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Once known solely from dental material and thought to represent an early ornithischian dinosaur, the early-diverging pseudosuchian Revueltosaurus callenderi is described from a minimum of 12 skeletons from a monodominant bonebed in the upper part of the Chinle Formation of Arizona. This material includes nearly the entire skeleton and possesses a combination of plesiomorphic and derived character states that help clarify ingroup relationships within Pseudosuchia. A phylogenetic analysis recovers R. callenderi in a clade with Aetosauria and Acaenasuchus geoffreyi that is named Aetosauriformes. Key autapomorphies of R. callenderi include a skull that is longer than the femur, a complete carapace of dermal armor including paramedian and lateral rows, as well as ventral osteoderms, and a tail end sheathed in bone. Histology of the femur and associated osteoderms demonstrate that R. callenderi was slow growing and that the individuals from the bonebed were not young juveniles but had not ceased growing. A review of other material assigned to Revueltosaurus concludes that the genus cannot be adequately diagnosed based on the type materials of the three assigned species and that only R. callenderi can be confidently referred to Revueltosaurus. Published version

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.

Morphology forms the most fundamental level of data in vertebrate palaeontology because it is through interpretations of morphology that taxa are identified, creating the basis for broad evolutionary and palaeobiological hypotheses. Assessing maturity is one of the most basic aspects of morphological interpretation and provides the means to study the evolution of ontogenetic changes, population structure and palaeoecology, life-history strategies, and heterochrony along evolutionary lineages that would otherwise be lost to time. Saurian reptiles (the least-inclusive clade containing Lepidosauria and Archosauria) have remained an incredibly diverse, numerous, and disparate clade through their ~260-million-year history. Because of the great disparity in this group, assessing maturity of saurian reptiles is difficult, fraught with methodological and terminological ambiguity. We compiled a novel database of literature, assembling >900 individual instances of saurian maturity assessment, to examine critically how saurian maturity has been diagnosed. We review the often inexact and inconsistent terminology used in saurian maturity assessment (e.g. 'juvenile', 'mature') and provide routes for better clarity and cross-study coherence. We describe the various methods that have been used to assess maturity in every major saurian group, integrating data from both extant and extinct taxa to give a full account of the current state of the field and providing method-specific pitfalls, best practices, and fruitful directions for future research. We recommend that a new standard subsection, 'Ontogenetic Assessment', be added to the Systematic Palaeontology portions of descriptive studies to provide explicit ontogenetic diagnoses with clear criteria. Because the utility of different ontogenetic criteria is highly subclade dependent among saurians, even for widely used methods (e.g. neurocentral suture fusion), we recommend that phylogenetic context, preferably in the form of a phylogenetic bracket, be used to justify the use of a maturity assessment method. Different methods should be used in conjunction as independent lines of evidence when assessing maturity, instead of an ontogenetic diagnosis resting entirely on a single criterion, which is common in the literature. Critically, there is a need for data from extant taxa with well-represented growth series to be integrated with the fossil record to ground maturity assessments of extinct taxa in well-constrained, empirically tested methods.

The Placerias/Downs' Quarry complex in eastern Arizona, USA, is the most diverse Upper Triassic vertebrate locality known. We report a new short-faced archosauriform, Syntomiprosopus sucherorum gen. et sp. nov., represented by four incomplete mandibles, that expands that diversity with a morphology unique among Late Triassic archosauriforms. The most distinctive feature of Syntomiprosopus gen. nov. is its anteroposteriorly short, robust mandible with 3-4 anterior, a larger caniniform, and 1-3 "postcanine" alveoli. The size and shape of the alveoli and the preserved tips of replacement teeth preclude assignment to any taxon known only from teeth. Additional autapomorphies of S. sucherorum gen. et sp. nov. include a large fossa associated with the mandibular fenestra, an interdigitating suture of the surangular with the dentary, fine texture ornamenting the medial surface of the splenial, and a surangular ridge that completes a 90 degrees arc. The external surfaces of the mandibles bear shallow, densely packed, irregular, fine pits and narrow, arcuate grooves. This combination of character states allows an archosauriform assignment; however, an associated and similarly sized braincase indicates that Syntomiprosopus n. gen. may represent previously unsampled disparity in early-diverging crocodylomorphs. The Placerias Quarry is Adamanian (Norian, maximum depositional age similar to 219 Ma), and this specimen appears to be an early example of shortening of the skull, which occurs later in diverse archosaur lineages, including the Late Cretaceous crocodyliform Simosuchus. This is another case where Triassic archosauriforms occupied morphospace converged upon by other archosaurs later in the Mesozoic and further demonstrates that even well-sampled localities can yield new taxa. Department of Geological Sciences at Appalachian State University; College of Arts and Sciences at Appalachian State University; Honors College at Appalachian State University Published version Funding to support open access publication of this contribution was provided by the Department of Geological Sciences, the College of Arts and Sciences, and the Honors College at Appalachian State University.

