The dawn of an Era: New contributions on comparative and functional anatomy of Triassic tetrapods.

The Triassic period stands as a crucial moment for understanding tetrapod evolution, marking the emergence and early diversification of numerous lineages that persist in today's ecosystems. Birds, crocodiles, testudines, lizards, and mammals can all trace their origins to the Triassic, which is distinguished by several adaptive radiation events that fostered unparalleled diversity in body plans and lifestyles. Beyond this macroevolutionary significance, the Triassic period serves as fertile ground for scientific inquiry, especially in tetrapod studies. The aim of this Special Issue is to assemble a diverse array of new contributions focused on continental Triassic tetrapods globally, encouraging collaboration among researchers across generations, pooling their efforts to comprehend this pivotal moment in tetrapod evolutionary history. This issue encompasses almost 40 varied contributions, spanning topics from comparative and functional anatomy, including descriptions of novel taxa, comprehensive anatomical reviews, systematic investigations, phylogenetic analyses, paleoneurological studies, biomechanical assessments, and detailed examinations of histology and ontogeny. Collectively, this Special Issue offers an extensive exploration of Triassic tetrapods from anatomical, ecological, and evolutionary perspectives, unveiling fresh insights into this intriguing moment in vertebrate evolutionary history.

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.

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.

New rhadinosuchine proterochampsids from the late Middle-early Late Triassic of southern Brazil enhance the diversity of archosauriforms.

Proterochampsidae is a clade of non-archosaurian archosauriforms restricted to the Middle to the Late Triassic of the Ischigualasto-Villa Unión Basin of Argentina and the Santa Maria Supersequence of Brazil. A reappraisal of proterochampsid specimens from the Brazilian Dinodontosaurus Assemblage Zone (AZ) of the Pinheiros-Chiniquá Sequence (late Ladinian-early Carnian) is presented here. One of the specimens was preliminary assigned to Chanaresuchus sp., whose type species comes from the Massetognathus-Chanaresuchus AZ of the Chañares Formation of Argentina. However, our revision indicates that it differs from Chanaresuchus, being more closely related to the middle-late Carnian Rhadinosuchus gracilis. We therefore propose the new taxon, Pinheirochampsa rodriguesi, to reallocate this specimen. Additionally, we present a revision of other putative Chanaresuchus occurrences in Brazil, including the only known specimen described for the Santacruzodon AZ (Santa Cruz do Sul Sequence; early Carnian), also proposing it as a new taxon: Kuruxuchampsa dornellesi. Both new species are characterized, among other features, by transverse expansion of the anterior end of the rostrum, similar to the condition present in Rhadinosuchus, but absent in Chanaresuchus, Gualosuchus, Pseudochampsa, and non-rhadinosuchine proterochampsids. These two new species expand the growing knowledge of the non-archosaurian archosauriform diversity during the Middle-Late Triassic in South America and enhance faunal and chronological comparisons between approximately coeval geological units between Argentina and Brazil.

The absence of an invasive air sac system in the earliest dinosaurs suggests multiple origins of vertebral pneumaticity.

The origin of the air sac system present in birds has been an enigma for decades. Skeletal pneumaticity related to an air sac system is present in both derived non-avian dinosaurs and pterosaurs. But the question remained open whether this was a shared trait present in the common avemetatarsalian ancestor. We analyzed three taxa from the Late Triassic of South Brazil, which are some of the oldest representatives of this clade (233.23 ± 0.73 Ma), including two sauropodomorphs and one herrerasaurid. All three taxa present shallow lateral fossae in the centra of their presacral vertebrae. Foramina are present in many of the fossae but at diminutive sizes consistent with neurovascular rather than pneumatic origin. Micro-tomography reveals a chaotic architecture of dense apneumatic bone tissue in all three taxa. The early sauropodomorphs showed more complex vascularity, which possibly served as the framework for the future camerate and camellate pneumatic structures of more derived saurischians. Finally, the evidence of the absence of postcranial skeletal pneumaticity in the oldest dinosaurs contradicts the homology hypothesis for an invasive diverticula system and suggests that this trait evolved independently at least 3 times in pterosaurs, theropods, and sauropodomorphs.

A new specimen provides insights into the anatomy of Irajatherium hernandezi, a poorly known probainognathian cynodont from the Late Triassic of southern Brazil.

