New reptile shows dinosaurs and pterosaurs evolved among diverse precursors.

Dinosaurs and pterosaurs have remarkable diversity and disparity through most of the Mesozoic Era1-3. Soon after their origins, these reptiles diversified into a number of long-lived lineages, evolved unprecedented ecologies (for example, flying, large herbivorous forms) and spread across Pangaea4,5. Recent discoveries of dinosaur and pterosaur precursors6-10 demonstrated that these animals were also speciose and widespread, but those precursors have few if any well-preserved skulls, hands and associated skeletons11,12. Here we present a well-preserved partial skeleton (Upper Triassic, Brazil) of the new lagerpetid Venetoraptor gassenae gen. et sp. nov. that offers a more comprehensive look into the skull and ecology of one of these precursors. Its skull has a sharp, raptorial-like beak, preceding that of dinosaurs by around 80 million years, and a large hand with long, trenchant claws that firmly establishes the loss of obligatory quadrupedalism in these precursor lineages. Combining anatomical information of the new species with other dinosaur and pterosaur precursors shows that morphological disparity of precursors resembles that of Triassic pterosaurs and exceeds that of Triassic dinosaurs. Thus, the 'success' of pterosaurs and dinosaurs was a result of differential survival among a broader pool of ecomorphological variation. Our results show that the morphological diversity of ornithodirans started to flourish among early-diverging lineages and not only after the origins of dinosaurs and pterosaurs.

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.

Exploring the effects of weighting against homoplasy in genealogies of palaeontological phylogenetic matrices.

Although simulations have shown that implied weighting (IW) outperforms equal weighting (EW) in phylogenetic parsimony analyses, weighting against homoplasy lacks extensive usage in palaeontology. Iterative modifications of several phylogenetic matrices in the last decades resulted in extensive genealogies of datasets that allow the evaluation of differences in the stability of results for alternative character weighting methods directly on empirical data. Each generation was compared against the most recent generation in each genealogy because it is assumed that it is the most comprehensive (higher sampling), revised (fewer misscorings) and complete (lower amount of missing data) matrix of the genealogy. The analyses were conducted on six different genealogies under EW and IW and extended implied weighting (EIW) with a range of concavity constant values (k) between 3 and 30. Pairwise comparisons between trees were conducted using Robinson-Foulds distances normalized by the total number of groups, distortion coefficient, subtree pruning and regrafting moves, and the proportional sum of group dissimilarities. The results consistently show that IW and EIW produce results more similar to those of the last dataset than EW in the vast majority of genealogies and for all comparative measures. This is significant because almost all of these matrices were originally analysed only under EW. Implied weighting and EIW do not outperform each other unambiguously. Euclidean distances based on a principal components analysis of the comparative measures show that different ranges of k-values retrieve the most similar results to the last generation in different genealogies. There is a significant positive linear correlation between the optimal k-values and the number of terminals of the last generations. This could be employed to inform about the range of k-values to be used in phylogenetic analyses based on matrix size but with the caveat that this emergent relationship still relies on a low sample size of genealogies.

The earliest bird-line archosaurs and the assembly of the dinosaur body plan.

The relationship between dinosaurs and other reptiles is well established, but the sequence of acquisition of dinosaurian features has been obscured by the scarcity of fossils with transitional morphologies. The closest extinct relatives of dinosaurs either have highly derived morphologies or are known from poorly preserved or incomplete material. Here we describe one of the stratigraphically lowest and phylogenetically earliest members of the avian stem lineage (Avemetatarsalia), Teleocrater rhadinus gen. et sp. nov., from the Middle Triassic epoch. The anatomy of T. rhadinus provides key information that unites several enigmatic taxa from across Pangaea into a previously unrecognized clade, Aphanosauria. This clade is the sister taxon of Ornithodira (pterosaurs and birds) and shortens the ghost lineage inferred at the base of Avemetatarsalia. We demonstrate that several anatomical features long thought to characterize Dinosauria and dinosauriforms evolved much earlier, soon after the bird-crocodylian split, and that the earliest avemetatarsalians retained the crocodylian-like ankle morphology and hindlimb proportions of stem archosaurs and early pseudosuchians. Early avemetatarsalians were substantially more species-rich, widely geographically distributed and morphologically diverse than previously recognized. Moreover, several early dinosauromorphs that were previously used as models to understand dinosaur origins may represent specialized forms rather than the ancestral avemetatarsalian morphology.