Steyer, Jean-Sébastien, Peecook, Brandon R., Arbez, Thomas, Nesbitt, Sterling J., Tolan, Steve, Stocker, Michelle R., Smith, Roger M. H., Angielczyk, Kenneth D., Sidor, Christian A. (2021): New data on the Triassic temnospondyls from the Karoo rift basins of Tanzania and Zambia. Geodiversitas 43 (12): 365-376, DOI: http://doi.org/10.5281/zenodo.4906293, {"references":["ATTRIDGE J., BALL H., CHARIG A. & COX C. B. 1964. - The BMNH/University of London joint palaeontological expedition to Northern Rhodesia and Tanganyika 1963. Nature 201: 445- 449. https://doi.org/10.1038/201445a0","BARBOLINI N., BAMFORD M. K. & TOLAN S. 2016. - Permo-Triassic palynology and palaeobotany of Zambia; a review.Palaeontologia Africana 50: 18-30.","BARRETT P. M., SCISCIO L., VIGLIETTI P. A., BRODERICK T. J., SUAREZ C. A., SHARMAN G. R., JONES A. S., MUNYIKWA D., EDWARDS S. F., CHAPELLE K. E. J., DOLLMAN K. N., ZONDO M. & CHOINIERE J. 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Zoological Journal of the Linnean Society 128: 77-121. https:// doi.org/10.1111/j.1096-3642.2000.tb00650.x","ZITTEL K. VON 1887-1890. - Handbuch der Palaontologie. Abteilung 1. Palaozoologie Band III. Vertebrata (Pisces, Amphibia, Reptilia, Aves). Oldenbourg, Munchen, and Leipzig, 890 p."]}

Synopsis Pygopodids are elongate, functionally limbless geckos found throughout Australia. The clade presents low taxonomic diversity (∼45 spp.), but a variety of cranial morphologies, habitat use, and locomotor abilities that vary between and within genera. In order to assess potential relationships between cranial morphology and ecology, computed tomography scans of 29 species were used for 3D geometric morphometric analysis. A combination of 24 static landmarks and 20 sliding semi-landmarks were subjected to Generalized Procrustes Alignment. Disparity in cranial shape was visualized through Principal Component Analysis, and a multivariate analysis of variance (MANOVA) was used to test for an association between shape, habitat, and diet. A subset of 27 species with well-resolved phylogenetic relationships was used to generate a phylomorphospace and conduct phylogeny-corrected MANOVA. Similar analyses were done solely on Aprasia taxa to explore species-level variation. Most of the variation across pygopodids was described by principal component (PC) 1(54%: cranial roof width, parabasisphenoid, and occipital length), PC2 (12%: snout elongation and braincase width), and PC3 (6%: elongation and shape of the palate and rostrum). Without phylogenetic correction, both habitat and diet were significant influencers of variation in cranial morphology. However, in the phylogeny-corrected MANOVA, habitat remained weakly significant, but not diet, which can be explained by generic-level differences in ecology rather than among species. Our results demonstrate that at higher levels, phylogeny has a strong effect on morphology, but that influence may be due to small sample size when comparing genera. However, because some closely related taxa occupy distant regions of morphospace, diverging diets, and use of fossorial habitats may contribute to variation seen in these geckos.

Dramatic early Cenozoic climatic shifts resulted in faunal reorganization on a global scale. Among vertebrates, multiple groups of mammals (e.g., adapiform and omomyiform primates, mesonychids, taeniodonts, dichobunid artiodactyls) are well known from the Western Interior of North America in the warm, greenhouse conditions of the early Eocene, but a dramatic drop in the diversity of these groups, along with the introduction of more dry-tolerant taxa, occurred near the Eocene–Oligocene boundary. Crocodyliforms underwent a striking loss of diversity at this time as well. Pre-Uintan crocodyliform assemblages in the central Western Interior are characterized by multiple taxa, whereas Chadronian assemblages are depauperate with onlyAlligator prenasalispreviously known. Crocodyliform diversity through the intervening Uintan and Duchesnean is not well understood. The middle Eocene Devil’s Graveyard Formation (DGF) of southwest Texas provides new data from southern latitudes during that crucial period. A new specimen from the middle member of the DGF (late Uintan–Duchesnean) is the most complete cranial material of an alligatorid known from Paleogene deposits outside the Western Interior. We identify this specimen as a caimanine based on notched descending laminae of the pterygoids posterior to the choanae and long descending processes of the exoccipitals that are in contact with the basioccipital tubera. UnlikeEocaiman cavernensis, the anterior palatine process is rounded rather than quadrangular. The relationships and age of this new taxon support the hypothesis that the modern distribution of caimanines represents a contraction of a more expansive early Cenozoic distribution. We hypothesize that the range of caimanines tracked shifting warm, humid climatic conditions that contracted latitudinally toward the hothouse-icehouse transition later in the Eocene.

International audience; The Lamy Quarry (= the Gunter bonebed) is known for its extensive accumulation of temnospondyl skulls and skeletons from the Norian (Upper Triassic) Garita Creek Formation, south of Lamy, New Mexico. Although the quarry is monodominant for metoposaurids, reptile fossils are also present. The reptile material, briefly described and identified to least-inclusive taxonomic levels, consists of several diagnostic elements from azendohsaurid and tanystropheid archosauromorphs based on apomorphies observed in the fossils. The most remarkable materials are two anterior cervical vertebrae and the posterior portion of a hemimandible that are identified as belonging to a Malerisaurus-like taxon within the Azendohsauridae. The vertebrae are two times longer than the holotype material of Malerisaurus robinsonaeChatterjee, 1980 and Malerisaurus langstoniChatterjee, 1986, which suggests that azendohsaurids in the Late Triassic of North America reached much larger sizes than previously documented. The presence of a tanystropheid and azendohsaurid together supports the Adamanian age of the Garita Creek Formation and shows that azendohsaurids were present in what is now New Mexico.