Irajatherium hernandezi is a poorly known non-mammaliaform cynodont from the Late Triassic of southern Brazil. A new specimen of this cynodont was found in recent fieldwork to the type-locality, Sesmaria do Pinhal (Candelária), providing new insights into the anatomy of this mammalian forerunner. This specimen comprises a partial skull preserving the left canine, two left and three right postcanines, and an isolated exoccipital; the left dentary with the canine and postcanines; a fragment of the right dentary; the proximal portion of the left partial humerus; the right scapula; and indeterminate fragments. Based on new material, it is here suggested that I. hernandezi presents: a rostrum broad and short, possibly long as the temporal region; three foramina on the lateral surface of the maxilla, that could correspond to the external openings of the rostral alveolar, infraorbital, and zygomaticofacial canals; a slender zygomatic arch and an absent postorbital bar; a posteriorly wide temporal fossa; a long secondary palate, slightly surpassing the level of the last postcanine tooth; the cerebral hemispheres of the cranial endocast divided by a median sulcus; the scapular blade long and straight, and the postscapular fossa absent in lateral aspect. Finally, I. hernandezi and other tritheledontids were included in a phylogenetic analysis of Eucynodontia. The analysis recovered unresolved relationships for ictidosaurs/tritheledontids, nested within a polytomy with Tritylodontidae and a clade composed by Pseudotherium argentinus, Botucaraitherium belarminoi, Brasilodon quadrangularis, and Mammaliaformes.

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.

The endocranial anatomy of Buriolestes schultzi (Dinosauria: Saurischia) and the early evolution of brain tissues in sauropodomorph dinosaurs.

Our knowledge on the anatomy of the first dinosaurs (Late Triassic, 235-205 Ma) has drastically increased in the last years, mainly due to several new findings of exceptionally well-preserved specimens. Nevertheless, some structures such as the neurocranium and its associated structures (brain, labyrinth, cranial nerves, and vasculature) remain poorly known, especially due to the lack of specimens preserving a complete and articulated neurocranium. This study helps to fill this gap by investigating the endocranial cavity of one of the earliest sauropodomorphs, Buriolestes schultzi, from the Upper Triassic (Carnian-c. 233 Ma) of Brazil. The endocranial anatomy of this animal sheds light on the ancestral condition of the brain of sauropodomorphs, revealing an elongated olfactory tract combined to a relatively small pituitary gland and well-developed flocculus of the cerebellum. These traits change drastically across the evolutionary history of sauropodomorphs, reaching the opposite morphology in Jurassic times. Furthermore, we present here the first calculations of the Reptile Encephalization Quotient (REQ) for a Triassic dinosaur. The REQ of B. schultzi is lower than that of Jurassic theropods, but higher than that of later sauropodomorphs. The combination of cerebral, dental, and postcranial data suggest that B. schultzi was an active small predator, able to track moving prey.

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.

A dinosaur ilium from the Late Triassic of Brazil with comments on key-character supporting Saturnaliinae.

Discoveries in Carnian-aged rocks are establishing a rich and diverse dinosaurian fauna at the so-called 'dawn of the age of dinosaurs' in the Late Triassic of Western Gondwana. Accordingly, Brazilian strata from the Candelária Sequence have contributed extensively to this trend. Here, we present a new dinosaurian specimen (CAPPA/UFSM 0200) from this geological unit. The material was collected at a fossiliferous site that had no previous record of dinosaurs. Our specimen comprises a single ilium, which we describe in detail. Its anatomy is consistent with Carnian sauropodomorph dinosaurs, but differs from coeval specimens by several features, although we do not discard the possibility of these features being the result of intraspecific variation. In part of our phylogenetic investigation, CAPPA/UFSM 0200 was recovered within Saturnaliinae, a group comprised of Carnian sauropodomorphs from South America. However, based on examination of better-sampled coeval taxa, a character (a strong rugosity in the ilium) supporting this less inclusive position might be related to intraspecific variation and so, should be carefully considered. This study increases the distribution of dinosaur remains in fossiliferous units from southern Brazil and adds to the discussion regarding intraspecific variation and its implications in the phylogeny of early dinosaurs.

A New Clevosaurid from the Triassic (Carnian) of Brazil and the Rise of Sphenodontians in Gondwana.

The early evolution of lepidosaurs is marked by an extremely scarce fossil record during the Triassic. Importantly, most Triassic lepidosaur specimens are represented by disarticulated individuals from high energy accretion deposits in Laurasia, thus greatly hampering our understanding of the initial stages of lepidosaur evolution. Here, we describe the fragmentary remains of an associated skull and mandible of Clevosaurus hadroprodon sp. nov., a new taxon of sphenodontian lepidosaur from the Late Triassic (Carnian; 237-228 Mya) of Brazil. Referral to Sphenodontia is supported by the combined presence of a marginal dentition ankylosed to the apex of the dentary, maxilla, and premaxilla; the presence of 'secondary bone' at the bases of the marginal dentition; and a ventrally directed mental process at the symphysis of the dentary. Our phylogenetic analyses recover Clevosaurus hadroprodon as a clevosaurid, either in a polytomy with the Late Triassic to Early Jurassic Clevosaurus and Brachyrhinodon (under Bayesian inference), or nested among different species of Clevosaurus (under maximum parsimony). Clevosaurus hadroprodon represents the oldest known sphenodontian from Gondwana, and its clevosaurid relationships indicates that these sphenodontians achieved a widespread biogeographic distribution much earlier than previously thought.