An enigmatic plant-eating theropod from the Late Jurassic period of Chile.

Theropod dinosaurs were the dominant predators in most Mesozoic era terrestrial ecosystems. Early theropod evolution is currently interpreted as the diversification of various carnivorous and cursorial taxa, whereas the acquisition of herbivorism, together with the secondary loss of cursorial adaptations, occurred much later among advanced coelurosaurian theropods. A new, bizarre herbivorous basal tetanuran from the Upper Jurassic of Chile challenges this conception. The new dinosaur was discovered at Aysén, a fossil locality in the Upper Jurassic Toqui Formation of southern Chile (General Carrera Lake). The site yielded abundant and exquisitely preserved three-dimensional skeletons of small archosaurs. Several articulated individuals of Chilesaurus at different ontogenetic stages have been collected, as well as less abundant basal crocodyliforms, and fragmentary remains of sauropod dinosaurs (diplodocids and titanosaurians).

The rise of the ruling reptiles and ecosystem recovery from the Permo-Triassic mass extinction.

One of the key faunal transitions in Earth history occurred after the Permo-Triassic mass extinction (ca 252.2 Ma), when the previously obscure archosauromorphs (which include crocodylians, dinosaurs and birds) become the dominant terrestrial vertebrates. Here, we place all known middle Permian-early Late Triassic archosauromorph species into an explicit phylogenetic context, and quantify biodiversity change through this interval. Our results indicate the following sequence of diversification: a morphologically conservative and globally distributed post-extinction 'disaster fauna'; a major but cryptic and poorly sampled phylogenetic diversification with significantly elevated evolutionary rates; and a marked increase in species counts, abundance, and disparity contemporaneous with global ecosystem stabilization some 5 million years after the extinction. This multiphase event transformed global ecosystems, with far-reaching consequences for Mesozoic and modern faunas.

A new Palaeocene crocodylian from southern Argentina sheds light on the early history of caimanines.

Caimanines are crocodylians currently restricted to South and Central America and the oldest members are from lower Palaeocene localities of the Salamanca Formation (Chubut Province, Argentina). We report here a new caimanine from this same unit represented by a skull roof and partial braincase. Its phylogenetic relationships were explored in a cladistic analysis using standard characters and a morphogeometric two-dimensional configuration of the skull roof. The phylogenetic results were used for an event-based supermodel quantitative palaeobiogeographic analysis. The new species is recovered as the most basal member of the South American caimanines, and the Cretaceous North American lineage 'Brachychampsa and related forms' as the most basal Caimaninae. The biogeographic results estimated north-central North America as the ancestral area of Caimaninae, showing that the Cretaceous and Palaeocene species of the group were more widespread than thought and became regionally extinct in North America around the Cretaceous-Palaeocene boundary. A dispersal event from north-central North America during the middle Late Cretaceous explains the arrival of the group to South America. The Palaeogene assemblage of Patagonian crocodylians is composed of three lineages of caimanines as a consequence of independent dispersal events that occurred between North and South America and within South America around the Cretaceous-Palaeogene boundary.

Unappreciated diversification of stem archosaurs during the Middle Triassic predated the dominance of dinosaurs.

BACKGROUND: Archosauromorpha originated in the middle-late Permian, radiated during the Triassic, and gave rise to the crown group Archosauria, a highly successful clade of reptiles in terrestrial ecosystems over the last 250 million years. However, scientific attention has mainly focused on the diversification of archosaurs, while their stem lineage (i.e. non-archosaurian archosauromorphs) has often been overlooked in discussions of the evolutionary success of Archosauria. Here, we analyse the cranial disparity of late Permian to Early Jurassic archosauromorphs and make comparisons between non-archosaurian archosauromorphs and archosaurs (including Pseudosuchia and Ornithodira) on the basis of two-dimensional geometric morphometrics. RESULTS: Our analysis recovers previously unappreciated high morphological disparity for non-archosaurian archosauromorphs, especially during the Middle Triassic, which abruptly declined during the early Late Triassic (Carnian). By contrast, cranial disparity of archosaurs increased from the Middle Triassic into the Late Triassic, declined during the end-Triassic extinction, but re-expanded towards the end of the Early Jurassic. CONCLUSIONS: Our study indicates that non-archosaurian archosauromorphs were highly diverse components of terrestrial ecosystems prior to the major radiation of archosaurs, including dinosaurs, while disparity patterns of the Ladinian and Carnian indicate a gradual faunal replacement of stem archosaurs by the crown group, including a short interval of partial overlap in morphospace during the Ladinian.