Pterosaurs were the first vertebrates to evolve powered flight1 and comprised one of the main evolutionary radiations in terrestrial ecosystems of the Mesozoic era (approximately 252–66 million years ago), but their origin has remained an unresolved enigma in palaeontology since the nineteenth century2–4. These flying reptiles have been hypothesized to be the close relatives of a wide variety of reptilian clades, including dinosaur relatives2–8, and there is still a major morphological gap between those forms and the oldest, unambiguous pterosaurs from the Upper Triassic series. Here, using recent discoveries of well-preserved cranial remains, microcomputed tomography scans of fragile skull bones (jaws, skull roofs and braincases) and reliably associated postcrania, we demonstrate that lagerpetids—a group of cursorial, non-volant dinosaur precursors—are the sister group of pterosaurs, sharing numerous synapomorphies across the entire skeleton. This finding substantially shortens the temporal and morphological gap between the oldest pterosaurs and their closest relatives and simultaneously strengthens the evidence that pterosaurs belong to the avian line of archosaurs. Neuroanatomical features related to the enhanced sensory abilities of pterosaurs9 are already present in lagerpetids, which indicates that these features evolved before flight. Our evidence illuminates the first steps of the assembly of the pterosaur body plan, whose conquest of aerial space represents a remarkable morphofunctional innovation in vertebrate evolution. Lagerpetids, bipedal archosaurs that are thought to be related to dinosaurs, are instead a sister group to pterosaurs, and although they have no obvious flight adaptations they share numerous synapomorphies with pterosaurs across the entire skeleton.

Focal adhesion kinase (FAK) integrates signaling from integrins, growth factor receptors and mechanical stress to control cell adhesion, motility, survival and proliferation. Here, we developed a single-component, photo-activatable FAK, termed optoFAK, by using blue light-induced oligomerization of cryptochrome 2 (CRY2) to activate FAK-CRY2 fusion proteins. OptoFAK functions uncoupled from physiological stimuli and activates downstream signaling rapidly and reversibly upon blue light exposure. OptoFAK stimulates SRC creating a positive feedback loop on FAK activation, facilitating phosphorylation of paxillin and p130Cas in adherent cells. In detached cells or in mechanically stressed adherent cells, optoFAK is autophosphorylated upon exposure to blue light, however, downstream signaling is hampered indicating that the accessibility to these substrates is disturbed. OptoFAK may prove to be a useful tool to study the biological function of FAK in growth factor and integrin signaling, tension-mediated focal adhesion maturation or anoikis and could additionally serve as test system for kinase inhibitors.

The evolution of the braincase and brain of early pseudosuchians through to the earliest crocodylomorphs is poorly understood given the paucity of specimens, lack of well-preserved material, and lack of consensus on the phylogenetic relationships of the major clades of Pseudosuchia. Here, we describe three differently sized braincases diagnosable as belonging to the archosaur Parringtonia gracilis from the Middle Triassic Manda Beds of Tanzania. One of them, a nearly complete and exceptionally preserved braincase, possesses a complex set of character states typically present in early-diverging Pseudosuchia, but it also possesses nearly all of the character states previously hypothesized to represent synapomorphies exclusive to aetosaurs and crocodylomorphs. Details of the internal anatomy and a reconstructed endocast were obtained through high-resolution computed tomographic data and show a high degree of conservation of brain architecture across Pseudosuchia, in stark contrast to the extensive shifts observed within Theropoda to crown Aves. Integrating cranial and postcranial data into a phylogenetic analysis, we find Parringtonia gracilis to be closely related to a Revueltosaurus-Aetosauria clade at the base of Suchia and distantly related to crocodylomorphs. No matter which pseudosuchian clade is the most closely related to crocodylomorphs, we deduce that homoplasy in the braincase appears to be common across Pseudosuchia.

The sacrum – consisting of those vertebrae that articulate with the ilia – is the exclusive skeletal connection between the hindlimbs and axial skeleton in tetrapods. Therefore, the morphology of this portion of the vertebral column plays a major role in the evolution of terrestrial locomotion. Whereas most extant reptiles only possess the two plesiomorphic sacral vertebrae, additional vertebrae have been incorporated into the sacrum multiple times independently among early‐diverging archosaurian (crocodylians + birds) clades. Phytosauria was a diverse, abundant, and cosmopolitan clade of archosauriforms throughout the Late Triassic, but postcrania of this clade are rarely described and few species‐level taxonomic placements of phytosaurian postcranial material are available, potentially hampering knowledge of morphological disparity in the postcranial skeleton among phytosaurs. Here, we describe the sacrum of Smilosuchus adamanensis, a phytosaur recovered from the Upper Triassic Chinle Formation of Arizona. This sacrum consists of the two primordial sacral vertebrae, but has a vertebra incorporated from the trunk into the sacrum (= a dorsosacral) and is therefore the first Late Triassic phytosaur and one of the first non‐archosaurian archosauromorphs to be described with more than two sacral vertebrae. Our interpretation of this element as a dorsosacral is justified by the lateral extent of the dorsosacral ribs, clear surfaces of articulation between the distal ends of the dorsosacral ribs and the first primordial sacral ribs, and the scar on the medial surface of each ilium for articulation with each dorsosacral rib. Additionally, we provide the first detailed description of the vertebral junction formed by two anteriorly projecting flanges on the first primordial sacral ribs and their corresponding facets on the centrum of the dorsosacral. Computed tomographic (CT) imaging reveals that the two primordial sacrals are not co‐ossified and that the dorsosacral morphology of this specimen is not the result of obvious pathology. We place this incorporation of a trunk vertebra into the phytosaurian sacrum in a broader evolutionary context, with this shift in vertebral identity occurring at least seven times independently among Triassic archosauriforms, including at least three times in early crocodylian‐line archosaurs and at least four times among bird‐line archosaurs. Additionally, anteriorly projecting flanges of sacral ribs which articulate with the anterior‐adjacent centrum have evolved several times in archosauriforms, and we interpret ‘shared’ sacral ribs (= a sacral rib that articulates with two adjacent sacral centra more or less equally) present in some archosaurian clades as a more extreme example of this morphology. In extant taxa the highly conserved Hox gene family plays a central role in the patterning of the axial skeleton, especially vertebral identity; therefore, the independent incorporation of a trunk vertebra into the sacrum across multiple archosauriform lineages may suggest a homologous underlying developmental mechanism for this evolutionary trend.

