Volcanically driven lacustrine ecosystem changes during the Carnian Pluvial Episode (Late Triassic).

The Late Triassic Carnian Pluvial Episode (CPE) saw a dramatic increase in global humidity and temperature that has been linked to the large-scale volcanism of the Wrangellia large igneous province. The climatic changes coincide with a major biological turnover on land that included the ascent of the dinosaurs and the origin of modern conifers. However, linking the disparate cause and effects of the CPE has yet to be achieved because of the lack of a detailed terrestrial record of these events. Here, we present a multidisciplinary record of volcanism and environmental change from an expanded Carnian lake succession of the Jiyuan Basin, North China. New U-Pb zircon dating, high-resolution chemostratigraphy, and palynological and sedimentological data reveal that terrestrial conditions in the region were in remarkable lockstep with the large-scale volcanism. Using the sedimentary mercury record as a proxy for eruptions reveals four discrete episodes during the CPE interval (ca. 234.0 to 232.4 Ma). Each eruptive phase correlated with large, negative C isotope excursions and major climatic changes to more humid conditions (marked by increased importance of hygrophytic plants), lake expansion, and eutrophication. Our results show that large igneous province eruptions can occur in multiple, discrete pulses, rather than showing a simple acme-and-decline history, and demonstrate their powerful ability to alter the global C cycle, cause climate change, and drive macroevolution, at least in the Triassic.

Sacral co-ossification in dinosaurs: The oldest record of fused sacral vertebrae in Dinosauria and the diversity of sacral co-ossification patterns in the group.

The fusion of the sacrum occurs in the major dinosaur lineages, i.e. ornithischians, theropods, and sauropodomorphs, but it is unclear if this trait is a common ancestral condition, or if it evolved independently in each lineage, or even how or if it is related to ontogeny. In addition, the order in which the different structures of the sacrum are fused, as well as the causes that lead to this co-ossification, are poorly understood. Herein, we described the oldest record of fused sacral vertebrae within dinosaurs, based on two primordial sacral vertebrae from the Late Triassic of Candelária Sequence, southern Brazil. We used computed microtomography (micro-CT) to analyze the extent of vertebral fusion, which revealed that it occurred only between the centra. We also assessed the occurrence of sacral fusion in Dinosauria and close relatives. The degree of fusion observed in representatives of the major dinosaur lineages suggested that there may be a sequential pattern of fusion of the elements of the sacrum, more clearly observed in Sauropodomorpha. Our analyses suggest that primordial sacral vertebrae fuse earlier in the lineage (as seen in Norian sauropodomorphs). Intervertebral fusion is observed to encompass progressively more vertebral units as sauropodomorphs evolve, reaching up to five or more fully fused sacrals in Neosauropoda. Furthermore, the new specimen described here indicates that the fusion of sacral elements occurred early in the evolution of dinosaurs. Factors such as ontogeny and the increase in body size, combined with the incorporation of vertebrae to the sacrum may have a significant role in the process and in the variation of sacral fusion observed.

New information on the mandibular anatomy of Agudotherium gassenae, a Late Triassic non-mammaliaform probainognathian from Brazil.

Agudotherium gassenae is a poorly known non-mammaliaform probainognathian cynodont from the Late Triassic of southern Brazil. It is known only by mandibular remains, and its affinities within Probainognathia are unclear. Furthermore, its phylogenetic affinities were never investigated through computational analyses. In this study, we described new lower jaw remains excavated from the type locality and performed the first phylogenetic investigation of this taxon. The new specimen provides further anatomical information. The rostral region of the lower jaw was poorly preserved in the type series, leading to the interpretation that A. gassenae had three lower incisors. The new specimen demonstrates the presence of four incisors. The phylogenetic analysis positioned A. gassenae as the sister group of Prozostrodontia. This hypothesis differs from that previously presented in the former description of the taxon, in which it was considered a non-mammaliaform prozostrodont by means of character-state comparisons.

An unusually robust specimen attributed to Buriolestes schultzi (Dinosauria: Sauropodomorpha) from the Late Triassic of southern Brazil.

