The incompleteness of the fossil record obscures the origin of many of the more derived clades of vertebrates. One such group is the Ichthyopterygia, a clade of obligatory marine reptiles that appeared in the Early Triassic epoch, without any known intermediates. Here we describe a basal ichthyosauriform from the upper Lower Triassic (about 248 million years ago) of China, whose primitive skeleton indicates possible amphibious habits. It is smaller than ichthyopterygians and had unusually large flippers that probably allowed limited terrestrial locomotion. It also retained characteristics of terrestrial diapsid reptiles, including a short snout and body trunk. Unlike more-derived ichthyosauriforms, it was probably a suction feeder. The new species supports the sister-group relationships between ichthyosauriforms and Hupehsuchia, the two forming the Ichthyosauromorpha. Basal ichthyosauromorphs are known exclusively from south China, suggesting that the clade originated in the region, which formed a warm and humid tropical archipelago in the Early Triassic. The oldest unequivocal record of a sauropterygian is also from the same stratigraphic unit of the region.
Largocephalosaurus polycarpon Cheng et al. 2012a was erected after the study of the skull and some parts of a skeleton and considered to be an eosauropterygian. Here we describe a new species of the genus, Largocephalosaurus qianensis, based on three specimens. The new species provides many anatomical details which were described only briefly or not at all in the type species, and clearly indicates that Largocephalosaurus is a saurosphargid. It differs from the type species mainly in having three premaxillary teeth, a very short retroarticular process, a large pineal foramen, two sacral vertebrae, and elongated small granular osteoderms mixed with some large ones along the lateral most side of the body. With additional information from the new species, we revise the diagnosis and the phylogenetic relationships of Largocephalosaurus and clarify a set of diagnostic features for the Saurosphargidae Li et al. 2011. Largocephalosaurus is characterized primarily by an oval supratemporal fenestra, an elongate dorsal ‘rib-basket’, a narrow and elongate transverse process of the dorsal vertebrae, and the lack of a complete dorsal carapace of osteoderms. The Saurosphargidae is distinct mainly in having a retracted external naris, a jugal–squamosal contact, a large supratemporal extensively contacting the quadrate shaft, a leaf-like tooth crown with convex labial surface and concave lingual surface, a closed dorsal ‘rib-basket’, many dorsal osteoderms, a large boomerang-like or atypical T-shaped interclavicle. Current evidence suggests that the Saurosphargidae is the sister-group of the Sauropterygia and that Largocephalosaurus is the sister-group of the Saurosphargis–Sinosaurosphargis clade within the family.
The study of the holotype and of a new specimen of Nanchangosaurus suni (Reptilia; Diapsida; Hupehsuchia) revealed a suite of hitherto unrecognized characters. For example, Nanchangosaurus has bipartite neural spines and its vertebral count is nearly identical to that of Hupehsuchus. It differs from the latter in having poorly developed forelimbs despite the advanced ossification in the rest of the skeleton. Other differences all pertain to hupehsuchian plesiomorphies retained in Nanchangosaurus, such as low neural spines. The relationship of Hupehsuchia within Diapsida was analyzed based on a data matrix containing 41 taxa coded for 213 characters, of which 18 were identified as aquatic adaptations from functional inferences. These aquatic adaptations may be vulnerable to the argumentation of character homology because expectation for homoplasy is high. There is an apparent incongruence between phylogenetic signals from aquatic adaptations and the rest of the data, with aquatic adaptations favoring all marine reptiles but Helveticosaurus to form a super-clade. However, this super-clade does not obtain when aquatic adaptations were deleted, whereas individual marine reptile clades are all derived without them. We examined all possible combinations of the 18 aquatic adaptations (n = 262143) and found that four lineages of marine reptiles are recognized almost regardless of which of these features were included in the analysis: Hupehsuchia-Ichthyopterygia clade, Sauropterygia-Saurosphargidae clade, Thalattosauria, and Helveticosaurus. The interrelationships among these four depended on the combination of aquatic adaptations to be included, i.e., assumed to be homologous a priori by bypassing character argumentation. Hupehsuchia always appeared as the sister taxon of Ichthyopterygia.