A complex pelvic morphology has been discovered in the fossils of one of the largest crocodylians.

Crown-group frogs (Anura) originated over 200 Ma according to molecular phylogenetic analyses, though only a few fossils from high latitudes chronicle the first approximately 60 Myr of frog evolution and distribution. We report fossils that represent both the first Late Triassic and the earliest equatorial record of Salientia, the group that includes stem and crown-frogs. These small fossils consist of complete and partial ilia with anteriorly directed, elongate and distally hollow iliac blades. These features of these ilia, including the lack of a prominent dorsal protuberance and a shaft that is much longer than the acetabular region, suggest a closer affinity to crown-group Anura than to Early Triassic stem anurans Triadobatrachus from Madagascar and Czatkobatrachus from Poland, both high-latitude records. The new fossils demonstrate that crown anurans may have been present in the Late Triassic equatorial region of Pangea. Furthermore, the presence of Early Jurassic anurans in the same stratigraphic sequence ( Prosalirus bitis from the Kayenta Formation) suggests that anurans survived the climatic aridification of this region in the early Mesozoic. These fossils highlight the importance of the targeted collection of microfossils and provide further evidence for the presence of crown-group representatives of terrestrial vertebrates prior to the end-Triassic extinction.

This project was funded by the Virginia Tech Department of Geosciences and is supported by the National Science Foundation Graduate Research Fellowship Program, grant number DGE-1418060.

Summary Living archosaurs (birds and crocodylians) have disparate locomotor strategies that evolved since their divergence ∼250 mya. Little is known about the early evolution of the sensory structures that are coupled with these changes, mostly due to limited sampling of early fossils on key stem lineages. In particular, the morphology of the semicircular canals (SCCs) of the endosseous labyrinth has a long-hypothesized relationship with locomotion. Here, we analyze SCC shapes and sizes of living and extinct archosaurs encompassing diverse locomotor habits, including bipedal, semi-aquatic, and flying taxa. We test form-function hypotheses of the SCCs and chronicle their evolution during deep archosaurian divergences. We find that SCC shape is statistically associated with both flight and bipedalism. However, this shape variation is small and is more likely explained by changes in braincase geometry than by locomotor changes. We demonstrate high disparity of both shape and size among stem-archosaurs and a deep divergence of SCC morphologies at the bird–crocodylian split. Stem-crocodylians exhibit diverse morphologies, including aspects also present in birds and distinct from other reptiles. Therefore, extant crocodylian SCC morphologies do not reflect retention of a "primitive" reptilian condition. Key aspects of bird SCC morphology that hitherto were interpreted as flight related, including large SCC size and enhanced sensitivity, appeared early on the bird stem-lineage in non-flying dinosaur precursors. Taken together, our results indicate a deep divergence of SCC traits at the bird–crocodylian split and that living archosaurs evolved from an early radiation with high sensory diversity. Video abstract

Allokotosauria, including Trilophosauridae and Azendohsauridae, is an extinct archosauromorph group that reached a near-Pangean distribution in the Middle Triassic to Late Triassic and evolved a broad range of cranial and dental morphologies. Within the Chinle Formation of western North America, allokotosaurs span the Norian-aged Blue Mesa Member (Adamanian estimated holochronozone) and Sonsela Member (Adamanian and Revueltian estimated holochronozone). The Adamanian-Revueltian faunal assemblage transition is hypothesized to represent extinction and faunal turnover in large-bodied archosauromorphs, but poor sampling of known localities has obscured any similar patterns in tetrapods

The neotype skull of the Indian phytosaur Parasuchus hislopi Lydekker, 1885 (ISI R42) is re-evaluated and compared with the type material of other basal phytosaurs. Parasuchus hislopi is extremely similar to species previously placed in Paleorhinus (P. bransoni and P. angustifrons), sharing with them such characters as a series of nodes on the lateral surface of the jugal, paired ridges on the squamosal and a frontal depression. Parasuchus hislopi represents a valid species: it can be distinguished from P. bransoni by a relatively low narial eminence and P. angustifrons by the absence of paired nasal depressions. Inclusion of Parasuchus hislopi in a phylogenetic analysis of phytosaurs recovers it in a well-supported clade with P. bransoni and P. angustifrons. Parasuchus is considered the senior synonym of Paleorhinus and Arganarhinus. Parasuchus (here considered to include P. hislopi, P. angustifrons, P. bransoni and P. magnoculus) has a broad circum-Pangaean distribution, with species occurring in the south-western United States, Morocco, central Europe and India. Phytosaur higher-level taxonomy is also revised: Parasuchidae is redefined to include ‘Paleorhinus-grade’ phytosaurs and the later-diverging Mystriosuchinae (the group formerly known as Phytosauridae), and Pseudopalatinae is renamed Mystriosuchini for reason of priority.

Following the end-Permian extinction, terrestrial vertebrate diversity recovered by the Middle Triassic, and that diversity was now dominated by reptiles. However, those reptilian clades, including archosaurs and their closest relatives, are not commonly found until ~30 million years post-extinction in Late Triassic deposits despite time-calibrated phylogenetic analyses predicting an Early Triassic divergence for those clades. One of these groups from the Late Triassic, Phytosauria, is well known from a near-Pangean distribution, and this easily recognized clade bears an elongated rostrum with posteriorly retracted nares and numerous postcranial synapomorphies that are unique compared with all other contemporary reptiles. Here, we recognize the exquisitely preserved, nearly complete skeleton of Diandongosuchus fuyuanensis from the Middle Triassic of China as the oldest and basalmost phytosaur. The Middle Triassic age and lack of the characteristically-elongated rostrum fill a critical morphological and temporal gap in phytosaur evolution, indicating that the characteristic elongated rostrum of phytosaurs appeared subsequent to cranial and postcranial modifications associated with enhanced prey capture, predating that general trend of morphological evolution observed within Crocodyliformes. Additionally, Diandongosuchus supports that the clade was present across Pangea, suggesting early ecosystem exploration for Archosauriformes through nearshore environments and leading to ease of dispersal across the Tethys.

