Comments on the pelvic girdle anatomy of Lagerpeton chanarensis Romer, 1971 (Archosauria) and its implications on the posture and gait of early pterosauromorphs.

Lagerpeton chanarensis is an early avemetatarsalian from the lower Carnian (lowermost Upper Triassic) levels of the Chañares Formation, La Rioja Province, Argentina. Lagerpeton and its kin were traditionally interpreted as dinosaur precursors of cursorial habits, with a bipedal posture and parasagittal gait. Some authors also speculated saltatorial capabilities for this genus. Recent analyses indicate that lagerpetids are early-diverging pterosauromorphs, a hypothesis that invites a review of most aspects of their anatomy and function. A revision of available specimens and additional preparation of previously known individuals indicate that Lagerpeton lacked a parasagittal gait and was probably a sprawling archosaur. This latter inference is based on the femoral head articulation with the acetabulum. The acetabular rim has a strongly laterally projected posteroventral antitrochanteric corner, which results in a position of the legs that recalls that of sprawling living reptiles, such as lizards, and departs from the parasagittally positioned limbs of dinosaurs. This may indicate that early pterosauromorphs had a sprawling posture of their hindlegs, casting doubts on the significance of bipedal posture and parasagittal gait for the radiation of early ornithodirans, given that both traits have been regarded as key features that triggered the ecological and evolutionary success of the clade. Our results bolster recent claims of a high ecomorphological diversity among early avemetatarsalians.

An unusually robust specimen attributed to Buriolestes schultzi (Dinosauria: Sauropodomorpha) from the Late Triassic of southern Brazil.

Buriolestes schultzi is a small sauropodomorph dinosaur from Carnian beds (ca., 233 Ma) of southern Brazil. It is one of the earliest members of that lineage and is a key taxon to investigate the initial evolution of Sauropodomorpha. Here, we attribute a new specimen to B. schultzi from Late Triassic of southern Brazil, which represents the first occurrence of the taxon outside the type locality. The new specimen comprises a disarticulated and partial skeleton, including cranial and postcranial elements. It is tentatively regarded as an additional specimen of B. schultzi according to a unique combination of traits (including autapomorphies). Conversely, the new specimen is stouter than the other specimens of B. schultzi, as shown by femoral Robustness Index. Based on femoral circumference, the estimated body mass of the new specimen is approximately 15 kg, which is far higher than the previous estimations for other specimens of B. schultzi (i.e., approximately 7 kg). In fact, the new specimen and some specimens of Eoraptor lunensis and Saturnalia tupiniquim were found to be significantly stouter than coeval sauropodomorphs. Therefore, instead of all being constructed as gracile, the earliest sauropodomorphs experienced an unappreciated intraspecific variation in robustness. This is interesting because more precise data on species body mass are crucial in order to better understand the complex terrestrial ecosystems in which dinosaurs originated.

A saurischian (Archosauria, Dinosauria) ilium from the Upper Triassic of southern Brazil and the rise of Herrerasauria.

The Carnian (Upper Triassic) rocks of the Candelária Sequence present a rich record of dinosaurs, including some of the oldest known dinosaurs worldwide. In this contribution we describe the first unequivocal dinosaur from the Pivetta site, located in the Restinga Sêca municipality, Southern Brazil. The specimen CAPPA/UFSM 0373 is an isolated but well-preserved left ilium. A thorough examination of the specimen's anatomy and a phylogenetic analysis provides evidence that CAPPA/UFSM 0373 belongs to the Herrerasauria. We were able to identify several similarities with potential non-herrerasaurid herrerasaurians (e.g., Tawa hallae, "Caseosaurus crosbyensis"), which were previously only known from North American deposits. In contrast, herrerasaurids (e.g., Herrerasaurus ischigualastensis) are almost exclusively known from South America. Our results support the nesting of CAPPA/UFSM 0373 as an early-diverging herrerasaurian. Furthermore, this is potentially the first record of a non-herrerasaurid herrerasaurian in unambiguous Carnian beds, suggesting a hidden diversity of dinosaurs in the Carnian rocks of the Candelária Sequence, which can be revealed even by fragmentary specimens.

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.

Development and evolution of the notarium in Pterosauria.

The notarium is the structure formed by fusion of the dorsal vertebrae which occurred independently in pterosaurs and birds. This ankylosis usually involves two to six elements and in many cases, also includes the last cervical vertebra. Fusion can occur in different degrees, uniting the vertebral centra, the neural spines, the transverse processes, the ventral processes, or a combination of these sites. A detailed assessment of the fusion process of pterosaur dorsal vertebrae is still lacking. Here we identify the fusion sequence of pterosaur notarial elements, demonstrating the order of ossification in vertebral bodies and neural spines based on fossils and extant birds. In both Pterosauria and Aves, the notarium generally develops in a antero-posterior direction, but the actual order of each fusion locus may present slight variations. Based on our data, we were able to identify seven developmental stages in the notarium formation, with broad implications for the prediction of ontogenetic stages for the Pterosauria. In addition, we report the occurrence of a notarium in Ardeadactylus longicollum (Kimmeridgian, Southern Germany), the oldest occurrence of this structure in pterosaurs.

A paraphyletic 'Silesauridae' as an alternative hypothesis for the initial radiation of ornithischian dinosaurs.