Buriolestes schultzi is a small sauropodomorph dinosaur from Carnian beds (ca., 233 Ma) of southern Brazil. It is one of the earliest members of that lineage and is a key taxon to investigate the initial evolution of Sauropodomorpha. Here, we attribute a new specimen to B. schultzi from Late Triassic of southern Brazil, which represents the first occurrence of the taxon outside the type locality. The new specimen comprises a disarticulated and partial skeleton, including cranial and postcranial elements. It is tentatively regarded as an additional specimen of B. schultzi according to a unique combination of traits (including autapomorphies). Conversely, the new specimen is stouter than the other specimens of B. schultzi, as shown by femoral Robustness Index. Based on femoral circumference, the estimated body mass of the new specimen is approximately 15 kg, which is far higher than the previous estimations for other specimens of B. schultzi (i.e., approximately 7 kg). In fact, the new specimen and some specimens of Eoraptor lunensis and Saturnalia tupiniquim were found to be significantly stouter than coeval sauropodomorphs. Therefore, instead of all being constructed as gracile, the earliest sauropodomorphs experienced an unappreciated intraspecific variation in robustness. This is interesting because more precise data on species body mass are crucial in order to better understand the complex terrestrial ecosystems in which dinosaurs originated.

First record of rhynchosaurs (Archosauromorpha: Rhynchosauria: Hyperodapedontinae) from the early Late Triassic Santacruzodon Assemblage Zone of the Santa Maria Supersequence, Brazil.

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

Skull anatomy and paleoneurology of a new traversodontid from the Middle-Late Triassic of Brazil.

Traversodontidae, a clade of gomphodont cynodonts, thrived during the Middle and Late Triassic, displaying a wide geographical distribution. During fieldwork in 2009, a new specimen was discovered in Ladinian/early Carnian stratigraphic layers in southern Brazil. Here, we describe this specimen and propose a new taxon closely related to Traversodon stahleckeri (Traversodontinae) but displaying a unique combination of traits (e.g., presence of a poorly developed suborbital process, mesiodistal length of the paracanine fossa similar to the length of the canine, short diastema between the fourth incisor and the upper canine, and coronoid process not entirely covering the distalmost lower postcanine). Furthermore, the endocranial anatomy of the new taxon was examined. The reconstruction of the cranial endocast revealed paleoneurological features consistent with non-Gomphodontosuchinae traversodontids. These features include the presence of a pineal body (but the absence of an open parietal foramen). These recent findings contribute significantly to our understanding of the evolutionary history and cranial anatomy of Middle-Late Triassic traversodontids, shedding light on the diversity and adaptations of non-mammaliaform cynodonts.

A saurischian (Archosauria, Dinosauria) ilium from the Upper Triassic of southern Brazil and the rise of Herrerasauria.

The Carnian (Upper Triassic) rocks of the Candelária Sequence present a rich record of dinosaurs, including some of the oldest known dinosaurs worldwide. In this contribution we describe the first unequivocal dinosaur from the Pivetta site, located in the Restinga Sêca municipality, Southern Brazil. The specimen CAPPA/UFSM 0373 is an isolated but well-preserved left ilium. A thorough examination of the specimen's anatomy and a phylogenetic analysis provides evidence that CAPPA/UFSM 0373 belongs to the Herrerasauria. We were able to identify several similarities with potential non-herrerasaurid herrerasaurians (e.g., Tawa hallae, "Caseosaurus crosbyensis"), which were previously only known from North American deposits. In contrast, herrerasaurids (e.g., Herrerasaurus ischigualastensis) are almost exclusively known from South America. Our results support the nesting of CAPPA/UFSM 0373 as an early-diverging herrerasaurian. Furthermore, this is potentially the first record of a non-herrerasaurid herrerasaurian in unambiguous Carnian beds, suggesting a hidden diversity of dinosaurs in the Carnian rocks of the Candelária Sequence, which can be revealed even by fragmentary specimens.

Reassessment of the enigmatic "Prestosuchus" loricatus (Archosauria: Pseudosuchia) from the Middle-Late Triassic of southern Brazil.

Our knowledge of the diversity and evolution of South American Triassic pseudosuchians has greatly improved in the past 15 years, due to new discoveries, but also to the revision of several historically important specimens. One of the earliest descriptions of pseudosuchians from the Triassic of Brazil stems from the classic work of Huene from the first half of the 20th century, who described several species, including such influential taxa as Rauisuchus tiradentes or Prestosuchus chiniquensis, which have recently been reviewed. The more poorly known proposed second species of Prestosuchus, P. loricatus, is the focus of the present work. The original material included some elements of the axial skeleton (cervical and caudal vertebrae, ribs, osteoderms) and the hindlimb (ischia, calcaneum, metatarsus), collected from the Dinodontosaurus Assemblage Zone of the Chiniquá area, west of São Pedro do Sul. "Prestosuchus" loricatus shows numerous differences to P. chiniquensis, including the shape of cervical neural spines, presence of epipophyses on the cervical vertebrae, presence of a pit in the iliac articulation of the ischium, lack of longitudinal furrows in the dorsolateral surface of the ischial shafts, the more slender calcaneal tuber and a less pronounced ventral pit in the calcaneum, and is thus referred to a new genus, Schultzsuchus gen. nov. Phylogenetic analysis indicates an early branching position within Poposauroidea for Schultzsuchus, making it the oldest known member of this clade in South America.