The archosaur species Teleocrater rhadinus, part of the new clade Aphanosauria, is an example of the earliest divergence of the avian stem lineage (Avemetatarsalia), the lineage that contains dinosaurs (including birds). The early history of the bird-line archosaurs, a group including dinosaurs, birds and pterosaurs, but excluding crocodilians, is not well defined. This is due in part to a fragmentary fossil record, but the distinctive morphology of pterosaurs has also obscured their ancestry. Sterling Nesbitt and colleagues describe a new species, Teleocraterrhadinus, from the Middle Triassic of Tanzania, that represents the most primitive known member of the bird-line archosaurs. Teleocrater provides the best guide so far to the ancestral bird-line condition. It was a lightly built, quadrupedal carnivore, so more like a crocodile than the small bipeds often depicted at this point in archosaur evolution. These are long-awaited findings on Teleocrater, which was undergoing study by the late Alan Charig of the Natural History Museum in London, and remained unpublished on his death in 1997. The relationship between dinosaurs and other reptiles is well established1,2,3,4, but the sequence of acquisition of dinosaurian features has been obscured by the scarcity of fossils with transitional morphologies. The closest extinct relatives of dinosaurs either have highly derived morphologies5,6,7 or are known from poorly preserved8,9 or incomplete material10,11. Here we describe one of the stratigraphically lowest and phylogenetically earliest members of the avian stem lineage (Avemetatarsalia), Teleocrater rhadinus gen. et sp. nov., from the Middle Triassic epoch. The anatomy of T. rhadinus provides key information that unites several enigmatic taxa from across Pangaea into a previously unrecognized clade, Aphanosauria. This clade is the sister taxon of Ornithodira (pterosaurs and birds) and shortens the ghost lineage inferred at the base of Avemetatarsalia. We demonstrate that several anatomical features long thought to characterize Dinosauria and dinosauriforms evolved much earlier, soon after the bird–crocodylian split, and that the earliest avemetatarsalians retained the crocodylian-like ankle morphology and hindlimb proportions of stem archosaurs and early pseudosuchians. Early avemetatarsalians were substantially more species-rich, widely geographically distributed and morphologically diverse than previously recognized. Moreover, several early dinosauromorphs that were previously used as models to understand dinosaur origins may represent specialized forms rather than the ancestral avemetatarsalian morphology.

The Triassic Period (252–201.5 Ma) records a great expansion of saurian diversity and disparity, particularly in skull morphology. Stem archosaurs exhibit substantial cranial disparity, especially ...

The shift from life in the water to life on land is regarded as a key transition in vertebrate evolution. This transition in tetrapodomorph vertebrates is marked by a number of morphological shifts...

Phytosaurs are a diverse and morphologically distinctive clade of superficially crocodile-like archosauriforms that had a near global distribution during the Late Triassic. Because their remains are among the most abundant vertebrate remains recovered in many Upper Triassic terrestrial formations, phytosaurs are used extensively in long-range biochronological and biostratigraphic correlations. The biochronologically oldest and earliest branching known phytosaurs include an array of nominal species from the early Late Triassic of the United States, Germany, Poland, Morocco, and India that have been synonymized within the genus Paleorhinus, and subsequently used to define a global ‘Paleorhinus biochron’. However, recent phylogenetic work suggested that the North American species previously referred to Paleorhinus are paraphyletic. Here, we reassess the systematics and anatomy of putative specimens of Paleorhinus from southern Germany. Two well-preserved basal phytosaur skulls from the Blasensandstein (Carnian) of Bavaria form the holotypes of Francosuchus angustifrons and Ebrachosuchus neukami, both of which were synonymized with Paleorhinus by previous workers. We demonstrate that Francosuchus angustifrons shares unique synapomorphies with specimens referred to Paleorhinus bransoni from the Late Triassic of Texas, and thus refer the species to Paleorhinus. By contrast, the longirostrine Ebrachosuchus is highly distinctive in morphology, and our new cladistic analysis of Phytosauria demonstrates that it represents a valid taxon that is more closely related to Phytosauridae than to Paleorhinus. We provide the first autapomorphy-based support for a monophyletic but restricted Paleorhinus (supported by a nodal row on the jugal, and low paired ridges on the squamosal) and confirm that previous broader conceptions of Paleorhinus are likely to be paraphyletic. © 2013 The Linnean Society of London

Reptiles have a long history of transitioning from terrestrial to semi-aquatic or aquatic environments that stretches back at least 250 million years. Within Archosauria, both living crocodylians and birds have semi-aquatic members. Closer to the root of Archosauria and within the closest relatives of the clade, there is a growing body of evidence that early members of those clades had a semi-aquatic lifestyle. However, the morphological adaptations to a semi-aquatic environment remain equivocal in most cases. Here, we introduce a new Middle Triassic (245-235 Ma) archosauriform, Litorosuchus somnii, gen. et sp. nov., based on a nearly complete skeleton from the Zhuganpo Member (Ladinian [241-235 Ma]) of the Falang Formation, Yunnan, China. Our phylogenetic analyses suggest that Litorosuchus is a stem archosaur closely related to the aberrant Vancleavea just outside of Archosauria. The well-preserved skeleton of L. somnii bears a number of morphological characters consistent with other aquatic-adapted tetrapods including: a dorsally directed external naris, tall neural spines and elongate chevrons in an elongated tail, a short and broad scapula, webbed feet, long cervical vertebrae with long slender ribs, and an elongated rostrum with long and pointed teeth. Together these features represent one of the best-supported cases of a semi-aquatic mode of life for a stem archosaur. Together with Vancleavea campi, the discovery of L. somnii demonstrates a growing body of evidence that there was much more diversity in mode of life outside Archosauria. Furthermore, L. somnii helps interpret other possible character states consistent with a semi-aquatic mode of life for archosauriforms, including archosaurs.

