitos gregarios pudieron haber constituido la condición ancestral para Theropoda. Bicentenaria se caracteriza por un hueso surangular con un margen dorsal prominente y con una repisa lateral bien desarrollada, un proceso retroarticular que es bajo, transversalmente amplio y en forma de cuchara, y un hueso cuadrado cuyo cóndilo lateral es de tamaño mucho mayor que el medial. El análisis filogenético recuperó al género chino Tugulusaurus y al patagónico Bicentenaria como grupos hermanos sucesivos de los restantes celurosaurios, revelando de este modo la importancia del nuevo taxón para el entendimiento de la radiación temprana de los Coelurosauria. En particular, Bicentenaria amplifica el elenco de celurosaurios basales que habitaron Gondwana durante el Cretácico, incluyendo entre otros a los compsognátidos, Aniksosaurus y Santanaraptor. A pesar de encontrarse restringida a un puñado de formas, la información disponible sugiere que Gondwana fue un importante centro para la evolución de diferentes linajes de celurosaurios basales, diferentes de aquellos representados en el Hemisferio Norte. Un análisis de la distribución del tamaño corporal en los terópodos averostros permite identificar dos episodios principales de drástica reducción del tamaño en el curso de la historia evolutiva de los Coelurosauria. El primero de estos episodios ocurrió durante la radiación inicial del grupo (tal como es representado por Bicentenaria, Zuolong, Tugulusaurus, compsognátidos, y Aniksosaurus), y el segundo episodio tomó lugar en la temprana diversificación de Paraves o Avialae. La reducción en la talla corporal pudo haber permitido a los celurosaurios adultos explotar nichos no ocupados anteriormente por terópodos averostros basales de mayor tamaño.

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.

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

Proterochampsids are crocodile-like, probably semi-aquatic, quadrupedal archosauriforms characterized by an elongated and dorsoventrally low skull. The group is endemic from the Middle-Late Triassic of South America. The most recently erected proterochampsid species is "Chanaresuchus ischigualastensis", based on a single, fairly complete skeleton from the early Late Triassic Ischigualasto Formation of northwestern Argentina. We describe here in detail the non-braincase cranial and postcranial anatomy of this species and revisit its taxonomy and phylogenetic relationships. The phylogenetic analysis recovered 'Chanaresuchus ischigualastensis' as part of a trichotomy together with Gualosuchus reigi and Chanaresuchus bonapartei. Accordingly, "Chanaresuchus ischigualastensis" can be potentially more closely related to Gualosuchus reigi, or even Rhadinosuchus gracilis, than to Chanaresuchus bonapartei. In addition, after discussing previously claimed synapomorphies of Chanaresuchus, we could not find unambiguous support for the monophyly of the genus. As a result, we propose here the erection of the new genus Pseudochampsa for 'Chanaresuchus ischigualastensis', which results in the new combination Pseudochampsa ischigualastensis. The information provided here about the anatomy and taxonomy of Pseudochampsa ischiguaslastensis will be useful for future quantitative analyses focused on the biogeography and macroevolutionary history of proterochampsids.

Sauria is the crown-group of Diapsida and is subdivided into Lepidosauromorpha and Archosauromorpha, comprising a high percentage of the diversity of living and fossil tetrapods. The split between lepidosauromorphs and archosauromorphs (the crocodile-lizard, or bird-lizard, divergence) is considered one of the key calibration points for molecular analyses of tetrapod phylogeny. Saurians have a very rich Mesozoic and Cenozoic fossil record, but their late Paleozoic (Permian) record is problematic. Several Permian specimens have been referred to Sauria, but the phylogenetic affinity of some of these records remains questionable. We reexamine and review all of these specimens here, providing new data on early saurian evolution including osteohistology, and present a new morphological phylogenetic dataset. We support previous studies that find that no valid Permian record for Lepidosauromorpha, and we also reject some of the previous referrals of Permian specimens to Archosauromorpha. The most informative Permian archosauromorph is Protorosaurus speneri from the middle Late Permian of Western Europe. A historically problematic specimen from the Late Permian of Tanzania is redescribed and reidentified as a new genus and species of basal archosauromorph: Aenigmastropheus parringtoni. The supposed protorosaur Eorasaurus olsoni from the Late Permian of Russia is recovered among Archosauriformes and may be the oldest known member of the group but the phylogenetic support for this position is low. The assignment of Archosaurus rossicus from the latest Permian of Russia to the archosauromorph clade Proterosuchidae is supported. Our revision suggests a minimum fossil calibration date for the crocodile-lizard split of 254.7 Ma. The occurrences of basal archosauromorphs in the northern (30°N) and southern (55°S) parts of Pangea imply a wider paleobiogeographic distribution for the group during the Late Permian than previously appreciated. Early archosauromorph growth strategies appear to be more diverse than previously suggested based on new data on the osteohistology of Aenigmastropheus.

Proterosuchidae are the most taxonomically diverse archosauromorph reptiles sampled in the immediate aftermath of the Permo-Triassic mass extinction and represent the earliest radiation of Archosauriformes (archosaurs and closely related species). Proterosuchids are potentially represented by approximately 15 nominal species collected from South Africa, China, Russia, Australia and India, but the taxonomic content of the group is currently in a state of flux because of the poor anatomic and systematic information available for several of its putative members. Here, the putative proterosuchid Tasmaniosaurus triassicus from the Lower Triassic of Hobart, Tasmania (Australia), is redescribed. The holotype and currently only known specimen includes cranial and postcranial remains and the revision of this material sheds new light on the anatomy of the animal, including new data on the cranial endocast. Several bones are re-identified or reinterpreted, contrasting with the descriptions of previous authors. The new information provided here shows that Tasmaniosaurus closely resembles the South African proterosuchid Proterosuchus, but it differed in the presence of, for example, a slightly downturned premaxilla, a shorter anterior process of maxilla, and a diamond-shaped anterior end of interclavicle. Previous claims for the presence of gut contents in the holotype of Tasmaniosaurus are considered ambiguous. The description of the cranial endocast of Tasmaniosaurus provides for the first time information about the anatomy of this region in proterosuchids. The cranial endocast preserves possibly part of the vomero-nasal ( = Jacobson's) system laterally to the olfactory bulbs. Previous claims of the absence of the vomero-nasal organs in archosaurs, which is suggested by the extant phylogenetic bracket, are questioned because its absence in both clades of extant archosaurs seems to be directly related with the independent acquisition of a non-ground living mode of life.