Carbon cycle perturbations and environmental change of the middle permian and Late Triassic Paleo-Antarctic circle.

During the middle Permian through the Triassic, Tasmania moved from paleo-latitudes of 78° to 69°S, wedged between Antarctica and Australia, within the paleo-South polar circle. During this time, significant global carbon cycle disturbances triggered major environmental and climatic changes and mass extinction events globally. The Bicheno-5 core from Eastern Tasmania, Australia, provides the opportunity to examine middle Permian and Upper Triassic sediments from the paleo-Antarctic, using high-resolution organic carbon isotope (δ13CTOC) chemostratigraphy, pXRF, and sedimentology, combined with new palynological data integrated with the existing radiometric age model. While there is a significant unconformity in the Upper Permian to the middle Triassic associated with eustatic sea-level fall as a result of regional uplift in eastern Australia, three distinct carbon isotope excursions (CIEs), characterized by negative shifts of up to - 6 ‰ were identified; the middle Permian Guadalupian Carbon Isotope Excursions (G-CIE), the Carnian Pluvial Episode (CPE), and the mid-Norian Event (MNE). These three events highlight a significant climate shift through glacial and interglacial cycles to warmer non-glacial intervals in the Late Triassic, with evidence of the polar record of the Carnian Pluvial Episode and the mid-Norian Event, which are poorly studied in the Southern Hemisphere, specifically within the Paleo-Antarctic circle.

The first record of chigutisaurid amphibian from the Late Triassic Tiki Formation and the probable Carnian pluvial episode in central India.

A new, partially preserved skull of chigutisaurid amphibian (temnospondyli) has been reported for the first time from the Late Triassic Tiki Formation of India. Chigutisaurids are now known to occur in Australia's Early and Late Triassic, the Late Triassic in India, Argentina, and Brazil, the Jurassic of South Africa and Australia, and the Cretaceous of Australia. In India, the first appearance of chigutisaurids marks the Carnian-middle Carnian/Norian Boundary. This work also attempts to correlate, again for the first time, the advent of chigutisaurids and the occurrence of Carnian Pluvial Episodes (CPE) in the Late Triassic Maleri and Tiki Formation of Central India. The new specimen belongs to the genus Compsocerops prevalent in the Late Triassic Maleri Formation occurring 700 km south. However, the chigutisaurid specimen recovered from the Tiki Formation is a new species when compared to that of the Maleri Formation. It has the presence of an inward curved process of the quadratojugal as opposed to the straight downward trending process of the quadratojugal, the presence of vomerine foramen, shorter and wider interpterygoid vacuities, wider subtemporal vacuities, and the base of the interpterygoid vacuities at the same level with the base of the subtemporal vacuity. It proves that the Tiki Formation is coeval with the Lower Maleri Formation and a part of Upper Maleri.

Skull osteology of the holotype of the rhynchosaur Hyperodapedon sanjuanensis (Sill, 1970) from the Upper Triassic Ischigualasto Formation of Argentina.

Rhynchosaurs are bulky quadrupedal herbivores that achieved a cosmopolitan distribution during the Middle and Late Triassic. Rhynchosaurids are characterized by a pair of premaxillae modified into an edentulous beak that had a bone-to-bone occlusion with the tips of the dentaries, and a specialized masticatory apparatus composed of groove(s) on the maxilla and ridges(s) on the dentary. The Argentinian fossil record of rhynchosaurs is abundant, but only two nominal species have been named so far. One of them, the hyperodapedontine Hyperodapedon sanjuanensis, is the most abundant tetrapod of the Ischigualasto Formation of northwestern Argentina. However, the anatomy of H. sanjuanensis remains poorly studied. The repreparation and computed tomographic (CT) scanning of its holotypic skull (MACN-Pv 18185) allowed a detailed description of its anatomy. Among the novel anatomical data presented here, we provide the first tridimensional complete reconstruction of a rhynchosaurid palate. This structure is extremely complex in comparison to the plesiomorphic archosauromorph condition, and more data on other rhynchosaurid palates are needed to shed light on the sequence of character states acquisition that resulted in this peculiar anatomy. The presence of a pair of septomaxillae in H. sanjuanensis is the first report of these bones in a rhynchosaurid, and this new information allowed to identify septomaxillae also in the hyperodapedontines Teyumbaita sulcognathus and Hyperodapedon mariensis. The better understanding of the skull anatomy of the holotype of H. sanjuanensis is a first step towards an improved knowledge of the morphology and taxonomy of the South American rhynchosaurs.