Summary Similarities in body plan evolution, such as wings in pterosaurs, birds, and bats or limblessness in snakes and amphisbaenians, can be recognized as classical examples of convergence among animals [1–3]. We introduce a new Triassic stem archosaur that is unexpectedly and remarkably convergent with the "dome-headed" pachycephalosaur dinosaurs that lived over 100 million years later. Surprisingly, numerous additional taxa in the same assemblage (the Otis Chalk assemblage from the Dockum Group of Texas) demonstrate the early acquisition of morphological novelties that were later convergently evolved by post-Triassic dinosaurs. As one of the most successful clades of terrestrial vertebrates, dinosaurs came to occupy an extensive morphospace throughout their diversification in the Mesozoic Era [4, 5], but their distant relatives were first to evolve many of those "dinosaurian" body plans in the Triassic Period [6–8]. Our analysis of convergence between archosauromorphs from the Triassic Period and post-Triassic archosaurs demonstrates the early and extensive exploration of morphospace captured in a single Late Triassic assemblage, and we hypothesize that many of the "novel" morphotypes interpreted to occur among archosaurs later in the Mesozoic already were in place during the initial Triassic archosauromorph, largely non-dinosaurian, radiation and only later convergently evolved in diverse dinosaurian lineages.

The Post Quarry, within the lower part of the type section of the Upper Triassic Cooper Canyon Formation in southern Garza County, western Texas, contains a remarkably diverse vertebrate assemblage. The Post Quarry has produced: the small temnospondylRileymillerus cosgriffi; the metoposauridApachesaurus gregorii; possible dicynodonts and eucynodonts; a clevosaurid sphenodontian; non-archosauriform archosauromorphs (Trilophosaurus dornorum, simiosaurians, and possiblyMalerisaurus); the phytosaurLeptosuchus; several aetosaurs (Calyptosuchus wellesi,Typothorax coccinarum,Paratypothorax, andDesmatosuchus smalli); the poposauroidShuvosaurus inexpectatus(“Chatterjeea elegans”); the rauisuchidPostosuchus kirkpatricki; an early crocodylomorph; several dinosauromorphs (the lagerpetidDromomeron gregorii, the silesauridTechnosaurus smalli, a herrerasaurid, and an early neotheropod); and several enigmatic small diapsids. Revised lithostratigraphic correlations of the lower Cooper Canyon Formation with the Tecovas Formation, the occurrence ofLeptosuchus, and the overall composition of the assemblage indicate that the Post Quarry falls within the Adamanian biozone, and not the Revueltian biozone. Stratigraphic subdivision of the Adamanian biozone may be possible, and the Post Quarry may be correlative with the upper part of the Adamanian biozone in Arizona. The age of the Post Quarry assemblage is possibly late Lacian or earliest Alaunian (late early Norian or earliest middle Norian), between 220 and 215 Ma.

The paraphyletic genus ‘Paleorhinus’ is understood currently as a cosmopolitan phytosaur taxon from the Late Triassic. There is no consensus regarding the number of species of ‘Paleorhinus,’ with multiple species and genera synonymised into a single genus or even a single species at various points in its published history. The taxonomy is confounded by historical descriptions without the benefit of comparisons to more recently collected specimens, emphasis on plesiomorphic cranial morphology as diagnostic features of the genus, and lack of cladistic analyses. When included in a recent explicitly cladistic phylogenetic analysis, the holotype of ‘Paleorhinus’ scurriensis (TTU P-00539) was found to be the earliest-branching phytosaur with respect to other North American specimens previously referred to ‘Paleorhinus,’ and is generically distinct from Paleorhinus. ‘Paleorhinus’ scurriensis differs from all known phytosaurs in five unambiguous characters: basitubera widely separated mediolaterally; ridge present on lateral surface of jugal; thickened shelf present along posteroventral edge of expanded pterygoid-quadrate wing; ‘septomaxillae’ separated and excluded from internarial septum; and nasal swelling present posterior to posterior borders of nares. This detailed morphological description of an early-branching phytosaur taxon is a first step towards resolving long-standing issues surrounding specific anatomical features and relationships among early members of the clade.

A new phytosaur taxon from Petrified Forest National Park, Arizona, is here described based on cranial material from a single individual. This specimen previously was included in an extensive phylogenetic analysis, and it was found to possess a combination of character states that differs from all known phytosaur taxa in addition to two autapomorphies within the braincase and an autapomorphy of the mandible. The new taxon adds to the taxonomic diversity recognized from the Sonsela Member of the Chinle Formation. The continued increase in phytosaur diversity emphasizes the need to more accurately characterize and identify taxa within a phylogenetic systematic context in order to produce a more refined signal for biostratigraphic correlations, biochronologic inferences, and faunal dynamics during the Late Triassic.

