An additional brain endocast of the ictidosaur Riograndia guaibensis (Eucynodontia: Probainognathia): intraspecific variation of endocranial traits.

Recently, the morphology and encephalization of the brain endocast of the Triassic non-mammaliaform probainognathian cynodont Riograndia guaibensis were studied. Here, we analyzed the brain endocast of an additional specimen of this species. The new endocast shows well-defined olfactory bulbs and a median sulcus dividing the hemispheres, traits that were not clearly observed in the first studied specimen. Encephalization quotients were also calculated, revealing similar values to other non-mammaliaform cynodonts and lower than those of the first analyzed specimen. The analyzed cranium is slightly larger than the first studied one and may represent an advanced ontogenetic stage. Hence, these differences may be related to the intraspecific variation of this cynodont or alternatively, to the preservation of each specimen.

Old fossil findings in the Upper Triassic rocks of southern Brazil improve diversity of traversodontid cynodonts (Therapsida, Cynodontia).

In this contribution we describe a new genus and species of gomphodontosuchine cynodont from the Late Triassic Hyperodapedon Assemblage Zone (AZ) of the Santa Maria Supersequence of southern Brazil, based on material collected 20 years ago. The new taxon, Santagnathus mariensis gen. et sp. nov., is based on numerous cranial and postcranial remains, which altogether provide data on several parts of the skeleton. Santagnathus mariensis is closely related to Siriusgnathus niemeyerorum and Exaeretodon spp. and expands the knowledge about gomphodontosuchine cynodonts. Morphologically, the new species has a skull bauplan particularly similar to S. niemeyerorum and E. riograndensis, differing from them by a unique combination of traits such as three upper incisors, the absence of a descending process in the jugal, a more posterior position of the postorbital bar and a preorbital region larger than the temporal one. The new traversodontid was found associated with the rhynchosaur Hyperodapedon sp., corroborating the assignment of the cynodont fossils to the Hyperodapedon AZ. We also provide comments on the status of the Argentine traversodontid cynodont Proexaeretodon vincei, typically considered a junior synonym of Exaeretodon argentinus, and here accepted as a valid taxon.

Replacement tooth in mesosaurs and new data on dental microanatomy and microstructure.

The Permian mesosaurs are well known for being the earliest amniotes to exhibit adaptations for living in a marine environment (Irati-Whitehill Sea). In addition to their set of skeletal features associated with aquatic dwelling life, their dentition includes important characteristics related to feeding in this habitat, which is described in this work, based on the analysis of mesosaur specimens from the Lower Permian Irati Formation of Brazil. Mesosaurs have several slender, conical teeth bordered by enamel apicobasal ridges, a feature predominantly found in aquatic amniotes. Internally, the dentine walls are formed by the arrangement of layers of orthodentine and globular dentine. To prevent tooth loss, the basal area is equipped with plicidentine, a particular type of orthodentine, allied with cementum, alveolar bone trabeculae, and periodontal space that reinforces anchorage and provides some flexibility. The teeth are replaced in a labio-vertical path, and the dentition replaces alternately. This feature is regarded as plesiomorphic, and it ensures the oral cavity is supplied with enough teeth. However, these features do not the assessment of whether mesosaurs teeth were capable of piercing prey with resistant tegument. Instead, we interpret this adaptation as a mechanism for catching prey, at least in adults, and we endorse a possible ontogenetic dietary shift from small to large forms.

Weighing in on miniaturization: New body mass estimates for Triassic eucynodonts and analyses of body size evolution during the cynodont-mammal transition.

Body size influences most aspects of an animal's biology, consequently, evolutionary diversification is often accompanied by differentiation of body sizes within a lineage. It is accepted that miniaturization, or the evolution of extremely small body sizes, played a key role in the origin and early evolution of different mammalian characters in non-mammaliaform cynodonts. However, while there are multiple studies on the biomechanical, behavioral, and physiological consequences of smaller sizes, few explore the evolutionary processes that lead to them. Here, we use body mass as a universal size measurement in phylogenetic comparative analyses to explore aspects of body size evolution in Cynodontia, focusing on the cynodont-mammal transition, and test the miniaturization hypothesis for the origin of Mammaliaformes. We estimated the body masses of 29 species, ranging from Theriocephalia to Mammaliaformes, providing the largest collection of Triassic cynodont body mass estimates that we know of, and used these estimates in analyses of disparity through time and RRphylo . Unexpectedly, our results did not support the miniaturization hypothesis. Even though cynodont body size disparity fell during the Late Triassic, and remained lower than expected under a purely Brownian motion model of evolution up until the Early Jurassic, we found that rates of body size evolution were significantly lower in prozostrodontians leading to the first Mammaliaformes than in other lineages. Evolution rates were higher in medium and large-sized taxa, indicating that size was changing more rapidly in those lineages and that small sizes were probably a persistent plesiomorphic character-state in Cynodontia.