Post-hatchling cranial ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi.

The phylogenetic position of Proterosuchus fergusi (Lower Triassic of South Africa) as one of the most basal archosauriforms means that it is critically important for understanding the successful evolutionary radiation of archosaurs during the Mesozoic. The excellent sample of the species provides a unique opportunity to understand early archosauriform ontogeny. Qualitative and quantitative analyses of cranial ontogenetic variation were conducted on an ontogenetic sequence, in which the smallest individual is 37% of the size of the largest one and osteohistological evidence suggests that four of 11 collected specimens had not reached sexual maturity. Through ontogeny the skull of Proterosuchus became proportionally taller, the infratemporal fenestra larger, and the teeth more isodont and numerous but with smaller crowns. The sequence of somatic maturity supports relatively high growth rates during early ontogeny. The skull of juvenile specimens of Proterosuchus closely resembles adults of the basal archosauromorph Prolacerta, whereas adult specimens resemble adults of more derived archosauriforms. As a result, a plausible hypothesis is that ontogenetic modification events (e.g., heterochrony) may have been key drivers of the evolution of the general shape of the skull at the base of Archosauriformes. These changes may have contributed to the occupation of a new morphospace by the clade around the Permo-Triassic boundary.

New clade of enigmatic early archosaurs yields insights into early pseudosuchian phylogeny and the biogeography of the archosaur radiation.

BACKGROUND: The origin and early radiation of archosaurs and closely related taxa (Archosauriformes) during the Triassic was a critical event in the evolutionary history of tetrapods. This radiation led to the dinosaur-dominated ecosystems of the Jurassic and Cretaceous, and the high present-day archosaur diversity that includes around 10,000 bird and crocodylian species. The timing and dynamics of this evolutionary radiation are currently obscured by the poorly constrained phylogenetic positions of several key early archosauriform taxa, including several species from the Middle Triassic of Argentina (Gracilisuchus stipanicicorum) and China (Turfanosuchus dabanensis, Yonghesuchus sangbiensis). These species act as unstable 'wildcards' in morphological phylogenetic analyses, reducing phylogenetic resolution. RESULTS: We present new anatomical data for the type specimens of G. stipanicicorum, T. dabanensis, and Y. sangbiensis, and carry out a new morphological phylogenetic analysis of early archosaur relationships. Our results indicate that these three previously enigmatic taxa form a well-supported clade of Middle Triassic archosaurs that we refer to as Gracilisuchidae. Gracilisuchidae is placed basally within Suchia, among the pseudosuchian (crocodile-line) archosaurs. The approximately contemporaneous and morphologically similar G. stipanicicorum and Y. sangbiensis may be sister taxa within Gracilisuchidae. CONCLUSIONS: Our results provide increased resolution of the previously poorly constrained relationships of early archosaurs, with increased levels of phylogenetic support for several key early pseudosuchian clades. Moreover, they falsify previous hypotheses suggesting that T. dabanensis and Y. sangbiensis are not members of the archosaur crown group. The recognition of Gracilisuchidae provides further support for a rapid phylogenetic diversification of crown archosaurs by the Middle Triassic. The disjunct distribution of the gracilisuchid clade in China and Argentina demonstrates that early archosaurs were distributed over much or all of Pangaea although they may have initially been relatively rare members of faunal assemblages.

The redescription of Malerisaurus robinsonae (Archosauromorpha: Allokotosauria) from the Upper Triassic lower Maleri Formation, Pranhita-Godavari Basin, India.

