Project description:Following the Permo-Triassic mass extinction, Archosauriformes-the clade that includes crocodylians, birds, and their extinct relatives outside crown Archosauria-rapidly diversified into many distinct lineages, became distributed globally, and, by the Late Triassic, filled a wide array of resource zones. Current scenarios of archosauriform evolution are ambiguous with respect to whether their taxonomic diversification in the Early-Middle Triassic coincided with the initial evolution of dietary specializations that were present by the Late Triassic or if their ecological disparity arose sometime after lineage diversification. Late Triassic archosauriform dietary specialization is recorded by morphological divergence from the plesiomorphic archosauriform tooth condition (laterally-compressed crowns with serrated carinae and a generally triangular lateral profile). Unfortunately, the roots of this diversification are poorly documented, with few known Early--Middle Triassic tooth assemblages, limiting characterizations of morphological diversity during this critical, early period in archosaur evolution. Recent fieldwork (2007-2017) in the Middle Triassic Manda Beds of the Ruhuhu Basin, Tanzania, recovered a tooth assemblage that provides a window into this poorly sampled interval. To investigate the taxonomic composition of that collection, we built a dataset of continuous quantitative and discrete morphological characters based on in situ teeth of known taxonomic status (e.g., Nundasuchus, Parringtonia: N = 65) and a sample of isolated teeth (N = 31). Using crown heights from known taxa to predict tooth base ratio (= base length/width), we created a quantitative morphospace for the tooth assemblage. The majority of isolated, unassigned teeth fall within a region of morphospace shared by several taxa from the Manda Beds (e.g., Nundasuchus, Parringtonia); two isolated teeth fall exclusively within a "Pallisteria" morphospace. A non-metric multidimensional scaling ordination (N = 67) of 11 binary characters reduced overlap between species. The majority of the isolated teeth from the Manda assemblage fall within the Nundasuchus morphospace. This indicates these teeth are plesiomorphic for archosauriforms as Nundasuchus exhibits the predicted plesiomorphic condition of archosauriform teeth. Our model shows that the conservative tooth morphologies of archosauriforms can be differentiated and assigned to species and/or genus, rendering the model useful for identifying isolated teeth. The large overlap in tooth shape among the species present and their overall similarity indicates that dietary specialization lagged behind species diversification in archosauriforms from the Manda Beds, a pattern predicted by Simpson's "adaptive zones" model. Although applied to a single geographic region, our methods offer a promising means to reconstruct ecological radiations and are readily transferable across a broad range of vertebrate taxa throughout Earth history.
Project description:Heteropelta boboi is a new archosauriform reptile from the upper Anisian of northeastern Italy represented by a fragment of dorsal armor with a row of neural arches of the dorsal vertebrae. The dorsal armor of the new taxon is composed of two columns of paramedian osteoderms and at least six columns of lateral osteoderms. Unlike other armored archosaurs, the osteoderms are imbricated with the posterior osteoderm overlapping the anterior one. The low neural arches bear small neural spines and long postzygapophyses. The osteoderms of the lateral columns increase in size and change in shape from the most medial to the most lateral columns. Among the Archosauriformes, only the non-archosaur proterochampsians Vancleavea campi, Litorosuchus somnii, and the doswelliids have dorsal armor comprised of more than two columns of osteoderms per side, but the morphology and arrangement of their osteoderms is unlike those of the new Italian taxon. A cladistic analysis of Archosauromorpha positions Heteropelta boboi as either a basal phytosaur or a basal suchian. However, a second cladistic analysis focused on armored archosaurs alternatively positions the new taxon as a basal archosauriform, basal suchian, basal loricatan or crocodylomorph. Better resolution of the phylogenetic relationships of Heteropelta boboi will likely be obtained only with the discovery of cranial and postcranial remains associated with its diagnostic armor elements.