Leptosuchus Case, 1922 (Reptilia: Phytosauria) from the Late Triassic of the American West is represented by many specimens. Here, I present complete morphological descriptions of the skull material of a new taxon from the Sonsela Member (Chinle Formation) of Petrified Forest National Park, Arizona, with the first rigorous phylogenetic analysis focused on the interrelationships of Leptosuchus. The new taxon is recovered as the sister taxon to Pseudopalatinae. It possesses one unambiguous synapomorphy (the ‘septomaxillae’ form part of the lateral borders of the nares) and shares the presence of a subsidiary opisthotic process with Pseudopalatinae. The new taxon does not fall within the restricted clade Leptosuchus. In my analysis, the previously proposed, but undemonstrated, sister taxon relationship between Angistorhinus and Rutiodon is not supported, Paleorhinus is recovered as paraphyletic, and a subset of taxa traditionally included within Leptosuchus are found to be more closely related to Pseudopalatinae, rendering Leptosuchus paraphyletic. ‘Leptosuchus’adamanensis emerges as sister taxon to Smilosuchus gregorii and is here referred to as Smilosuchus adamanensis nov. comb., and ‘Machaeroprosopus’lithodendrorum is also transferred to Smilosuchus lithodendrorum nov. comb. Documentation of the variation present within Phytosauria, and specifically within Leptosuchus sensu lato, demonstrates higher diversity within Phytosauria than previously appreciated and places the character states previously proposed for Pseudopalatinae into a broader context of shared characters.

The pathological findings are described in three cases of infectious canine hepatitis in free-ranging red foxes (Vulpes vulpes) in England. The foxes died after short periods of clinical illness. Mild jaundice and hepatic congestion were evident grossly. On histopathological examination, intranuclear inclusion bodies were visible in hepatocytes, in association with hepatocyte dissociation and necrosis, as well as in renal glomeruli, renal tubular epithelial cells and vascular endothelial cells. Canine adenovirus type 1 (CAV-1) was isolated from all three foxes. In a serological study, antibodies to CAV-1 were detected in tissue fluid extracts taken from 11 of 58 (19 per cent) frozen red fox carcases from England and Scotland.

Vancleavea campi Long & Murry, 1995, from the Late Triassic of western North America, represents the latest surviving non-archosaurian archosauriform known to date. We present here a detailed comparative description based on a nearly complete, articulated skeleton from the Coelophysis Quarry in north-central New Mexico and other fragmentary specimens. The unique combination of morphological features of Vancleavea is unparalleled within Reptilia; it has four unique morphologies of imbricated osteoderms covering the entire body, a short, highly ossified skull, relatively small limbs and morphological features consistent with a semi-aquatic lifestyle. Vancleavea is placed in a rigorous phylogenetic analysis examining the relationships of non-archosaurian archosauriforms, and is found to be more closely related to Archosauria than both Erythrosuchus and Proterosuchus, but outside of the crown group. The analysis confirms previously hypothesized relationships, which found Euparkeria to be the closest sister taxon of Archosauria. It is not clear whether specimens referred to Vancleavea campi represent a single species-level taxon or a clade of closely related taxa that lived through much of the Late Triassic of North America, given the poor fossil record of the taxon.

The Upper Triassic Canjilon Quarry in northern New Mexico preserves a vertebrate assemblage typical of the Norian Stage of North America. Although dominated in relative abundance by the phytosaur Pseudopalatus and the aetosaur Typothorax, a more diverse assemblage of smaller forms was previously reported. Additions to the Canjilon Quarry vertebrate assemblage are critically reviewed and described using an apomorphy-based method for assigning taxa. An apomorphy-based approach for compiling and comparing assemblages allows each taxonomic assignment to be a testable hypothesis. Fragmentary yet diagnostic elements that may not be assignable to a species-level taxon can be assigned to a larger clade and thus provide useful information. Additionally, each taxon listed in the assemblage must be tied to a diagnostic specimen listed with a museum specimen number. Using this method for the Canjilon Quarry, we establish the presence of an assemblage that is more similar to other vertebrate quarries near Gho...

Dual control of cellular heme levels by extracellular scavenger proteins and degradation by heme oxygenases is essential in diseases associated with increased heme release. During severe hemolysis or rhabdomyolysis, uncontrolled heme exposure can cause acute kidney injury and endothelial cell damage. The toxicity of heme was primarily attributed to its pro-oxidant effects; however additional mechanisms of heme toxicity have not been studied systematically. In addition to redox reactivity, heme may adversely alter cellular functions by binding to essential proteins and impairing their function. We studied inducible heme oxygenase (Hmox1)-deficient mouse embryo fibroblast cell lines as a model to systematically explore adaptive and disruptive responses that were triggered by intracellular heme levels exceeding the homeostatic range. We extensively characterized the proteome phenotype of the cellular heme stress responses by quantitative mass spectrometry of stable isotope-labeled cells that covered more than 2000 individual proteins. The most significant signals specific to heme toxicity were consistent with oxidative stress and impaired protein degradation by the proteasome. This ultimately led to an activation of the response to unfolded proteins. These observations were explained mechanistically by demonstrating binding of heme to the proteasome that was linked to impaired proteasome function. Oxidative heme reactions and proteasome inhibition could be differentiated as synergistic activities of the porphyrin. Based on the present data a novel model of cellular heme toxicity is proposed, whereby proteasome inhibition by heme sustains a cycle of oxidative stress, protein modification, accumulation of damaged proteins and cell death., Cell Death and Differentiation, 22 (4), ISSN:1350-9047, ISSN:1476-5403

