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  1. Abstract Living amphibians (Lissamphibia) include frogs and salamanders (Batrachia) and the limbless worm-like caecilians (Gymnophiona). The estimated Palaeozoic era gymnophionan–batrachian molecular divergence 1 suggests a major gap in the record of crown lissamphibians prior to their earliest fossil occurrences in the Triassic period 2–6 . Recent studies find a monophyletic Batrachia within dissorophoid temnospondyls 7–10 , but the absence of pre-Jurassic period caecilian fossils 11,12 has made their relationships to batrachians and affinities to Palaeozoic tetrapods controversial 1,8,13,14 . Here we report the geologically oldest stem caecilian—a crown lissamphibian from the Late Triassic epoch of Arizona, USA—extending the caecilian record by around 35 million years. These fossils illuminate the tempo and mode of early caecilian morphological and functional evolution, demonstrating a delayed acquisition of musculoskeletal features associated with fossoriality in living caecilians, including the dual jaw closure mechanism 15,16 , reduced orbits 17 and the tentacular organ 18 . The provenance of these fossils suggests a Pangaean equatorial origin for caecilians, implying that living caecilian biogeography reflects conserved aspects of caecilian function and physiology 19 , in combination with vicariance patterns driven by plate tectonics 20 . These fossils reveal a combination of features that is unique to caecilians alongside features that are shared with batrachian and dissorophoid temnospondyls, providing new and compelling evidence supporting a single origin of living amphibians within dissorophoid temnospondyls. 
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  2. Abstract Non-archosaur archosauromorphs are a paraphyletic group of diapsid reptiles that were important members of global Middle and Late Triassic continental ecosystems. Included in this group are the azendohsaurids, a clade of allokotosaurians (kuehneosaurids and Azendohsauridae + Trilophosauridae) that retain the plesiomorphic archosauromorph postcranial body plan but evolved disparate cranial features that converge on later dinosaurian anatomy, including sauropodomorph-like marginal dentition and ceratopsian-like postorbital horns. Here we describe a new malerisaurine azendohsaurid from two monodominant bonebeds in the Blue Mesa Member, Chinle Formation (Late Triassic, ca. 218–220 Ma); the first occurs at Petrified Forest National Park and preserves a minimum of eight individuals of varying sizes, and the second occurs near St. Johns, Arizona. Puercosuchus traverorum n. gen. n. sp. is a carnivorous malerisaurine that is closely related to Malerisaurus robinsonae from the Maleri Formation of India and to Malerisaurus langstoni from the Dockum Group of western Texas. Dentigerous elements from Puercosuchus traverorum n. gen. n. sp. confirm that some Late Triassic tooth morphotypes thought to represent early dinosaurs cannot be differentiated from, and likely pertain to, Puercosuchus -like malerisaurine taxa. These bonebeds from northern Arizona support the hypothesis that non-archosauriform archosauromorphs were locally diverse near the middle Norian and experienced an extinction event prior to the end-Triassic mass extinction coincidental with the Adamanian-Revueltian boundary recognized at Petrified Forest National Park. The relatively late age of this early-diverging taxon (Norian) suggests that the diversity of azendohsaurids is underrepresented in Middle and Late Triassic fossil records around the world. UUID: http://zoobank.org/e6eeefd2-a0ae-47fc-8604-9f45af8c1147 . 
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  3. Abstract

    The anatomy of the braincase and associated soft tissues of the lagerpetidDromomeron gregorii(Archosauria: Avemetatarsalia) from the Late Triassic of the United States is here described. This corresponds to the first detailed description of cranial materials of Lagerpetidae, an enigmatic group of Late Triassic (c. 236–200 Million years ago) animals that are the closest known relatives of pterosaurs, the flying reptiles. The braincase ofD. gregoriiis characterized by the presence of an anteriorly elongated laterosphenoid and a postparietal, features observed in stem‐archosaurs but that were still unknown in early members of the avian lineage of archosaurs. Using micro‐computed tomography (CT‐scan data), we present digital reconstructions of the brain and endosseous labyrinth ofD. gregorii. The brain ofD. gregoriiexhibits a floccular lobe of the cerebellum that projects within the space of the semicircular canals. The semicircular canals are relatively large when compared to other archosauromorphs, with the anterior canal exhibiting a circular shape. These features of the sensory structures ofD. gregoriiare more similar to those of pterosaurs than to those of other early avemetatarsalians. In sum, the braincase anatomy ofD. gregoriishows a combination of plesiomorphic and apomorphic features in the phylogenetic context of Archosauria and suggests that the still poorly understood early evolution of the braincase in avemetatarsalians is complex, with a scenario of independent acquisitions and losses of character states.