Photosymbiosis in Late Triassic scleractinian corals from the Italian Dolomites.

During the Carnian, oligotrophic shallow-water regions of the western Tethys were occupied by small, coral-rich patch reefs. Scleractinian corals, which already contributed to the formation of the reef structure, owed their position most probably to the symbiosis with dinoflagellate algae (zooxanthellae). Using microstructural (regularity of growth increments) and geochemical (oxygen and carbon stable isotopes) criteria of zooxanthellae symbiosis, we investigated whether this partnership was widespread among Carnian scleractinians from the Italian Dolomites (locality Alpe di Specie). Although corals from this locality are renowned from excellent mineralogical preservation (aragonite), their skeletons were rigorously tested against traces of diagenesis Irrespective of their growth forms, well preserved skeletons of corals from the Dolomites, most frequently revealed regular growth bands (low values of coefficient of variation) typical of modern zooxanthellate corals. Paradoxically, some Carnian taxa (Thamnasteriomorpha frechi and Thamnasteriomorphasp.)with highly integrated thamnasterioid colonies which today are formed exclusively by zooxanthellate corals, showed irregular fine-scale growth bands (coefficient of variation of 40% and 41% respectively) that could suggest their asymbiotic status. However, similar irregular skeletal banding is known also in some modern agariciids (Leptoseris fragilis) which are symbiotic with zooxanthellae. This may point to a similar ecological adaptation of Triassic taxa with thamnasterioid colonies. Contrary to occasionally ambiguous interpretation of growth banding, all examined Carnian corals exhibited lack of distinct correlation between carbon (δ 13C range between 0.81‰ and 5.81‰) and oxygen (δ 18O values range between -4.21‰ and -1.06‰) isotope composition of the skeleton which is consistent with similar pattern in modern zooxanthellates. It is therefore highly likely, that Carnian scleractinian corals exhibited analogous ecological adaptations as modern symbiotic corals and that coral-algal symbiosis that spread across various clades of Scleractinia preceded the reef bloom at the end of the Triassic.

Gnathovorax cabreirai: a new early dinosaur and the origin and initial radiation of predatory dinosaurs.

Predatory dinosaurs were an important ecological component of terrestrial Mesozoic ecosystems. Though theropod dinosaurs carried this role during the Jurassic and Cretaceous Periods (and probably the post-Carnian portion of the Triassic), it is difficult to depict the Carnian scenario, due to the scarcity of fossils. Until now, knowledge on the earliest predatory dinosaurs mostly relies on herrerasaurids recorded in Carnian strata of South America. Phylogenetic investigations recovered the clade in different positions within Dinosauria, whereas fewer studies challenged its monophyly. Although herrerasaurid fossils are much better recorded in present-day Argentina than in Brazil, Argentinean strata so far yielded no fairly complete skeleton representing a single individual. Here, we describe Gnathovorax cabreirai, a new herrerasaurid based on an exquisite specimen found as part of a multitaxic association form southern Brazil. The type specimen comprises a complete and well-preserved articulated skeleton, preserved in close association (side by side) with rhynchosaur and cynodont remains. Given its superb state of preservation and completeness, the new specimen sheds light into poorly understood aspects of the herrerasaurid anatomy, including endocranial soft tissues. The specimen also reinforces the monophyletic status of the group, and provides clues on the ecomorphology of the early carnivorous dinosaurs. Indeed, an ecomorphological analysis employing dental traits indicates that herrerasaurids occupy a particular area in the morphospace of faunivorous dinosaurs, which partially overlaps the area occupied by post-Carnian theropods. This indicates that herrerasaurid dinosaurs preceded the ecological role that later would be occupied by large to medium-sized theropods.

Skull anatomy and phylogenetic assessment of a large specimen of Ecteniniidae (Eucynodontia: Probainognathia) from the Upper Triassic of southern Brazil.