Hypotheses of feeding behaviors and community structure are testable with rare direct evidence of trophic interactions in the fossil record (e.g., bite marks). We present evidence of four predation, scavenging, and/or interspecific fighting events involving two large paracrocodylomorphs (=‘rauisuchians’) from the Upper Triassic Chinle Formation (∼220–210 Ma). The larger femur preserves a rare history of interactions with multiple actors prior to and after death of this ∼8–9-m individual. A large embedded tooth crown and punctures, all of which display reaction tissue formed through healing, record evidence of a failed attack on this individual. The second paracrocodylomorph femur exhibits four unhealed bite marks, indicating the animal either did not survive the attack or was scavenged soon after death. The combination of character states observed (e.g., morphology of the embedded tooth, ‘D’-shaped punctures, evidence of bicarination of the marking teeth, spacing of potentially serial marks) indicates that large phytosaurs were actors in both cases. Our analysis of these specimens demonstrates phytosaurs targeted large paracrocodylomorphs in these Late Triassic ecosystems. Previous distinctions between ‘aquatic’ and ‘terrestrial’ Late Triassic trophic structures were overly simplistic and built upon mistaken paleoecological assumptions; we show they were intimately connected at the highest trophic levels. Our data also support that size cannot be the sole factor in determining trophic status. Furthermore, these marks provide an opportunity to start exploring the seemingly unbalanced terrestrial ecosystems from the Late Triassic of North America, in which large carnivores far outnumber herbivores in terms of both abundance and diversity.

1. Instantaneous leaf photosynthesis and land area-based net ecosystem CO2 exchange (NEC) are almost universally increased at elevated CO2 concentrations, at least in the short term and under high light conditions. This raises the possibility that terrestrial ecosystems sequester extra C in response to elevated CO2, and it has been hypothesized that part of this extra C is stored in soils. 2. Attempts to quantify ecosystem C sequestration experimentally are based on (i) ecosystem CO2 exchange measurements; (ii) C isotope tracking; and (iii) direct C stock measurements. 3. Because direct C stock measurements are insensitive to increases in ecosystem C storage in the range expected, and C isotope tracking is methodologically difficult because of the need to account for new and old soil C pools, NEC measurements were considered to be a more direct and unbiased method to estimate net ecosystem production and C sequestration at elevated CO2. 4. Here we present a case study in calcareous grassland under long-term CO2 enrichment in which we demonstrate that calculated C balances are extremely sensitive to systematic experimental biases inherent in any CO2 flux-based study at elevated CO2. A sensitivity analysis demonstrates that these systematic errors tend to result in severe overestimation of ecosystem C accretion under elevated CO2. 5. Carbon isotope data and soil pool C and N measurements from the same study add to the evidence that the C balance derived from CO2 flux measurements at elevated CO2 is overestimated. 6. Based on this evidence, and the wide attention given to C sequestration at elevated CO2, we suggest a critical reconsideration of the appropriateness of NEC measurement in CO2-enriched ecosystems as a basis for calculating ecosystem C balances at elevated CO2.

As part of a long-term study of the effects of elevated CO 2 on biodiversity and ecosystem function in a calcareous grassland, we measured ecosystem carbon dioxide and water-vapour fluxes over 24-h periods during the 1994 and 1995 growing seasons. Data were used to derive CO 2 and H 2 O gas-exchange response functions to quantum flux density (QFD). The relative increase in net ecosystem CO 2 flux (NEC) owing to CO 2 enrichment increased as QFD rose. Daytime NEC at high QFD under elevated CO 2 increased by 25% to 60%, with the greatest increases in the spring and after mowing in June when above-ground biomass was lowest. There was much less stimulation of NEC in early June and again in October when the canopy was fully developed. Night-time NEC was not significantly altered under elevated CO 2 . Short-term reversal of CO 2 concentrations between treatments after two seasons of CO 2 exposure provided evidence for a 50% downward adjustment of NEC expressed per unit above-ground plant dry weight. However, when expressed on a land area basis, this difference disappeared because of a c. 20% increase in above-ground biomass under elevated CO 2 . Ecosystem evapotranspiration (ET) was not significantly altered by elevated CO 2 when averaged over all measurement dates and positions. However, ET was reduced 3-18% at high QFD in plots at the top of the slope at our study site. In summary, CO 2 enrichment resulted in a large stimulation of ecosystem CO 2 capture, especially during periods of a large demand of carbon in relationship to its supply, and resulted in a relatively small and variable effect on ecosystem water consumption.

Paraplegia affected 14 hedgehogs (Erinaceus europaeus) in a wildlife rescue hospital over a period of six months. Postmortem examination revealed demyelination in the brain and spinal cord and an inflammatory response in the meninges, choroid plexus and CNS. The peripheral nervous system was not affected. In the spleen, lungs and liver there was an accumulation of megakaryocytes and other evidence of extramedullary haemopoiesis, but there was no haematological evidence of anaemia. The pattern of disease incidence and the nature of the changes in the CNS suggest they were of viral origin, but no causal agent was isolated and the possibility of a neurotoxin cause cannot be ruled out.

The mechanism underlying demyelination in inflammatory canine distemper encephalitis is uncertain. Macrophages and their secretory products are thought to play an important effector role in this lesion. Recently, we have shown that anti-canine distemper virus antibodies, known to occur in chronic inflammatory lesions, stimulate macrophages leading to the secretion of reactive oxygen species (ROS). To investigate whether ROS could be involved in demyelination, dog glial cell cultures were exposed to xanthine/xanthine oxidase (X/XO), a system capable of generating O2-. This treatment resulted in a specific time-dependent degeneration and loss of oligodendrocytes, the myelin producing cells of the central nervous system. Initial degeneration was not associated with a decrease in viability of oligodendrocytes as judged by trypan blue and propidium iodide exclusion. Astrocytes and brain macrophages were not affected morphologically by this treatment. Further, an evaluation of the effect of several ROS scavengers, transition metal chelators and inhibitors of poly (ADP-ribose) polymerase suggests that a metal dependent formation of .OH or a similar highly oxidizing species could be responsible for the observed selective damage to oligodendrocytes.