Bone histology supports gregarious behavior and an early ontogenetic stage to Decuriasuchus quartacolonia (Pseudosuchia: Loricata) from the Middle-Late Triassic of Brazil.

Decuriasuchus quartacolonia is a middle-sized basal "rauisuchian" (Pseudosuchia, Loricata) from the Triassic beds of Brazil, whose original description was based on 10 specimens of equivalent size found in aggregation. In this contribution, we explore the osteohistology of its appendicular bones and a rib, aiming to infer growth patterns and ontogeny. In all analyzed bones (except in the rib) from three individuals, we found fibrolamellar bone (FLB) throughout the cortices, a pattern shared with other histologically sampled basal loricatans. We found evidence that suggests an early ontogenetic stage for all studied specimens: absence of secondary osteons, lack of transition from an inner highly vascularized FLB matrix to a poor vascularized parallel-fibred bone, one line of arrested growth, open vascular canals in the external surface of the cortex and absence of an external fundamental system. In addition, we observed that the neurocentral sutures in the caudal to cervical vertebrae of the holotype are unfused, strongly suggesting that these individuals were not skeletally mature, as had been previously assumed. In addition, our data support the prior hypothesis that Decuriasuchus is the oldest-known archosaur to show evidence of gregarious behavior and adds a probable cause of this as a strategy to obtain food and avoid predation until becoming independent as adults. Furthermore, our results open the possibility that Decuriasuchus may represent an earlier growth stage of the larger Prestosuchus chiniquensis, with two specimens recovered from the same stratigraphic level and paleontological site. Since the adult form of Decuriasuchus remains unknown, a deeper anatomical study is needed to discuss the validity of the species. This study case exemplifies the importance of paleohistology as an essential complementary tool to describe a new morphospecies in vertebrate paleontology.

Reassessment of Faxinalipterus minimus, a purported Triassic pterosaur from southern Brazil with the description of a new taxon.

Faxinalipterus minimus was originally described as a purported pterosaur from the Late Triassic (early Norian) Caturrita Formation of southern Brazil. Its holotype comprises fragmentary postcranial elements, whereas a partial maxilla was referred to the species. The assignment of Faxinalipterus minimus to Pterosauria has been questioned by some studies, but the specimen has never been accessed in detail after its original description. Here we provide a reassessment of Faxinalipterus minimus after additional mechanical preparation of the holotype. Our interpretations on the identity of several bones differ from those of the original description, and we found no support favoring pterosaur affinities for the taxon. The maxilla previously referred to Faxinalipterus minimus is disassociated from this taxon and referred to a new putative pterosauromorph described here from a partial skull and fragmentary postcranial elements. Maehary bonapartei gen. et sp. nov. comes from the same fossiliferous site that yielded Faxinalipterus minimus, but the lack of overlapping bones hampers comparisons between the two taxa. Our phylogenetic analysis places Faxinalipterus minimus within Lagerpetidae and Maehary bonapartei gen. et sp. nov. as the earliest-diverging member of Pterosauromorpha. Furthermore, the peculiar morphology of the new taxon reveals a new dental morphotype for archosaurs, characterized by conical, unserrated crowns, with a pair of apicobasally oriented grooves. These two enigmatic archosaurs expand our knowledge on the Caturrita Formation fauna and reinforce the importance of its beds on the understanding of Late Triassic ecosystems.

The history of vaginal birth.

Vaginal delivery, as known today, is a still unfinished product that originated hundreds of million years ago, much before mammals evolved on land. In this article, we will discuss the way in which our direct ancestors were born over the eons until the present day, focusing on the factors that presented substantial changes in how birth occurred, in relation to our earlier ancestors. The history begins with the first amniotes around 300 million years ago and ends with the appearance of the first Homo sapiens around 160,000 years ago.