Ecteniniidae comprises an endemic radiation of carnivore probainognathian cynodonts from the Late Triassic of South America. Three taxa have been included in this clade: Ecteninion lunensis Martínez et al., 1996 and Diegocanis elegans Martínez et al., 2013 from Argentina, and Trucidocynodon riograndensis Oliveira et al., 2010 from Brazil. Herein, a new specimen (skull and mandible) assigned to T. riograndensis from the Carnian of the Candelária Sequence (Southern Brazil) is described. A phylogenetic analysis recovered the new specimen as the sister taxon of the holotype of T. riograndensis, and both in a trichotomy with E. lunensis and D. elegans, all supporting the monophyly of Ecteniniidae. The new specimen of T. riograndensis is almost 20% larger than its holotype. Therefore, it represents one of the largest specimens of a carnivorous probainognathian from the Late Triassic known to date and contributes to knowledge of size variation in ecteniniids.

Under the armor: X-ray computed tomographic reconstruction of the internal skeleton of Coahomasuchus chathamensis (Archosauria: Aetosauria) from the Upper Triassic of North Carolina, USA, and a phylogenetic analysis of Aetosauria.

Aetosauria is a clade of heavily armored, quadrupedal omnivorous to herbivorous archosaurs known from the Late Triassic across what was the supercontinent of Pangea. Their abundance in many deposits relative to the paucity of other Triassic herbivores indicates that they were key components of Late Triassic ecosystems. However, their evolutionary relationships remain contentious due, in large part, to their extensive dermal armor, which often obstructs observation of internal skeletal anatomy and limits access to potentially informative characters. In an attempt to address this problem we reanalyzed the holotype of a recently described species of Coahomasuchus, C. chathamensis, from the Sanford sub-basin of North Carolina using computed tomography (CT). CT scans of the holotype specimen clarify preservation of the skeleton, revealing several articulated vertebrae and ribs, an isolated vertebra, left ulna, left scapula, and the right humerus, though none of the material resulted in updated phylogenetic scorings. Reexamination of aetosaur materials from the holotype locality also indicates that several isolated osteoderms and elements of the appendicular skeleton are newly referable. Based on these results, we update the Coahomasuchus chathamensis hypodigm and conduct a revised phylogenetic analysis with improved character scorings for Coahomasuchus and several other aetosaurs. Our study recovers Coahomasuchus in a polytomy with Aetosaurus and the Typothoracinae, in contrast with a recent analysis that recovered Coahomasuchus as a wild-card taxon.

Homeotic shift at the dawn of the turtle evolution.

All derived turtles are characterized by one of the strongest reductions of the dorsal elements among Amniota, and have only 10 dorsal and eight cervical vertebrae. I demonstrate that the Late Triassic turtles, which represent successive stages of the shell evolution, indicate that the shift of the boundary between the cervical and dorsal sections of the vertebral column occurred over the course of several million years after the formation of complete carapace. The more generalized reptilian formula of at most seven cervicals and at least 11 dorsals is thus plesiomorphic for Testudinata. The morphological modifications associated with an anterior homeotic change of the first dorsal vertebra towards the last cervical vertebra in the Triassic turtles are partially recapitulated by the reduction of the first dorsal vertebra in crown-group Testudines, and they resemble the morphologies observed under laboratory conditions resulting from the experimental changes of Hox gene expression patterns. This homeotic shift hypothesis is supported by the, unique to turtles, restriction of Hox-5 expression domains, somitic precursors of scapula, and brachial plexus branches to the cervical region, by the number of the marginal scute-forming placodes, which was larger in the Triassic than in modern turtles, and by phylogenetic analyses.

Supradapedon revisited: geological explorations in the Triassic of southern Tanzania.

The upper Triassic deposits of the Selous Basin in south Tanzania have not been prospected for fossil tetrapods since the middle of last century, when Gordon M. Stockley collected two rhynchosaur bone fragments from the so called "Tunduru beds". Here we present the results of a field trip conducted in July 2015 to the vicinities of Tunduru and Msamara, Ruvuma Region, Tanzania, in search for similar remains. Even if unsuccessful in terms of fossil discoveries, the geological mapping conducted during the trip improved our knowledge of the deposition systems of the southern margin of the Selous Basin during the Triassic, allowing tentative correlations to its central part and to neighbouring basins. Moreover, we reviewed the fossil material previously collected by Gordon M. Stockley, confirming that the remains correspond to a valid species, Supradapedon stockleyi, which was incorporated into a comprehensive phylogeny of rhynchosaurs and found to represent an Hyperodapedontinae with a set of mostly plesiomorphic traits for the group. Data gathered form the revision and phylogenetic placement of Su. stockleyi helps understanding the acquisition of the typical dental traits of Late Triassic rhynchosaurs, corroborating the potential of hyperodapedontines as index fossils of the Carnian-earliest Norian.