Project description:Neck elongation has appeared independently in several tetrapod groups, including giraffes and sauropod dinosaurs on land, birds and pterosaurs in the air, and sauropterygians (plesiosaurs and relatives) in the oceans. Long necks arose in Early Triassic sauropterygians, but the nature and rate of that elongation has not been documented. Here, we report a new species of pachypleurosaurid sauropterygian, Chusaurus xiangensis gen. et sp. nov., based on two new specimens from the Early Triassic Nanzhang-Yuan'an Fauna in the South China Block. The new species shows key features of its Middle Triassic relatives, but has a relatively short neck, measuring 0.48 of the trunk length, compared to > 0.8 from the Middle Triassic onwards. Comparative phylogenetic analysis shows that neck elongation occurred rapidly in all Triassic eosauropterygian lineages, probably driven by feeding pressure in a time of rapid re-establishment of new kinds of marine ecosystems.
Project description:The Manda Beds of southwest Tanzania have yielded key insights into the early evolutionary radiation of archosaurian reptiles. Many key archosaur specimens were collected from the Manda Beds in the 1930s and 1960s, but until recently, few of these had been formally published. Here, we describe an archosaur specimen collected in 1963 which has previously been referred to informally as Pallisteria angustimentum. We recognize this specimen as the type of a new taxon, Mambawakale ruhuhu gen. et sp. nov. The holotype and only known specimen of M. ruhuhu comprises a partial skull of large size (greater than 75 cm inferred length), lower jaws and fragments of the postcranium, including three anterior cervical vertebrae and a nearly complete left manus. Mambawakale ruhuhu is characterized by several cranial autapomorphies that allow it to be distinguished with confidence from all other Manda Beds archosaurs, with the possible exception of Stagonosuchus nyassicus for which comparisons are highly constrained due to very limited overlapping material. Phylogenetic analysis suggests that M. ruhuhu is an early diverging pseudosuchian, but more precise resolution is hampered by missing data. Mambawakale ruhuhu is one of the largest known pseudosuchians recovered to date from the Middle Triassic.
Project description:The Placerias/Downs' Quarry complex in eastern Arizona, USA, is the most diverse Upper Triassic vertebrate locality known. We report a new short-faced archosauriform, Syntomiprosopus sucherorum gen. et sp. nov., represented by four incomplete mandibles, that expands that diversity with a morphology unique among Late Triassic archosauriforms. The most distinctive feature of Syntomiprosopus gen. nov. is its anteroposteriorly short, robust mandible with 3-4 anterior, a larger caniniform, and 1-3 "postcanine" alveoli. The size and shape of the alveoli and the preserved tips of replacement teeth preclude assignment to any taxon known only from teeth. Additional autapomorphies of S. sucherorum gen. et sp. nov. include a large fossa associated with the mandibular fenestra, an interdigitating suture of the surangular with the dentary, fine texture ornamenting the medial surface of the splenial, and a surangular ridge that completes a 90° arc. The external surfaces of the mandibles bear shallow, densely packed, irregular, fine pits and narrow, arcuate grooves. This combination of character states allows an archosauriform assignment; however, an associated and similarly sized braincase indicates that Syntomiprosopus n. gen. may represent previously unsampled disparity in early-diverging crocodylomorphs. The Placerias Quarry is Adamanian (Norian, maximum depositional age ~219 Ma), and this specimen appears to be an early example of shortening of the skull, which occurs later in diverse archosaur lineages, including the Late Cretaceous crocodyliform Simosuchus. This is another case where Triassic archosauriforms occupied morphospace converged upon by other archosaurs later in the Mesozoic and further demonstrates that even well-sampled localities can yield new taxa.
Project description:The Middle Triassic Luoping Biota in south-west China represents the inception of modern marine ecosystems, with abundant and diverse arthropods, fishes and marine reptiles, indicating recovery from the Permian-Triassic mass extinction. Here we report a new specimen of the predatory marine reptile Diandongosaurus, based on a nearly complete skeleton. The specimen is larger than most other known pachypleurosaurs, and the body shape, caniniform teeth, clavicle with anterior process, and flat distal end of the anterior caudal ribs show its affinities with Diandongosaurus acutidentatus, while the new specimen is approximately three times larger than the holotype. The morphological characters indicate that the new specimen is an adult of D. acutidentatus, allowing for ontogenetic variation. The fang-like teeth and large body size confirm it was a predator, but the amputated hind limb on the right side indicate itself had been predated by an unknown hunter. Predation on such a large predator reveals that predation pressure in the early Mesozoic was intensive, a possible early hint of the Mesozoic Marine Revolution.
