Braincase and neuroanatomy of the lagerpetid Dromomeron gregorii (Archosauria, Pterosauromorpha) with comments on the early evolution of the braincase and associated soft tissues in Avemetatarsalia.

The anatomy of the braincase and associated soft tissues of the lagerpetid Dromomeron gregorii (Archosauria: Avemetatarsalia) from the Late Triassic of the United States is here described. This corresponds to the first detailed description of cranial materials of Lagerpetidae, an enigmatic group of Late Triassic (c. 236-200 Million years ago) animals that are the closest known relatives of pterosaurs, the flying reptiles. The braincase of D. gregorii is characterized by the presence of an anteriorly elongated laterosphenoid and a postparietal, features observed in stem-archosaurs but that were still unknown in early members of the avian lineage of archosaurs. Using micro-computed tomography (CT-scan data), we present digital reconstructions of the brain and endosseous labyrinth of D. gregorii. The brain of D. gregorii exhibits a floccular lobe of the cerebellum that projects within the space of the semicircular canals. The semicircular canals are relatively large when compared to other archosauromorphs, with the anterior canal exhibiting a circular shape. These features of the sensory structures of D. gregorii are more similar to those of pterosaurs than to those of other early avemetatarsalians. In sum, the braincase anatomy of D. gregorii shows a combination of plesiomorphic and apomorphic features in the phylogenetic context of Archosauria and suggests that the still poorly understood early evolution of the braincase in avemetatarsalians is complex, with a scenario of independent acquisitions and losses of character states.

Comment on "The early origin of a birdlike inner ear and the evolution of dinosaurian movement and vocalization".

Hanson et al. (Research Articles, 7 May 2021, p. 601) claim that the shape of the vestibular apparatus reflects the evolution of reptilian locomotion. Using biomechanics, we demonstrate that semicircular canal shape is a dubious predictor of semicircular duct function. Additionally, we show that the inference methods used by Hanson et al. largely overestimate relationships between semicircular canal shape and locomotion.

Deep evolutionary diversification of semicircular canals in archosaurs.

Living archosaurs (birds and crocodylians) have disparate locomotor strategies that evolved since their divergence ∼250 mya. Little is known about the early evolution of the sensory structures that are coupled with these changes, mostly due to limited sampling of early fossils on key stem lineages. In particular, the morphology of the semicircular canals (SCCs) of the endosseous labyrinth has a long-hypothesized relationship with locomotion. Here, we analyze SCC shapes and sizes of living and extinct archosaurs encompassing diverse locomotor habits, including bipedal, semi-aquatic, and flying taxa. We test form-function hypotheses of the SCCs and chronicle their evolution during deep archosaurian divergences. We find that SCC shape is statistically associated with both flight and bipedalism. However, this shape variation is small and is more likely explained by changes in braincase geometry than by locomotor changes. We demonstrate high disparity of both shape and size among stem-archosaurs and a deep divergence of SCC morphologies at the bird-crocodylian split. Stem-crocodylians exhibit diverse morphologies, including aspects also present in birds and distinct from other reptiles. Therefore, extant crocodylian SCC morphologies do not reflect retention of a "primitive" reptilian condition. Key aspects of bird SCC morphology that hitherto were interpreted as flight related, including large SCC size and enhanced sensitivity, appeared early on the bird stem-lineage in non-flying dinosaur precursors. Taken together, our results indicate a deep divergence of SCC traits at the bird-crocodylian split and that living archosaurs evolved from an early radiation with high sensory diversity. VIDEO ABSTRACT.

Erratum: Cretaceous blind snake from Brazil fills major gap in snake evolution.

[This corrects the article DOI: 10.1016/j.isci.2020.101834.].

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.

Cretaceous Blind Snake from Brazil Fills Major Gap in Snake Evolution.

Blind snakes (Scolecophidia) are minute cryptic snakes that diverged at the base of the evolutionary radiation of modern snakes. They have a scant fossil record, which dates back to the Upper Paleocene-Lower Eocene (∼56 Ma); this late appearance conflicts with molecular evidence, which suggests a much older origin for the group (during the Mesozoic: 160-125 Ma). Here we report a typhlopoid blind snake from the Late Cretaceous of Brazil, Boipeba tayasuensis gen. et sp. nov, which extends the scolecophidian fossil record into the Mesozoic and reduces the fossil gap predicted by molecular data. The new species is estimated to have been over 1 m long, much larger than typical modern scolecophidians (<30 cm). This finding sheds light on the early evolution of blind snakes, supports the hypothesis of a Gondwanan origin for the Typhlopoidea, and indicates that early scolecophidians had large body size, and only later underwent miniaturization.

Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria.