Taphonomy and palaeoecology of Late Triassic (Carnian) ammonoid concentrations from the Taurus Mountains, Turkey.

The deposits of the Carnian Kasimlar Formation within the Taurus Platform Units of south-western Turkey represent an important archive of a Late Triassic ecosystem. New palaeontological information was obtained by analysing the Kasimlarceltites mass occurrence, located within the Kasimlar Formation and named after the Lower Carnian (Julian) ammonoid genus Kasimlarceltites. This is the dominant taxon (> 94%) within the mass occurrence: nearly 775 million ammonoids and 50 million gastropods were extrapolated for the whole extension (at least 5 km2) of the Kasimlarceltites beds. This calculation is one of the main findings within this study, as it is the first time that such a fossil mass occurrence was quantified. Additionally, orientation measurements of the planispiral ammonoids and the helical gastropods enabled reconstructing the history of the mass occurrence and interpreting the underlying transport mechanisms. Further taphonomic aspects (e.g. biofabric, preservation, bioerosion or genetic classification) as well as comparisons with samples of the same acme zone from different localities near Asagiyaylabel (AS IV, KA I-II) point to a two-phased genetic history. Accordingly, local mass mortality within the Kasimlarceltites fauna due to oxygen fluctuations or methane degassing may have initially led to a primary accumulation. These deposits were then reworked and redeposited basinward by gravity flows to create the present-day secondary allochthonous concentrations.

A Unique Late Triassic Dinosauromorph Assemblage Reveals Dinosaur Ancestral Anatomy and Diet.

Dinosauromorpha includes dinosaurs and other much less diverse dinosaur precursors of Triassic age, such as lagerpetids [1]. Joint occurrences of these taxa with dinosaurs are rare but more common during the latest part of that period (Norian-Rhaetian, 228-201 million years ago [mya]) [2, 3]. In contrast, the new lagerpetid and saurischian dinosaur described here were unearthed from one of the oldest rock units with dinosaur fossils worldwide, the Carnian (237-228 mya) Santa Maria Formation of south Brazil [4], a record only matched in age by much more fragmentary remains from Argentina [5]. This is the first time nearly complete dinosaur and non-dinosaur dinosauromorph remains are found together in the same excavation, clearly showing that these animals were contemporaries since the first stages of dinosaur evolution. The new lagerpetid preserves the first skull, scapular and forelimb elements, plus associated vertebrae, known for the group, revealing how dinosaurs acquired several of their typical anatomical traits. Furthermore, a novel phylogenetic analysis shows the new dinosaur as the most basal Sauropodomorpha. Its plesiomorphic teeth, strictly adapted to faunivory, provide crucial data to infer the feeding behavior of the first dinosaurs.

A new herrerasaurid (Dinosauria, Saurischia) from the Upper Triassic Ischigualasto Formation of northwestern Argentina.

Herrerasauridae comprises a basal clade of dinosaurs best known from the Upper Triassic of Argentina and Brazil, which have yielded remains of Herrerasaurus ischigualastensis and Staurikosaurus pricei, respectively. Systematic opinion regarding the position of Herrerasauridae at the base of Dinosauria has varied. Here we describe a new herrerasaurid, Sanjuansaurus gordilloi gen. n., sp. n., based on a partial skeleton from Carnian-age strata of the the Upper Triassic Ischigualasto Formation of northwestern Argentina. The new taxon is diagnosed by numerous features, including long, band-shaped and posterolaterally oriented transverse process on the posterior cervical vertebrae; neural spines of the sixth to eighth dorsal vertebrae, at least, bearing acute anterior and posterior processes; scapula and coracoid with everted lateral margins of the glenoid; and short pubis (63% of the femoral length). Phylogenetic analysis placed Sanjuansaurus within a monophyletic Herrerasauridae, at the base of Theropoda and including Herrerasaurus and Staurikosaurus. The presence of Sanjuansaurus at the base of the Ischigualasto Formation, along with other dinosaurs such as Herrerasaurus, Eoraptor, Panphagia, and Chromogisaurus suggests that saurischian dinosaurs in southwestern Pangea were already widely diversified by the late Carnian rather than increasing in diversity across the Carnian-Norian boundary.