The first edentulous ceratosaur from South America.

The recognition of ontogenetic edentulism in the Jurassic noasaurid Limusaurus inextricabilis shed new light on the dietary diversity within Ceratosauria, a stem lineage of non-avian theropod dinosaurs known for peculiar craniomandibular adaptations. Until now, edentulism in Ceratosauria was exclusive to adult individuals of Limusaurus. Here, an exceptionally complete skeleton of a new toothless ceratosaur, Berthasaura leopoldinae gen. et sp. nov., is described from the Cretaceous aeolian sandstones of the Bauru Basin, Southern Brazil. The specimen resembles adult individuals of Limusaurus by the absence of teeth but based on the unfused condition of several elements (e.g., skull, vertebral column) it clearly represents an ontogenetically immature individual, indicating that it might never have had teeth. The phylogenetic analysis performed here has nested Berthasaura leopoldinae as an early-divergent Noasauridae, not closely related to Limusaurus. It represents the most complete non-avian theropod from the Brazilian Cretaceous and preserves the most complete noasaurid axial series known so far. Moreover, the new taxon exhibits many novel osteological features, uncommon in non-avian theropods, and unprecedented even among South American ceratosaurs. These include not only toothless jaws but also a premaxilla with cutting occlusal edge, and a slightly downturned rostral tip. This indicate that B. leopoldinae unlikely had the same diet as other ceratosaurs, most being regarded as carnivorous. As the ontogenetically more mature specimens of Limusaurus, Berthasaura might have been herbivorous or at least omnivorous, corroborating with an early evolutionary divergence of noasaurids from the ceratosaurian bauplan by disparate feeding modes.

Osteohistology and growth dynamics of the Brazilian noasaurid Vespersaurus paranaensis Langer et al., 2019 (Theropoda: Abelisauroidea).

Although the knowledge of bone histology of non-avian theropods has advanced considerably in recent decades, data about the bone tissue patterns, growth dynamics and ontogeny of some taxa such as abelisauroids are still limited. Here we describe the bone microstructure and growth dynamics of the Brazilian noasaurine Vespersaurus paranaensis using five femora and six tibiae and quantify the annual growth marks through retrocalculation of missing ones to estimate ontogenetic ages. The femoral series comprises four femoral histological classes (FHC I-IV), varying from two annuli or LAGs to seven LAGs. Femora show that sexual maturity was achieved around the seventh to tenth year of life, whereas the tibiae suggest it was earlier (around three to five years old). Tibiae represent three histological classes (THC I-III) displaying from three to nine LAGs. Two tibiae (THC III) exhibit an external fundamental system indicating that these specimens reached full skeletal size. The heterogeneous maturity observed in Vespersaurus hind limb bones could result from differential allometry scaling between femora and tibiae length with the body length. The predominant parallel-fibered bone matrix suggests that Vespersaurus grew more slowly than most theropods, including other abelisauroids, in a pattern shared with the noasaurines Masiakasaurus knopfleri from Madagascar and CPPLIP 1490 from Brazil. This deviation from the typical theropod growth pattern may be mainly correlated with small body size, but also may related to resource limitation imposed by the arid climate prevailing in southwestern Gondwana during Cretaceous. Moreover, given the ecological and phylogenetic similarities among these taxa, such features would probably be apomorphic within Noasauridae.

Early evidence of molariform hypsodonty in a Triassic stem-mammal.

Hypsodonty, the occurrence of high-crowned teeth, is widespread among mammals with diets rich in abrasive material, such as plants or soil, because it increases the durability of dentitions against wear. Hypsodont postcanine teeth evolved independently in multiple mammalian lineages and in the closely related mammaliaforms since the Jurassic period. Here, we report the oldest record, to our knowledge, of hypsodont postcanines in the non-mammaliaform stem-mammal, Menadon besairiei, from the early Late Triassic. The postcanines are long and columnar, with open roots. They were not replaced in older individuals and remained functional after the total wear of the crown enamel. Dental histology suggests that, convergently to hypsodont mammals, wear was compensated by the prolonged growth of each postcanine, resulting in dentine hypsodont teeth most similar to extant xenarthran mammals. These findings highlight the constraints imposed by limited tooth replacement and tooth wear in the evolutionary trajectories of herbivorous mammals and stem-mammals.