Pterosaurs were the first vertebrates to evolve powered flight1 and comprised one of the main evolutionary radiations in terrestrial ecosystems of the Mesozoic era (approximately 252-66 million years ago), but their origin has remained an unresolved enigma in palaeontology since the nineteenth century2-4. These flying reptiles have been hypothesized to be the close relatives of a wide variety of reptilian clades, including dinosaur relatives2-8, and there is still a major morphological gap between those forms and the oldest, unambiguous pterosaurs from the Upper Triassic series. Here, using recent discoveries of well-preserved cranial remains, microcomputed tomography scans of fragile skull bones (jaws, skull roofs and braincases) and reliably associated postcrania, we demonstrate that lagerpetids-a group of cursorial, non-volant dinosaur precursors-are the sister group of pterosaurs, sharing numerous synapomorphies across the entire skeleton. This finding substantially shortens the temporal and morphological gap between the oldest pterosaurs and their closest relatives and simultaneously strengthens the evidence that pterosaurs belong to the avian line of archosaurs. Neuroanatomical features related to the enhanced sensory abilities of pterosaurs9 are already present in lagerpetids, which indicates that these features evolved before flight. Our evidence illuminates the first steps of the assembly of the pterosaur body plan, whose conquest of aerial space represents a remarkable morphofunctional innovation in vertebrate evolution.

Skull remains of the dinosaur Saturnalia tupiniquim (Late Triassic, Brazil): With comments on the early evolution of sauropodomorph feeding behaviour.

Saturnalia tupiniquim is a sauropodomorph dinosaur from the Late Triassic (Carnian-c. 233 Ma) Santa Maria Formation of Brazil. Due to its phylogenetic position and age, it is important for studies focusing on the early evolution of both dinosaurs and sauropodomorphs. The osteology of Saturnalia has been described in a series of papers, but its cranial anatomy remains mostly unknown. Here, we describe the skull bones of one of its paratypes (only in the type-series to possess such remains) based on CT Scan data. The newly described elements allowed estimating the cranial length of Saturnalia and provide additional support for the presence of a reduced skull (i.e. two thirds of the femoral length) in this taxon, as typical of later sauropodomorphs. Skull reduction in Saturnalia could be related to an increased efficiency for predatory feeding behaviour, allowing fast movements of the head in order to secure small and elusive prey, a hypothesis also supported by data from its tooth and brain morphology. A principal co-ordinates analysis of the sauropodomorph jaw feeding apparatus shows marked shifts in morphospace occupation in different stages of the first 30 million years of their evolutionary history. One of these shifts is observed between non-plateosaurian and plateosaurian sauropodomorphs, suggesting that, despite also having an omnivorous diet, the feeding behaviour of some early Carnian sauropodomorphs, such as Saturnalia, was markedly different from that of later Triassic taxa. A second shift, between Late Triassic and Early Jurassic taxa, is congruent with a floral turnover hypothesis across the Triassic-Jurassic boundary.

The multi-peak adaptive landscape of crocodylomorph body size evolution.

BACKGROUND: Little is known about the long-term patterns of body size evolution in Crocodylomorpha, the > 200-million-year-old group that includes living crocodylians and their extinct relatives. Extant crocodylians are mostly large-bodied (3-7 m) predators. However, extinct crocodylomorphs exhibit a wider range of phenotypes, and many of the earliest taxa were much smaller (< 1.2 m). This suggests a pattern of size increase through time that could be caused by multi-lineage evolutionary trends of size increase or by selective extinction of small-bodied species. Here, we characterise patterns of crocodylomorph body size evolution using a model fitting-approach (with cranial measurements serving as proxies). We also estimate body size disparity through time and quantitatively test hypotheses of biotic and abiotic factors as potential drivers of crocodylomorph body size evolution. RESULTS: Crocodylomorphs reached an early peak in body size disparity during the Late Jurassic, and underwent an essentially continual decline since then. A multi-peak Ornstein-Uhlenbeck model outperforms all other evolutionary models fitted to our data (including both uniform and non-uniform), indicating that the macroevolutionary dynamics of crocodylomorph body size are better described within the concept of an adaptive landscape, with most body size variation emerging after shifts to new macroevolutionary regimes (analogous to adaptive zones). We did not find support for a consistent evolutionary trend towards larger sizes among lineages (i.e., Cope's rule), or strong correlations of body size with climate. Instead, the intermediate to large body sizes of some crocodylomorphs are better explained by group-specific adaptations. In particular, the evolution of a more aquatic lifestyle (especially marine) correlates with increases in average body size, though not without exceptions. CONCLUSIONS: Shifts between macroevolutionary regimes provide a better explanation of crocodylomorph body size evolution on large phylogenetic and temporal scales, suggesting a central role for lineage-specific adaptations rather than climatic forcing. Shifts leading to larger body sizes occurred in most aquatic and semi-aquatic groups. This, combined with extinctions of groups occupying smaller body size regimes (particularly during the Late Cretaceous and Cenozoic), gave rise to the upward-shifted body size distribution of extant crocodylomorphs compared to their smaller-bodied terrestrial ancestors.

Phylogeny, biogeography and diversification patterns of side-necked turtles (Testudines: Pleurodira).

