Birds, dinosaurs, crocodilians, pterosaurs and their close relatives form the highly diverse clade Archosauriformes. Archosauriforms have a deep evolutionary history, originating in the late Permian, prior to the end-Permian mass extinction, and radiating in the Triassic to dominate Mesozoic ecosystems. However, the origins of this clade and its extraordinarily successful body plan remain obscure. Here, we describe an exceptionally preserved fossil skull from the Lower Triassic of Brazil, representing a new species, Teyujagua paradoxa, transitional in morphology between archosauriforms and more primitive reptiles. This skull reveals for the first time the mosaic assembly of key features of the archosauriform skull, including the antorbital and mandibular fenestrae, serrated teeth, and closed lower temporal bar. Phylogenetic analysis recovers Teyujagua as the sister taxon to Archosauriformes, and is congruent with a two-phase model of early archosauriform evolution, in response to two mass extinctions occurring at the end of the Guadalupian and the Permian.
Archosauriformes are a major group of fossil and living reptiles that include the crown group Archosauria (birds, crocodilians and their extinct relatives) and closely related taxa. Archosauriformes are characterized by a highly diagnostic skull architecture, which is linked to the predatory habits of their early representatives, and the development of extensive cranial pneumaticity associated with the nasal capsule. The evolution of the archosauriform skull from the more plesiomorphic configuration present ancestrally in the broader clade Archosauromorpha was, until recently, elusive. This began to change with the discovery and description of Teyujagua paradoxa, an early archosauromorph from the Lower Triassic Sanga do Cabral Formation of Brazil. Here, we provide a detailed osteological description of the holotype and, thus far, only known specimen of T. paradoxa. In addition to providing new details of the anatomy of T. paradoxa, our study also reveals an early development of skull pneumaticity prior to the emergence of the antorbital fenestra. We use these new data to discuss the evolution of antorbital openings within Archosauriformes. Reappraisal of the phylogenetic position of T. paradoxa supports previous hypotheses of a close relationship with Archosauriformes. The data presented here provide new insights into character evolution during the origin of the archosauriform skull.
Background Anhanguerids comprise an important clade of pterosaurs, mostly known from dozens of three-dimensionally preserved specimens recovered from the Lower Cretaceous Romualdo Formation (northeastern Brazil). They are remarkably diverse in this sedimentary unit, with eight named species, six of them belonging to the genus Anhanguera. However, such diversity is likely overestimated, as these species have been historically diagnosed based on subtle differences, mainly based on the shape and position of the cranial crest. In spite of that, recently discovered pterosaur taxa represented by large numbers of individuals, including juveniles and adults, as well as presumed males and females, have crests of sizes and shapes that are either ontogenetically variable or sexually dimorphic. Methods We describe in detail the skull of one of the most complete specimens referred to Anhanguera, AMNH 22555, and use it as a case study to review the diversity of anhanguerids from the Romualdo Formation. In order to accomplish that, a geometric morphometric analysis was performed to assess size-dependent characters with respect to the premaxillary crest in the 12 most complete skulls bearing crests that are referred in, or related to, this clade, almost all of them analyzed first hand. Results Geometric morphometric regression of shape on centroid size was highly statistically significant (p = 0.0091) and showed that allometry accounts for 25.7% of total shape variation between skulls of different centroid sizes. Premaxillary crests are both taller and anteroposteriorly longer in larger skulls, a feature consistent with ontogenetic growth. A new diagnosis is proposed for Anhanguera, including traits that are nowadays known to be widespread within the genus, as well as ontogenetic changes. AMNH 22555 cannot be referred to “Anhanguera santanae” and, in fact, “Anhanguera santanae”, “Anhanguera araripensis”, and “Anhanguera robustus” are here considered nomina dubia. Discussion Historically, minor differences in crest morphology have been used in the definition of new anhanguerid species. Nowadays, this practice resulted in a considerable difficulty in referring well-preserved skulls into known taxa. When several specimens are analyzed, morphologies previously believed to be disparate are, in fact, separated by a continuum, and are thus better explained as individual or temporal variations. Stratigraphically controlled excavations on the Romualdo Formation have showed evidence for faunal turnover regarding fish communities. It is thus possible that some of the pterosaurs from this unit were not coeval, and might even represent anagenetic morphotypes. Unfortunately, amateur collecting of Romualdo Formation fossils, aimed especially at commerce, resulted in the lack of stratigraphic data of virtually all its pterosaurs and precludes testing of these further hypotheses.