Allokotosauria, a clade of non-archosauriform archosauromorphs with a broad diversity of body plans, plays a crucial role in better understanding the evolutionary history of early diverging stem-archosaurs. Here we provide a detailed redescription of Malerisaurus robinsonae, a malerisaurine allokotosaur from the middle Carnian-lowermost Norian lower Maleri Formation, Pranhita-Godavari Basin, India. The new anatomical information available from recently discovered and well-preserved skeletons of various allokotosaurs, such as Azendohsaurus madagaskarensis, Shringasaurus indicus, Puercosuchus traverorum, and Malerisaurus-like taxa, and their comparison with Malerisaurus robinsonae enriches our understanding of the anatomy of this species. To reassess the phylogenetic relationships of Malerisaurus robinsonae, we revised its scorings and included eight additional allokotosaurian species to the already most comprehensive phylogenetic dataset focused on Permo-Triassic archosauromorphs. We modified 70 scorings for Malerisaurus robinsonae and the new analysis recovered this species at the base of Malerisaurinae and this group as the earliest branch of Azendohsauridae. Pamelaria dolichotrachela is found as the earliest diverging non-malerisaurine azendohsaurid and sister taxon to the Shringasaurus indicus + Azendohsaurus spp. clade. Trilophosaurid interrelationships are well resolved, with Teraterpeton hrynewichorum, Coelodontognathus ricovi, and Rutiotomodon tytthos as their successive earliest-branching species. The position of Anisodontosaurus greeri as a sister taxon to Variodens inopinatus bolsters long ghost lineages in the Late Triassic trilophosaurid record. A disparity analysis of tooth crown morphology shows that Allokotosauria is the most disparate Permo-Triassic archosauromorph clade, exploring the almost complete range of basic crown morphologies. Trilophosaurids occupy an area of the dental morphospace unique among archosauromorphs.

Osteology of the sauropodomorph dinosaur Jaklapallisaurus asymmetricus from the Late Triassic of central India.

The Gondwana formations exposed in the Pranhita-Godavari Valley of central India include Middle Triassic to Lower Jurassic continental deposits that provide essential information about the tetrapod assemblages of that time, documenting some of the oldest known dinosaurs and the first faunas numerically dominated by this group. The Upper Maleri Formation of the Pranhita-Godavari Basin preserves an early-middle Norian dinosaur assemblage that provides information about the early evolutionary history of this group in central-south Gondwana. This assemblage comprises sauropodomorph dinosaurs and an herrerasaurian, including two nominal species. Here, we describe in detail the anatomy of one of those early dinosaurs, the bagualosaurian sauropodomorph Jaklapallisaurus asymmetricus. The new anatomical information is used to investigate the position of the species in an updated quantitative phylogenetic analysis focused on early sauropodomorphs. The analysis recovered Jaklapallisaurus asymmetricus as a member of Unaysauridae, at the base of Plateosauria, together with Macrocollum itaquii and Unaysaurus tolentinoi from the early Norian of southern Brazil. This phylogenetic result indicates that the dispersal of early plateosaurian sauropodomorphs between the Southern Hemisphere and what nowadays is Europe would have occurred shortly after Ischigualastian times because of the extension of their ghost lineage. Thus, the presence of early plateosaurians in the early Norian of South America and India reduces a previously inferred diachrony between the biogeographic dispersals of theropods and sauropodomorphs during post-Ischigualastian times.

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.

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.

A new early-diverging probainognathian cynodont and a revision of the occurrence of cf. Aleodon from the Chañares Formation, northwestern Argentina: New clues on the faunistic composition of the latest Middle-?earliest Late Triassic Tarjadia Assemblage Zone.

The Chañares Formation (Ischigualasto-Villa Unión Basin) is worldwide known by its exquisitely preserved fossil record of latest Middle-to-early Late Triassic tetrapods, including erpetosuchids, "rauisuchians," proterochampsids, gracilisuchids, dinosauromorphs, pterosauromorphs, kannemeyeriiform dicynodonts, and traversodontid, chiniquodontid and probainognathid cynodonts, coming from the Tarjadia (bottom) and Massetognathus-Chanaresuchus (top) Assemblage Zones of its lower member. Regarding cynodonts, most of its profuse knowledge comes from the traditional layers discovered by Alfred Romer and his team in the 1960s that are now enclosed in the Massetognathus-Chanaresuchus Assemblage Zone (AZ). In this contribution we focus our study on the probainognathian cynodonts discovered in levels of the Tarjadia Assemblage Zone. We describe a new chiniquodontid cynodont with transversely broad postcanine teeth (Riojanodon nenoi gen. et sp. nov.) which is related to the genus Aleodon. In addition, the specimen CRILAR-Pv 567 previously referred to cf. Aleodon is here described, compared, and included in a phylogenetic analysis. It is considered as an indeterminate Aleodontinae nov., a clade here proposed to included chiniquodontids with transversely broad upper and lower postcanines, by having a cuspidated sectorial labial margin and a lingual platform that is twice broader than a lingual cingulum. Cromptodon mamiferoides, from the Cerro de Las Cabras Formation (Cuyo Basin), was also included in the phylogenetic analysis and recovered as an Aleodontinae. The new cynodont and the record of Aleodontinae indet. reinforce the faunal differentiation between the Tarjadia and Massetognathus-Chanaresuchus Assemblage Zones, in the lower member of the Chañares Formation, and inform on the diverse chiniquodontid clade with both sectorial and transversely broad postcanine teeth.