Whereas ornithischian dinosaurs are well known from Jurassic and Cretaceous deposits, deciphering the origin and early evolution of the group remains one of the hardest challenges for palaeontologists. So far, there are no unequivocal records of ornithischians from Triassic beds. Here, we present an alternative evolutionary hypothesis that suggests consideration of traditional 'silesaurids' as a group of low-diversity clades representing a stem group leading to core ornithischians (i.e. unambiguous ornithischians, such as Heterodontosaurus tucki). This is particularly interesting because it fills most of the ghost lineages that emerge from the Triassic. Following the present hypothesis, the lineage that encompasses the Jurassic ornithischians evolved from 'silesaurids' during the Middle to early Late Triassic, while typical 'silesaurids' shared the land ecosystems with their relatives until the Late Triassic, when the group completely vanished. Therefore, Ornithischia changes from an obscure to a well-documented clade in the Triassic and is represented by records from Gondwana and Laurasia. Furthermore, according to the present hypothesis, Ornithischia was the first group of dinosaurs to adopt an omnivorous/herbivorous diet. However, this behaviour was achieved as a secondary step instead of an ancestral condition for ornithischians, as the earliest member of the clade is a faunivorous taxon. This pattern was subsequently followed by sauropodomorph dinosaurs. Indeed, the present scenario favours the independent acquisition of an herbivorous diet for ornithischians and sauropodomorphs during the Triassic, whereas the previous hypotheses suggested the independent acquisition for sauropodomorphs, ornithischians, and 'silesaurids'.

An exceptionally preserved association of complete dinosaur skeletons reveals the oldest long-necked sauropodomorphs.

The rise of sauropodomorphs is still poorly understood due to the scarcity of well-preserved fossils in early Norian rocks. Here, we present an association of complete and exceptionally well-preserved dinosaur skeletons that helps fill that gap. They represent a new species, which is recovered as a member of a clade solely composed of Gondwanan Triassic taxa. The new species allows the definition of a set of anatomical changes that shaped sauropodomorph evolution along a period from 233 to 225 Ma, as recorded in the well dated Late Triassic beds of Brazil. In that time span, apart from achieving a more herbivorous diet, sauropodomorph dinosaurs increased their size in a ratio of 230% and their typical long neck was also established, becoming proportionally twice longer than those of basal taxa. Indeed, the new dinosaur is the oldest-known sauropodomorph with such an elongated neck, suggesting that the ability to feed on high vegetation was a key trait achieved along the early Norian. Finally, the clustered preservation mode of the skeletons represents the oldest evidence of gregarious behaviour among sauropodomorphs.

Skull anatomy and phylogenetic assessment of a large specimen of Ecteniniidae (Eucynodontia: Probainognathia) from the Upper Triassic of southern Brazil.

Ecteniniidae comprises an endemic radiation of carnivore probainognathian cynodonts from the Late Triassic of South America. Three taxa have been included in this clade: Ecteninion lunensis Martínez et al., 1996 and Diegocanis elegans Martínez et al., 2013 from Argentina, and Trucidocynodon riograndensis Oliveira et al., 2010 from Brazil. Herein, a new specimen (skull and mandible) assigned to T. riograndensis from the Carnian of the Candelária Sequence (Southern Brazil) is described. A phylogenetic analysis recovered the new specimen as the sister taxon of the holotype of T. riograndensis, and both in a trichotomy with E. lunensis and D. elegans, all supporting the monophyly of Ecteniniidae. The new specimen of T. riograndensis is almost 20% larger than its holotype. Therefore, it represents one of the largest specimens of a carnivorous probainognathian from the Late Triassic known to date and contributes to knowledge of size variation in ecteniniids.

Ingroup relationships of Lagerpetidae (Avemetatarsalia: Dinosauromorpha): a further phylogenetic investigation on the understanding of dinosaur relatives.

Despite representing a key-taxon in dinosauromorph phylogeny, Lagerpertidae is one of the most obscure and enigmatic branches from the stem that leads to the dinosaurs. Recent new findings have greatly increased our knowledge about lagerpetids, but no phylogenetic analysis has so far included all known members of this group. Here, we present the most inclusive phylogenetic study so far conducted for Lagerpetidae. Phylogenetic analyses were performed based on three independent data matrixes. In two of them, Lagerpeton chanarensis Romer, 1971 is the sister taxon to all other known Lagerpetidae, whereas Ixalerpeton polesinensis Cabreira et al., 2016 is in a sister group relationship with a clade that includes PVSJ 883 and Dromomeron. Conversely, the other analysis supports an alternative topology, where I. polesinensis is the sister taxon to either L. chanarensis or all other Lagerpetidae. Although coeval and geographically close, I. polesinensis and PVSJ 883 do not form a clade exclusive of other lagerpetids. As previously suggested D. gigas Martínez, Apaldetti, Correa Abelín, 2016 is the sister taxon of D. romeri Irmis et al., 2007. The phylogenetic analyses also indicate that the earliest lagerpetids are restricted to southwestern Pangea, whereas later forms spread across the entire western portion of the supercontinent. Finally, quantification of the codified characters of our analysis reveals that Lagerpetidae is one of the poorest known among the Triassic dinosauromorph groups in terms of their anatomy, so that new discoveries of more complete specimens are awaited to establish a more robust phylogeny.

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.