Scientific practices stemming from colonialism, whereby middle- and low-income countries supply data for high-income countries and the contributions of local expertise are devalued, are still prevalent today in the field of palaeontology. In response to these unjust practices, countries such as Mexico and Brazil adopted protective laws and regulations during the twentieth century to preserve their palaeontological heritage. However, scientific colonialism is still reflected in many publications describing fossil specimens recovered from these countries. Here, we present examples of ‘palaeontological colonialism’ from publications on Jurassic–Cretaceous fossils from NE Mexico and NE Brazil spanning the last three decades. Common issues that we identified in these publications are the absence of both fieldwork and export permit declarations and the lack of local experts among authorships. In Mexico, access to many fossil specimens is restricted on account of these specimens being housed in private collections, whereas a high number of studies on Brazilian fossils are based on specimens illegally reposited in foreign collections, particularly in Germany and Japan. Finally, we outline and discuss the wider academic and social impacts of these research practices, and propose exhaustive recommendations to scientists, journals, museums, research institutions and government and funding agencies in order to overcome these practices.
The oldest archosauromorphs (dinosaurs, birds, crocodiles, and their stem-taxa) are recorded in middle− upper Permian rocks, but it was not after the Permo− Triassic mass extinction that the group shows a substantially high taxonomic richness and ecomorphological disparity. The early evolutionary history of the Archosauromorpha during the Early and Middle Triassic is mainly based on fossils recovered from rocks in southern Africa, Europe and Asia, whereas South America possesses a more complete fossil record of the group only in the Late Triassic.Here we revisit, discuss, and reanalyse the non-archosaurian archosauromorph fossil record of the current-day South America. The Early Triassic archosauromorph record in this continent is still scarce, but it documents the early evolution of the group in western Pangaea and is crucial to understand more globally the biotic recovery after the Permo− Triassic mass extinction. The Middle Triassic record is extremely scarce, but the Late Triassic archosauromorph assemblage of South America is among the most diverse and abundant worldwide. The last decade has witnessed a considerable improvement in our knowledge of the record, taxonomy, phylogeny, and macroevolution of the group with the input from the South American fossils. Nevertheless, a considerable amount of research is needed and ideally should be focused on some particular aspects of the Triassic evolutionary radiation of Archosauromorpha. Among them, the Early Triassic record should be expanded, more numerous and more complete Middle Triassic archosauromorph specimens are crucial to have a more complete picture of the evolution of the group, and the taxonomy of groups like proterochampsids and hyperodapedontine rhynchosaurs should be clarified through detailed anatomical work.
A remarkably well-preserved, almost complete and articulated new specimen (GP/2E 9266) of Tupandactylus navigans is here described for the Early Cretaceous Crato Formation of Brazil. The new specimen comprises an almost complete skeleton, preserving both the skull and post-cranium, associated with remarkable preservation of soft tissues, which makes it the most complete tapejarid known thus far. CT-Scanning was performed to allow the assessment of bones still covered by sediment. The specimen can be assigned to Tupa. navigans due to its vertical supra-premaxillary bony process and short and rounded parietal crest. It also bears the largest dentary crest among tapejarine pterosaurs and a notarium, which is absent in other representatives of the clade. The new specimen is here regarded as an adult individual. This is the first time that postcranial remains of Tupa. navigans are described, being also an unprecedented record of an articulated tapejarid skeleton from the Araripe Basin.
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