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.

A new global palaeobiogeographical model for the late Mesozoic and early Tertiary.

Late Mesozoic palaeobiogeography has been characterized by a distinction between the northern territories of Laurasia and the southern landmasses of Gondwana. The repeated discovery of Gondwanan lineages in Laurasia has led to the proposal of alternative scenarios to explain these anomalous occurrences. A new biogeographical model for late Mesozoic terrestrial ecosystems is here proposed in which Europe and "Gondwanan" territories possessed a common Eurogondwanan fauna during the earliest Cretaceous. Subsequently, following the Hauterivian, the European territories severed from Africa and then connected to Asiamerica resulting in a faunal interchange. This model explains the presence of Gondwanan taxa in Laurasia and the absence of Laurasian forms in the southern territories during the Cretaceous. In order to test this new palaeobiogeographical model, tree reconciliation analyses (TRAs) were performed based on biogeographical signals provided by a supertree of late Mesozoic archosaurs. The TRAs found significant evidence for the presence of an earliest Cretaceous Eurogondwanan fauna followed by a relatively short-term Gondwana-Laurasia dichotomy. The analysis recovered evidence for a biogeographical reconnection of the European territories with Africa and South America-Antarctica during the Campanian to Maastrichtian time-slice. This biogeographical scenario appears to continue through the early Tertiary and sheds light on the trans-Atlantic disjunct distributions of several extant plant and animal groups.

The better to eat you with: morphological disparity and enamel ultrastructure in odontocetes.

Variations in the shape and size of teeth have been associated with changes in enamel ultrastructure across odontocetes. Characterizing these features in extinct taxa can elucidate their functional morphology and feeding strategy, while also shedding light into macroevolutionary patterns during the evolutionary history of cetaceans. This study aimed to (1) describe the enamel and dentine ultrastructure of the Early Miocene odontocetes Notocetus vanbenedeni and Phoberodon arctirostris from Patagonia (Argentina) and (2) quantify tooth and enamel ultrastructure morphological disparity among odontocetes. Enamel was predominantly prismatic, thin in the anterior tooth of N. vanbenedeni and P. arctirostris; whilst thick on the posterior tooth of N. vanbenedeni. Together with skull morphology, data suggests a raptorial feeding strategy for P. arctirostris and a combination suction feeding method for N. vanbenedeni. Statistical analyses supported these inferences, indicating that enamel characters are useful for paleoecological research. Morphological disparity analyses showed that extant odontocetes occupy a larger morphospace and have more disparate morphologies, whilst extinct odontocetes were more similar among each other than with the extant group. There was no clear phylogenetic-based grouping, suggesting that tooth and enamel ultrastructure disparity were mainly driven by ecological pressures. These results highlight enamel ultrastructure as a source for broader-scale paleoecological studies in cetaceans.

A new archosauriform species from the Panchet Formation of India and the diversification of Proterosuchidae after the end-Permian mass extinction.

Proterosuchidae represents the oldest substantial diversification of Archosauromorpha and plays a key role in understanding the biotic recovery after the end-Permian mass extinction. Proterosuchidae was long treated as a wastebasket taxon, but recent revisions have reduced its taxonomic content to five valid species from the latest Permian of Russia and the earliest Triassic (Induan) of South Africa and China. In addition to these occurrences, several isolated proterosuchid bones have been reported from the Induan Panchet Formation of India for over 150 years. Following the re-study of historical specimens and newly collected material from this unit, we erect the new proterosuchid species Samsarasuchus pamelae, which is represented by most of the presacral vertebral column. We also describe cf. proterosuchid and proterosuchid cranial, girdle and limb bones that are not referred to Samsarasuchus pamelae. Phylogenetic analyses recovered Samsarasuchus pamelae within the new proterosuchid clade Chasmatosuchinae. The taxonomic diversity of Proterosuchidae is substantially expanded here, with at least 11 nominal species and several currently unnamed specimens, and a biogeographical range encompassing present-day South Africa, China, Russia, India, Brazil, Uruguay and Australia. This indicates a broader taxonomic, phylogenetic and biogeographic diversification of Proterosuchidae than previously thought in the aftermath of the end-Permian mass extinction.