Pleurodires or side-necked turtles are today restricted to freshwater environments of South America, Africa-Madagascar and Australia, but in the past they were distributed much more broadly, being found also on Eurasia, India and North America, and marine environments. Two hypotheses were proposed to explain this distribution; in the first, vicariance would have shaped the current geographical distribution and, in the second, extinctions constrained a previously widespread distribution. Here, we aim to reconstruct pleurodiran biogeographic history and diversification patterns based on a new phylogenetic hypothesis recovered from the analysis of the largest morphological dataset yet compiled for the lineage, testing which biogeographical process prevailed during its evolutionary history. The resulting topology generally agrees with previous hypotheses of the group and shows that most diversification shifts were related to the exploration of new niches, e.g. littoral or marine radiations. In addition, as other turtles, pleurodires do not seem to have been much affected by either the Cretaceous-Palaeogene or the Eocene-Oligocene mass extinctions. The biogeographic analyses highlight the predominance of both anagenetic and cladogenetic dispersal events and support the importance of transoceanic dispersals as a more common driver of area changes than previously thought, agreeing with previous studies with other non-turtle lineages.

Endocast of the Late Triassic (Carnian) dinosaur Saturnalia tupiniquim: implications for the evolution of brain tissue in Sauropodomorpha.

The evolutionary history of dinosaurs might date back to the first stages of the Triassic (c. 250-240 Ma), but the oldest unequivocal records of the group come from Late Triassic (Carnian - c. 230 Ma) rocks of South America. Here, we present the first braincase endocast of a Carnian dinosaur, the sauropodomorph Saturnalia tupiniquim, and provide new data regarding the evolution of the floccular and parafloccular lobe of the cerebellum (FFL), which has been extensively discussed in the field of palaeoneurology. Previous studies proposed that the development of a permanent quadrupedal stance was one of the factors leading to the volume reduction of the FFL of sauropods. However, based on the new data for S. tupiniquim we identified a first moment of FFL volume reduction in non-sauropodan Sauropodomorpha, preceding the acquisition of a fully quadrupedal stance. Analysing variations in FFL volume alongside other morphological changes in the group, we suggest that this reduction is potentially related to the adoption of a more restricted herbivore diet. In this context, the FFL of sauropods might represent a vestigial trait, retained in a reduced version from the bipedal and predatory early sauropodomorphs.

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.

Postcranial anatomy of Pissarrachampsa sera (Crocodyliformes, Baurusuchidae) from the Late Cretaceous of Brazil: insights on lifestyle and phylogenetic significance.

The postcranial anatomy of Crocodyliformes has historically been neglected, as most descriptions are based solely on skulls. Yet, the significance of the postcranium in crocodyliforms evolution is reflected in the great lifestyle diversity exhibited by the group, with members ranging from terrestrial animals to semi-aquatic and fully marine forms. Recently, studies have emphasized the importance of the postcranium. Following this trend, here we present a detailed description of the postcranial elements of Pissarrachampsa sera (Mesoeucrocodylia, Baurusuchidae), from the Adamantina Formation (Bauru Group, Late Cretaceous of Brazil). The preserved elements include dorsal vertebrae, partial forelimb, pelvic girdle, and hindlimbs. Comparisons with the postcranial anatomy of baurusuchids and other crocodyliforms, together with body-size and mass estimates, lead to a better understanding of the paleobiology of Pissarrachampsa sera, including its terrestrial lifestyle and its role as a top predator. Furthermore, the complete absence of osteoderms in P. sera, a condition previously known only in marine crocodyliforms, suggests osteoderms very likely played a minor role in locomotion of baurusuchids, unlike other groups of terrestrial crocodyliforms. Finally, a phylogenetic analysis including the newly recognized postcranial features was carried out, and exploratory analyses were performed to investigate the influence of both cranial and postcranial characters in the phylogeny of Crocodyliformes. Our results suggest that crocodyliform relationships are mainly determined by cranial characters. However, this seems to be a consequence of the great number of missing entries in the data set with only postcranial characters and not of the lack of potential (or synapomorphies) for this kind of data to reflect the evolutionary history of Crocodyliformes.

Diversification events and the effects of mass extinctions on Crocodyliformes evolutionary history.

The rich fossil record of Crocodyliformes shows a much greater diversity in the past than today in terms of morphological disparity and occupation of niches. We conducted topology-based analyses seeking diversification shifts along the evolutionary history of the group. Our results support previous studies, indicating an initial radiation of the group following the Triassic/Jurassic mass extinction, here assumed to be related to the diversification of terrestrial protosuchians, marine thalattosuchians and semi-aquatic lineages within Neosuchia. During the Cretaceous, notosuchians embodied a second diversification event in terrestrial habitats and eusuchian lineages started diversifying before the end of the Mesozoic. Our results also support previous arguments for a minor impact of the Cretaceous/Palaeogene mass extinction on the evolutionary history of the group. This argument is not only based on the information from the fossil record, which shows basal groups surviving the mass extinction and the decline of other Mesozoic lineages before the event, but also by the diversification event encompassing only the alligatoroids in the earliest period after the extinction. Our results also indicate that, instead of a continuous process through time, Crocodyliformes diversification was patchy, with events restricted to specific subgroups in particular environments and time intervals.