Leptosuchus Case, 1922 (Reptilia: Phytosauria) from the Late Triassic of the American West is represented by many specimens. Here, I present complete morphological descriptions of the skull material of a new taxon from the Sonsela Member (Chinle Formation) of Petrified Forest National Park, Arizona, with the first rigorous phylogenetic analysis focused on the interrelationships of Leptosuchus. The new taxon is recovered as the sister taxon to Pseudopalatinae. It possesses one unambiguous synapomorphy (the 'septomaxillae' form part of the lateral borders of the nares) and shares the presence of a subsidiary opisthotic process with Pseudopalatinae. The new taxon does not fall within the restricted clade Leptosuchus. In my analysis, the previously proposed, but undemonstrated, sister taxon relationship between Angistorhinus and Rutiodon is not supported, Paleorhinus is recovered as paraphyletic, and a subset of taxa traditionally included within Leptosuchus are found to be more closely related to Pseudopalatinae, rendering Leptosuchus paraphyletic. 'Leptosuchus' adamanensis emerges as sister taxon to Smilosuchus gregorii and is here referred to as Smilosuchus adamanensis nov. comb., and 'Machaeroprosopus' lithodendrorum is also transferred to Smilosuchus lithodendrorum nov. comb. Documentation of the variation present within Phytosauria, and specifically within Leptosuchus sensu lato, demonstrates higher diversity within Phytosauria than previously appreciated and places the character states previously proposed for Pseudopalatinae into a broader context of shared characters.
Vancleavea campi Long & Murry, 1995, from the Late Triassic of western North America, represents the latest surviving non-archosaurian archosauriform known to date. We present here a detailed comparative description based on a nearly complete, articulated skeleton from the Coelophysis Quarry in north-central New Mexico and other fragmentary specimens. The unique combination of morphological features of Vancleavea is unparalleled within Reptilia; it has four unique morphologies of imbricated osteoderms covering the entire body, a short, highly ossified skull, relatively small limbs and morphological features consistent with a semi-aquatic lifestyle. Vancleavea is placed in a rigorous phylogenetic analysis examining the relationships of non-archosaurian archosauriforms, and is found to be more closely related to Archosauria than both Erythrosuchus and Proterosuchus, but outside of the crown group. The analysis confirms previously hypothesized relationships, which found Euparkeria to be the closest sister taxon of Archosauria. It is not clear whether specimens referred to Vancleavea campi represent a single species-level taxon or a clade of closely related taxa that lived through much of the Late Triassic of North America, given the poor fossil record of the taxon.
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 Post Quarry, within the lower part of the type section of the Upper Triassic Cooper Canyon Formation in southern Garza County, western Texas, contains a remarkably diverse vertebrate assemblage. The Post Quarry has produced: the small temnospondylRileymillerus cosgriffi; the metoposauridApachesaurus gregorii; possible dicynodonts and eucynodonts; a clevosaurid sphenodontian; non-archosauriform archosauromorphs (Trilophosaurus dornorum, simiosaurians, and possiblyMalerisaurus); the phytosaurLeptosuchus; several aetosaurs (Calyptosuchus wellesi,Typothorax coccinarum,Paratypothorax, andDesmatosuchus smalli); the poposauroidShuvosaurus inexpectatus(“Chatterjeea elegans”); the rauisuchidPostosuchus kirkpatricki; an early crocodylomorph; several dinosauromorphs (the lagerpetidDromomeron gregorii, the silesauridTechnosaurus smalli, a herrerasaurid, and an early neotheropod); and several enigmatic small diapsids. Revised lithostratigraphic correlations of the lower Cooper Canyon Formation with the Tecovas Formation, the occurrence ofLeptosuchus, and the overall composition of the assemblage indicate that the Post Quarry falls within the Adamanian biozone, and not the Revueltian biozone. Stratigraphic subdivision of the Adamanian biozone may be possible, and the Post Quarry may be correlative with the upper part of the Adamanian biozone in Arizona. The age of the Post Quarry assemblage is possibly late Lacian or earliest Alaunian (late early Norian or earliest middle Norian), between 220 and 215 Ma.
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.
Phytosaurs are a diverse and morphologically distinctive clade of superficially crocodile-like archosauriforms that had a near global distribution during the Late Triassic. Because their remains are among the most abundant vertebrate remains recovered in many Upper Triassic terrestrial formations, phytosaurs are used extensively in long-range biochronological and biostratigraphic correlations. The biochronologically oldest and earliest branching known phytosaurs include an array of nominal species from the early Late Triassic of the United States, Germany, Poland, Morocco, and India that have been synonymized within the genus Paleorhinus, and subsequently used to define a global 'Paleorhinus biochron'. However, recent phylogenetic work suggested that the North American species previously referred to Paleorhinus are paraphyletic. Here, we reassess the systematics and anatomy of putative specimens of Paleorhinus from southern Germany. Two well-preserved basal phytosaur skulls from the Blasensandstein (Carnian) of Bavaria form the holotypes of Francosuchus angustifrons and Ebrachosuchus neukami, both of which were synonymized with Paleorhinus by previous workers. We demonstrate that Francosuchus angustifrons shares unique synapomorphies with specimens referred to Paleorhinus bransoni from the Late Triassic of Texas, and thus refer the species to Paleorhinus. By contrast, the longirostrine Ebrachosuchus is highly distinctive in morphology, and our new cladistic analysis of Phytosauria demonstrates that it represents a valid taxon that is more closely related to Phytosauridae than to Paleorhinus. We provide the first autapomorphybased support for a monophyletic but restricted Paleorhinus (supported by a nodal row on the jugal, and low paired ridges on the squamosal) and confirm that previous broader conceptions of Paleorhinus are likely to be paraphyletic.
The neotype skull of the Indian phytosaur Parasuchus hislopi Lydekker, 1885 (ISI R42) is re‐evaluated and compared with the type material of other basal phytosaurs. Parasuchus hislopi is extremely similar to species previously placed in Paleorhinus (P. bransoni and P. angustifrons), sharing with them such characters as a series of nodes on the lateral surface of the jugal, paired ridges on the squamosal and a frontal depression. Parasuchus hislopi represents a valid species: it can be distinguished from P. bransoni by a relatively low narial eminence and P. angustifrons by the absence of paired nasal depressions. Inclusion of Parasuchus hislopi in a phylogenetic analysis of phytosaurs recovers it in a well‐supported clade with P. bransoni and P. angustifrons. Parasuchus is considered the senior synonym of Paleorhinus and Arganarhinus. Parasuchus (here considered to include P. hislopi, P. angustifrons, P. bransoni and P. magnoculus) has a broad circum‐Pangaean distribution, with species occurring in the south‐western United States, Morocco, central Europe and India. Phytosaur higher‐level taxonomy is also revised: Parasuchidae is redefined to include ‘Paleorhinus‐grade’ phytosaurs and the later‐diverging Mystriosuchinae (the group formerly known as Phytosauridae), and Pseudopalatinae is renamed Mystriosuchini for reason of priority.
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