Live birth has evolved many times independently in vertebrates, such as mammals and diverse groups of lizards and snakes. However, live birth is unknown in the major clade Archosauromorpha, a group that first evolved some 260 million years ago and is represented today by birds and crocodilians. Here we report the discovery of a pregnant long-necked marine reptile (Dinocephalosaurus) from the Middle Triassic (∼245 million years ago) of southwest China showing live birth in archosauromorphs. Our discovery pushes back evidence of reproductive biology in the clade by roughly 50 million years, and shows that there is no fundamental reason that archosauromorphs could not achieve live birth. Our phylogenetic models indicate that Dinocephalosaurus determined the sex of their offspring by sex chromosomes rather than by environmental temperature like crocodilians. Our results provide crucial evidence for genotypic sex determination facilitating land-water transitions in amniotes.
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.
The vertebrate recovery after the end-Permian mass extinction can be approached through the ichnological record, which is much more abundant than body fossils. The late Olenekian (Early Triassic) tetrapod ichnoassemblage of the Catalan Pyrenean Basin is the most complete and diverse of this age from Western Tethys. This extensional basin, composed of several depocenters, was formed in the latest phases of the Variscan orogeny (Pangea breakup) and was infilled by braided and meandering fluvial systems of the red-beds Buntsandstein facies. Abundant and diverse tetrapod ichnites are recorded in these facies, including Prorotodactylus mesaxonichnus isp. nov. (tracks possibly produced by euparkeriids), cf. Rotodactylus, at least two large chirotheriid morphotypes (archosauriform trackmakers), Rhynchosauroides cf. schochardti, two other undetermined Rhynchosauroides forms, an undetermined Morphotype A (archosauromorph trackmakers) and two types of Characichnos isp. (swimming traces, here associated to archosauromorph trackmakers). The Pyrenean ichnoassemblage suggests a relatively homogeneous ichnofaunal composition through the late Early Triassic of Central Pangea, characterized by the presence of Prorotodactylus and Rotodactylus. Small archosauromorph tracks dominate and present a wide distribution through the different fluviatile facies of the Triassic Pyrenean Basin, with large archosaurian footprints being present in a lesser degree. Archosauromorphs radiated and diversified through the Triassic vertebrate recovery, which ultimately lead to the archosaur and dinosaur dominance of the Mesozoic.
The rise of dinosaurs was a major event in vertebrate history, but the timing of the origin and early diversification of the group remain poorly constrained. Here, we describe Nyasasaurus parringtoni gen. et sp. nov., which is identified as either the earliest known member of, or the sister-taxon to, Dinosauria. Nyasasaurus possesses a unique combination of dinosaur character states and an elevated growth rate similar to that of definitive early dinosaurs. It demonstrates that the initial dinosaur radiation occurred over a longer timescale than previously thought (possibly 15 Myr earlier), and that dinosaurs and their immediate relatives are better understood as part of a larger Middle Triassic archosauriform radiation. The African provenance of Nyasasaurus supports a southern Pangaean origin for Dinosauria.
Similarities in body plan evolution, such as wings in pterosaurs, birds, and bats or limblessness in snakes and amphisbaenians, can be recognized as classical examples of convergence among animals [1-3]. We introduce a new Triassic stem archosaur that is unexpectedly and remarkably convergent with the “dome-headed” pachycephalosaur dinosaurs that lived over 100 million years later. Surprisingly, numerous additional taxa in the same assemblage (the Otis Chalk assemblage from the Dockum Group of Texas) demonstrate the early acquisition of morphological novelties that were later convergently evolved by post-Triassic dinosaurs. As one of the most successful clades of terrestrial vertebrates, dinosaurs came to occupy an extensive morphospace throughout their diversification in the Mesozoic Era [4, 5], but their distant relatives were first to evolve many of those “dinosaurian” body plans in the Triassic Period [6-8]. Our analysis of convergence between archosauromorphs from the Triassic Period and post-Triassic archosaurs demonstrates the early and extensive exploration of morphospace captured in a single Late Triassic assemblage, and we hypothesize that many of the “novel” morphotypes interpreted to occur among archosaurs later in the Mesozoic already were in place during the initial Triassic archosauromorph, largely non-dinosaurian, radiation and only later convergently evolved in diverse dinosaurian lineages.
