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.
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.
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.
This paper describes the oldest record of fused sacral vertebrae for Dinosauria, based on two primordial sacrals from the Brazilian Upper Triassic. We also review the occurrence of sacral fusion in Dinosauria and close relatives. The new specimen indicates that sacral fusion occurred early in the evolution of dinosaurs and our review suggests that, at least in Sauropodomorpha, the primordial sacrals fuse prior to the additional sacral vertebrae, which fuse later along the lineage.
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.
The first complete endocast of an early dinosaur is presented. The endocranium of Buriolestes schultzi reveals relatively small olfactory bulbs, an elongated olfactory tract, a small pituitary fossa, and well‐developed flocculus of the cerebellum. The data support B. schultzi as an active small predator, which was able to track moving prey. These endocranial traits change drastically across the evolutionary history of sauropodomorphs, reaching the opposite morphology in Jurassic times.
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.
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.
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.
The Paraná Basin was filled by a sedimentary package deposited in successive sedimentation episodes related to the tectonic events that hit the SW portion of the Gondwana. The Triassic portion of ...this package, known worldwide for its continental tetrapod fauna, occurs only in the southern portion of the basin and is represented by 2 s-order sequences: the Sanga do Cabral Supersequence (SCS - Early Triassic) and the Santa Maria Supersequence (SMS - Middle-Late Triassic). The SCS fauna, including temnospondyls, parareptiles (mainly Procolophon), archosauromorphs, putative synapsids, and a number of indeterminate specimens, is traditionally considered Early Triassic and corresponds to the “Procolophon abundant zone” of the Karoo Basin (the upper levels of the Lystrosaurus AZ), in the upper Katberg Formation, which is Induan to early Olenekian in age. The sedimentary environment of the SCS is thought to be a wide alluvial plain, in which small and shallow channels spread northwards into a vast semiarid environment. By its turn, the Middle-Upper Triassic Santa Maria Supersequence is divided into four third-order sequences, from base to top: Pinheiros-Chiniquá, Santa Cruz, Candelária and Mata. Each of these sequences begins with fluvial deposition (low sinuosity rivers) that is overlain by transgressive shallow lacustrine deposits. The first three of these sequences present a very rich record of fossil tetrapods, including four successive faunal associations: Dinodontosaurus Assemblage Zone (Ladinian, within the Pinheiros-Chiniquá Sequence), Santacruzodon AZ (Ladinian/Carnian, in the Santa Cruz Sequence), Hyperodapedon AZ (Carnian) and the Riograndia AZ (early Norian), the latter two respectively at the base and top of the Candelária Sequence. In general, the lower portion of the package (Pinheiros-Chiniquá and Santa Cruz Sequences) was deposited under more basic and dried environmental conditions and are dominated by synapsids, while the top of the section (Candelária Sequence) is characterized by more acid and humid conditions and by a shift in the faunal content, with diapsids as dominating forms and presenting an increase of diversity compared to the lower biozones.
•In Brazil, Triassic tetrapods are known only in the extreme south of the country.•The south Brazilian Triassic package includes four biozones based on tetrapods.•The lower tetrapod biozone of the Triassic package is Early Triassic and corresponds to the “Procolophon abundant zone” of the Karoo Basin.•Lower portion of the Middle-Late Triassic was deposited under more basic and dried environmental conditions and is dominated by synapsids.•Upper portion of the Middle-Late Triassic is characterized by more acid and humid conditions and presents an increase of diapsids compared to the lower portion.
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