Project description:The presence of gigantic apex predators in the eastern Panthalassic and western Tethyan oceans suggests that complex ecosystems in the sea had become re-established in these regions at least by the early Middle Triassic, after the Permian-Triassic mass extinction (PTME). However, it is not clear whether oceanic ecosystem recovery from the PTME was globally synchronous because of the apparent lack of such predators in the eastern Tethyan/western Panthalassic region prior to the Late Triassic. Here we report a gigantic nothosaur from the lower Middle Triassic of Luoping in southwest China (eastern Tethyan ocean), which possesses the largest known lower jaw among Triassic sauropterygians. Phylogenetic analysis suggests parallel evolution of gigantism in Triassic sauropterygians. Discovery of this gigantic apex predator, together with associated diverse marine reptiles and the complex food web, indicates global recovery of shallow marine ecosystems from PTME by the early Middle Triassic.
Project description: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.
Project description:Erythrosuchidae is a clade of early archosauriform reptiles, which were apex predators in many late Early and Middle Triassic ecosystems, following the Permo-Triassic mass extinction. Erythrosuchids had a worldwide distribution, with well-preserved fossil material known from South Africa, European Russia, and China. We here redescribe the anatomy and revise the taxonomy of Guchengosuchus shiguaiensis, which is one of the stratigraphically oldest erythrosuchids and is known from a single partial skeleton from the lowermost Middle Triassic (lower Anisian) lower Ermaying Formation of Shaanxi Province, China. We provide a new differential diagnosis for Guchengosuchus shiguaiensis, and identify a series of autapomorphies relating to the morphologies of the skull roof and vertebrae. Incorporating updated anatomical information for Guchengosuchus into the most comprehensive morphological phylogenetic analysis available for early archosauromorphs recovers it as an early branching member of Erythrosuchidae, outside of the clade formed by Garjainia, Erythrosuchus, Chalishevia, and Shansisuchus. Fugusuchus hejiapanensis, from the uppermost Lower Triassic to lower Middle Triassic Heshanggou Formation of China, is recovered as the earliest branching member of Erythrosuchidae.
Project description:A new species of the erythrosuchid archosauriform reptile Garjainia Ochev, 1958 is described on the basis of disarticulated but abundant and well-preserved cranial and postcranial material from the late Early Triassic (late Olenekian) Subzone A of the Cynognathus Assemblage Zone of the Burgersdorp Formation (Beaufort Group) of the Karoo Basin of South Africa. The new species, G. madiba, differs from its unique congener, G. prima from the late Olenekian of European Russia, most notably in having large bony bosses on the lateral surfaces of the jugals and postorbitals. The new species also has more teeth and a proportionately longer postacetabular process of the ilium than G. prima. Analysis of G. madiba bone histology reveals thick compact cortices comprised of highly vascularized, rapidly forming fibro-lamellar bone tissue, similar to Erythrosuchus africanus from Subzone B of the Cynognathus Assemblage Zone. The most notable differences between the two taxa are the predominance of a radiating vascular network and presence of annuli in the limb bones of G. madiba. These features indicate rapid growth rates, consistent with data for many other Triassic archosauriforms, but also a high degree of developmental plasticity as growth remained flexible. The diagnoses of Garjainia and of Erythrosuchidae are addressed and revised. Garjainia madiba is the geologically oldest erythrosuchid known from the Southern Hemisphere, and demonstrates that erythrosuchids achieved a cosmopolitan biogeographical distribution by the end of the Early Triassic, within five million years of the end-Permian mass extinction event. It provides new insights into the diversity of the Subzone A vertebrate assemblage, which partially fills a major gap between classic 'faunal' assemblages from the older Lystrosaurus Assemblage Zone (earliest Triassic) and the younger Subzone B of the Cynognathus Assemblage Zone (early Middle Triassic).