     
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  4. Abstract 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° 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 ~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. 
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  5. Abstract

    A snake‐like body plan and burrowing lifestyle characterize numerous vertebrate groups as a result of convergent evolution. One such group is the amphisbaenians, a clade of limbless, fossorial lizards that exhibit head‐first burrowing behavior. Correlated with this behavior, amphisbaenian skulls are more rigid and coossified than those of nonburrowing lizards. However, due to their lifestyle, there are many gaps in our understanding of amphisbaenian anatomy, including how their cranial osteology varies among individuals of the same species and what that reveals about constraints on the skull morphology of head‐first burrowing taxa. We investigated intraspecific variation in the cranial osteology of amphisbaenians using seven individuals of the trogonophidDiplometopon zarudnyi. Variation in both skull and individual skull element morphology was examined qualitatively and quantitatively through three‐dimensional (3D) models created from microcomputed tomography data. Qualitative examination revealed differences in the number and position of foramina, the interdigitation between the frontals and parietal, and the extent of coossification among the occipital complex, fused basioccipital and parabasisphenoid (“parabasisphenoid‐basioccipital complex”), and elements X. We performed 3D landmark‐based geometric morphometrics for the quantitative assessment, revealing shape differences in the skull, premaxilla, maxilla, frontal, and parietal. The observed intraspecific variation may be the result of different stages of ontogenetic development or biomechanical optimization for head‐first burrowing. For example, variation in the coossification of the occipital region suggests a potential ontogenetic coossification sequence. Examination of these areas of variation across other head‐first burrowing taxa will help determine if the variation is clade‐specific or part of a broader macroevolutionary pattern of head‐first burrowing.

     
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  6. Abstract

    The femora of diapsids have undergone morphological changes related to shifts in postural and locomotor modes, such as the transition from plesiomorphic amniote and diapsid taxa to the apomorphic conditions related to a more erect posture within Archosauriformes. One remarkable clade of Triassic diapsids is the chameleon‐like Drepanosauromorpha. This group is known from numerous articulated but heavily compressed skeletons that have the potential to further inform early reptile femoral evolution. For the first time, we describe the three‐dimensional osteology of the femora of Drepanosauromorpha, based on undistorted fossils from the Upper Triassic Chinle Formation and Dockum Group of North America. We identify apomorphies and a combination of character states that link these femora to those in crushed specimens of drepanosauromorphs and compare our sample with a range of amniote taxa. Several characteristics of drepanosauromorph femora, including a hemispherical proximal articular surface, prominent asymmetry in the proximodistal length of the tibial condyles, and a deep intercondylar sulcus, are plesiomorphies shared with early diapsids. The femora contrast with those of most diapsids in lacking a crest‐like, distally tapering internal trochanter. They bear a ventrolaterally positioned tuberosity on the femoral shaft, resembling the fourth trochanter in Archosauriformes. The reduction of an internal trochanter parallels independent reductions in therapsids and archosauriforms. The presence of a ventrolaterally positioned trochanter is also similar to that of chameleonid squamates. Collectively, these features demonstrate a unique femoral morphology for drepanosauromorphs, and suggest an increased capacity for femoral adduction and protraction relative to most other Permo‐Triassic diapsids.

     
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  7. null (Ed.)
    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. 
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  8. Abstract

    Archosauromorph reptiles underwent rapid lineage diversification, increases in morphological and body size disparity, and expansion into new adaptive landscapes. Several of the primary early archosauromorph clades (e.g. rhynchosaurs) are easy to differentiate from others because of their characteristic body types, whereas the more lizard‐like and carnivorous forms with long necks (e.g. tanystropheids) were historically all relegated to the groups Protorosauria or Prolacertiformes. However, it is now clear that these groups are polyphyletic and that a lizard‐like, carnivorous form is plesiomorphic for Archosauromorpha, and multiple subclades started with that body plan. Among these early forms isMalerisaurusfrom the Upper Triassic of India (M. robinsonae) and the Upper Triassic of south‐western USA (M. langstoni). In this paper, we critically re‐evaluate the genus. We find both species ofMalerisaurusas valid, and identifyMalerisaurusas an early diverging, but late‐surviving, carnivorous member of Azendohsauridae within Allokotosauria. Our histological analysis and assessment of ontogenetic changes of limb bones of small and large individuals demonstrate that the skeletons of the small forms grew slowly and became more robust through ontogeny, and that the larger recovered bones are at or near the maximum size of the taxon.MalerisaurusandMalerisaurus‐like taxa were common members of the Otischalkian–Adamanian (late Carnian to mid‐Norian) faunal assemblages from Upper Triassic strata of the south western USA, but they are absent from the younger Revueltian holochronozone. Specimens from western North America show that Allokotosauria had a near‐Pangaean distribution for much of the Middle Triassic to Late Triassic.

     
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