The holotype and only known specimen of the enigmatic small reptile Elachistosuchus huenei Janensch, 1949 from the Upper Triassic (Norian) Arnstadt Formation of Saxony-Anhalt (Germany) is redescribed using μCT scans of the material. This re-examination revealed new information on the morphology of this taxon, including previously unknown parts of the skeleton such as the palate, braincase, and shoulder girdle. Elachistosuchus is diagnosed especially by the presence of the posterolateral process of the frontal, the extension of the maxillary tooth row to the posterior margin of the orbit, the free posterior process of the jugal, and the notched anterior margin of the interclavicle. Phylogenetic analyses using two recently published character-taxon matrices recovered conflicting results for the phylogenetic position of Elachistosuchus-either as an archosauromorph, as a lepidosauromorph or as a more basal, non-saurian diapsid. These different placements highlight the need of a thorough revision of critical taxa and new character sets used for inferring neodiapsid relationships.
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.
Macelognathus vagans Marsh, 1884 from the Late Jurassic Morrison Fm. of Wyoming was originally described as a dinosaur by Marsh and in 1971 Ostrom suggested crocodilian affinities. In 2005, Göhlich and collaborators identified new material of this species from Colorado as a basal crocodylomorph. However, a partial skull found in association with mandibular and postcranial remains was not described.
Proterosuchidae are the most taxonomically diverse archosauromorph reptiles sampled in the immediate aftermath of the Permo-Triassic mass extinction and represent the earliest radiation of Archosauriformes (archosaurs and closely related species). Proterosuchids are potentially represented by approximately 15 nominal species collected from South Africa, China, Russia, Australia and India, but the taxonomic content of the group is currently in a state of flux because of the poor anatomic and systematic information available for several of its putative members. Here, the putative proterosuchid Tasmaniosaurus triassicus from the Lower Triassic of Hobart, Tasmania (Australia), is redescribed. The holotype and currently only known specimen includes cranial and postcranial remains and the revision of this material sheds new light on the anatomy of the animal, including new data on the cranial endocast. Several bones are re-identified or reinterpreted, contrasting with the descriptions of previous authors. The new information provided here shows that Tasmaniosaurus closely resembles the South African proterosuchid Proterosuchus, but it differed in the presence of, for example, a slightly downturned premaxilla, a shorter anterior process of maxilla, and a diamond-shaped anterior end of interclavicle. Previous claims for the presence of gut contents in the holotype of Tasmaniosaurus are considered ambiguous. The description of the cranial endocast of Tasmaniosaurus provides for the first time information about the anatomy of this region in proterosuchids. The cranial endocast preserves possibly part of the vomero-nasal ( = Jacobson’s) system laterally to the olfactory bulbs. Previous claims of the absence of the vomero-nasal organs in archosaurs, which is suggested by the extant phylogenetic bracket, are questioned because its absence in both clades of extant archosaurs seems to be directly related with the independent acquisition of a non-ground living mode of life.
Archosauria and their closest relatives, the non-archosaurian archosauriforms, diversified in the Early and Middle Triassic, soon after the end-Permian extinction. This diversification is poorly documented in most Lower and Middle Triassic rock sequences because fossils of early groups of archosauriforms are relatively rare compared to those of other amniotes. The early Middle Triassic (? late Anisian) Manda beds of southwestern Tanzania form an exception, with early archosaur skeletons being relatively common and preserved as articulated or associated specimens. The Manda archosaur assemblage is exceptionally diverse for the Middle Triassic. However, to date, no non-archosaurian archosauriforms have been reported from these rocks.