The African cynodont Aleodon (Cynodontia, Probainognathia) in the Triassic of southern Brazil and its biostratigraphic significance.

In this contribution we report the first occurrence of the enigmatic African probainognathian genus Aleodon in the Middle-early Late Triassic of several localities from the state of Rio Grande do Sul in southern Brazil. Aleodon is unusual among early probainognathians in having transversely-expanded postcanine teeth, similar to those of gomphodont cynognathians. This genus was previously known from the Manda Beds of Tanzania and the upper Omingonde Formation of Namibia. The Brazilian record of this genus is based upon multiple specimens representing different ontogenetic stages, including three that were previously referred to the sectorial-toothed probainognathian Chiniquodon theotonicus. We propose a new species of Aleodon (A. cromptoni sp. nov.) based on the specimens from Brazil. Additionally, we tentatively refer one specimen from the upper Omingonde Formation of Namibia to this new taxon, strengthening biostratigraphic correlations between these strata. Inclusion of A. cromptoni in a phylogenetic analysis of eucynodonts recovers it as the sister-taxon of A. brachyrhamphus within the family Chiniquodontidae. The discovery of numerous specimens of Aleodon among the supposedly monospecific Chiniquodon samples of Brazil raises concerns about chiniquodontid alpha taxonomy, particularly given the extremely broad geographic distribution of Chiniquodon. The discovery of Brazilian Aleodon and new records of the traversodontid Luangwa supports the hypothesis that at least two subzones can be recognized in the Dinodontosaurus Assemblage Zone.

Two New Cynodonts (Therapsida) from the Middle-Early Late Triassic of Brazil and Comments on South American Probainognathians.

We describe two new cynodonts from the early Late Triassic of southern Brazil. One taxon, Bonacynodon schultzi gen. et sp. nov., comes from the lower Carnian Dinodontosaurus AZ, being correlated with the faunal association at the upper half of the lower member of the Chañares Formation (Ischigualasto-Villa Unión Basin, Argentina). Phylogenetically, Bonacynodon is a closer relative to Probainognathus jenseni than to any other probainognathian, bearing conspicuous canines with a denticulate distal margin. The other new taxon is Santacruzgnathus abdalai gen. et sp. nov. from the Carnian Santacruzodon AZ. Although based exclusively on a partial lower jaw, it represents a probainognathian close to Prozostrodon from the Hyperodapedon AZ and to Brasilodon, Brasilitherium and Botucaraitherium from the Riograndia AZ. The two new cynodonts and the phylogenetic hypothesis presented herein indicate the degree to which our knowledge on probainognathian cynodonts is incomplete and also the relevance of the South American fossil record for understanding their evolutionary significance. The taxonomic diversity and abundance of probainognathians from Brazil and Argentina will form the basis of deep and complex studies to address the evolutionary transformations of cynodonts leading to mammals.

Dentary morphological variation in Clevosaurus brasiliensis (Rhynchocephalia, Clevosauridae) from the Upper Triassic of Rio Grande do Sul, Brazil.

Clevosaurus was a cosmopolitan rhynchocephalian genus, known from the Late Triassic to the Early Jurassic. In South America this genus is represented by C. brasiliensis, an important component of the Linha São Luiz taphocoenosis, on the top of the Norian Santa Maria 2 Sequence of Southern Brazil. The best preserved and most abundant bone elements of C. brasiliensis are dentaries, in which variations of shape and size are observed. The aim of this study is to describe and evaluate the variation, using geometric morphometrics methods. Geometric morphometric analysis of 10 specimens highlights variations in relative size of the dentary. Most of the variation observed for PC1 (83.3%) is likely related to ontogeny, and PC2 (10.0%) is likely related to taphonomic signatures. The development patterns observed, such as the growth of the dentary, consists of differential growth in length between the posterior portion of the dentary, that grows at a higher rate, regarding the anterior portion of the element. This allometric growth is similar to what is observed in other rhynchocephalians and is accompanied by the allometric skull growth, similar to the trend exhibited by clevosaurs. The taphocoenosis is bimodal (juveniles and adults) with a bias towards adult preservation. Some diagenetic influence is reflected in deformed skulls and this is observed in the tangent-plot. Finally, a strong correlation was detected between the taphonomic signatures and the PC2, regarding specially disarticulation and degree of fragmentation.