In the last 15 years, the discovery of several new actinopterygian fish faunas from the Early and Middle Triassic of the Tethys, cast new light on the timing, speed and range of their recovery after the end-Permian crisis. In addition to several new taxa having been described, the stratigraphical and geographical record of many others have been greatly extended. In fact, most of the new fossiliferous sites are in southern China, thus at the Eastern end of the Tethys, and furthermore a few are somewhat older (Chaohu, Panxian, Luoping) than the major classical Western Tethys sites (Monte San Giorgio). Following these new finds, it is possible to have a better definition of the Triassic recovery stages. Indeed, after a quite short phase till the end of the Smithian (Olenekian, Early Triassic) in which a rather consistent fauna was present all around the Pangea coasts, a major radiation occurred in the Early-Middle Anisian after the new Middle Triassic fish fauna already appeared in the late Early Triassic, thus occuring well before what was previously supposed from the Alps localities. Furthermore, the new assemblages from southern China point to an early broader differentiation among the basal neopterygians rather than in the 'subholosteans', the group that was then dominant in the Western Tethys since the Late Anisian. It stands that during the Norian a new basal neopterygian radiation gave rise to several new branches that dominated the remaining part of the Mesozoic.
Parahupehsuchus longus is a new species of marine reptile from the Lower Triassic of Yuan’an County, Hubei Province, China. It is unique among vertebrates for having a body wall that is completely surrounded by a bony tube, about 50 cm long and 6.5 cm deep, comprising overlapping ribs and gastralia. This tube and bony ossicles on the back are best interpreted as anti-predatory features, suggesting that there was predation pressure upon marine tetrapods in the Early Triassic. There is at least one sauropterygian that is sufficiently large to feed on Parahupehsuchus in the Nanzhang-Yuan’an fauna, together with six more species of potential prey marine reptiles with various degrees of body protection. Modern predators of marine tetrapods belong to the highest trophic levels in the marine ecosystem but such predators did not always exist through geologic time. The indication of marine-tetrapod feeding in the Nanzhang-Yuan’an fauna suggests that such a trophic level emerged for the first time in the Early Triassic. The recovery from the end-Permian extinction probably proceeded faster than traditionally thought for marine predators. Parahupehsuchus has superficially turtle-like features, namely expanded ribs without intercostal space, very short transverse processes, and a dorsal outgrowth from the neural spine. However, these features are structurally different from their turtle counterparts. Phylogeny suggests that they are convergent with the condition in turtles, which has a fundamentally different body plan that involves the folding of the body wall. Expanded ribs without intercostal space evolved at least twice and probably even more among reptiles.
Contrary to the fast radiation of most metazoans after the end-Permian mass extinction, it is believed that early marine reptiles evolved slowly during the same time interval. However, emerging discoveries of Early Triassic marine reptiles are questioning this traditional view. Here we present an aberrant basal ichthyosauriform with a hitherto unknown body design that suggests a fast radiation of early marine reptiles. The new species is larger than coeval marine reptiles and has an extremely small head and a long tail without a fluke. Its heavily-built body bears flattened and overlapping gastral elements reminiscent of hupehsuchians. A phylogenetic analysis places the new species at the base of ichthyosauriforms, as the sister taxon of Cartorhynchus with which it shares a short snout with rostrally extended nasals. It now appears that ichthyosauriforms evolved rapidly within the first one million years of their evolution, in the Spathian (Early Triassic), and their true diversity has yet to be fully uncovered. Early ichthyosauromorphs quickly became extinct near the Early-Middle Triassic boundary, during the last large environmental perturbation after the end-Permian extinction involving redox fluctuations, sea level changes and volcanism. Marine reptile faunas shifted from ichthyosauromorph-dominated to sauropterygian-dominated composition after the perturbation.
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