Da-Yong Jiang

A basal ichthyosauriform with a short snout from the Lower Triassic of China

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.

First record of Placodontoidea (Reptilia, Sauropterygia, Placodontia) from the Eastern Tethys

DA-YONG JIANG, RYOSUKE MOTANI, WEI-CHENG HAO, OLIVIER RIEPPEL, YUAN-LIN SUN, LARS SCHMITZ, and ZUO-YU SUN Department of Geology and Geological Museum, Peking University, Beijing 100871, P. R. China, djiang@pku.edu.cn; Department of Geology, University of California, Davis, California 95616-8605, U.S.A; Department of Geology, The Field Museum, Chicago, Illinois 60605-2496, U.S.A., rieppel@fieldmuseum.org.

Terrestrial Origin of Viviparity in Mesozoic Marine Reptiles Indicated by Early Triassic Embryonic Fossils

Viviparity in Mesozoic marine reptiles has traditionally been considered an aquatic adaptation. We report a new fossil specimen that strongly contradicts this traditional interpretation. The new specimen contains the oldest fossil embryos of Mesozoic marine reptile that are about 10 million years older than previous such records. The fossil belongs to Chaohusaurus (Reptilia, Ichthyopterygia), which is the oldest of Mesozoic marine reptiles (ca. 248 million years ago, Early Triassic). This exceptional specimen captures an articulated embryo in birth position, with its skull just emerged from the maternal pelvis. Its headfirst birth posture, which is unlikely to be a breech condition, strongly indicates a terrestrial origin of viviparity, in contrast to the traditional view. The tail-first birth posture in derived ichthyopterygians, convergent with the conditions in whales and sea cows, therefore is a secondary feature. The unequivocally marine origin of viviparity is so far not known among amniotes, a subset of vertebrate animals comprising mammals and reptiles, including birds. Therefore, obligate marine amniotes appear to have evolved almost exclusively from viviparous land ancestors. Viviparous land reptiles most likely appeared much earlier than currently thought, at least as early as the recovery phase from the end-Permian mass extinction.

A NEW MIXOSAURID ICHTHYOSAUR FROM THE MIDDLE TRIASSIC OF CHINA

Abstract We describe the new ichthyosaur taxon Mixosaurus panxianensis, sp. nov., from the Middle Triassic of Guizhou Province, China. Diagnostic characters of the new species include a short posteroventral jugal process and the absence of external contact between jugal and quadratojugal. The morphologic description of the type specimens amends the knowledge of the postorbital region and the postcranium of the Mixosauridae. The holotype of Mixosaurus maotaiensis, which is very fragmentary and has no taxonomic value at the species level, is found to be undiagnostic, and hence the lately introduced, monospecific genus name for this species needs to be abandoned. Phylogenetic analysis strongly supports the monophyly of the family Mixosauridae. Furthermore, the analysis shows a bifurcation of the family into two sister groups, suggesting the presence of two genera, Mixosaurus and Phalarodon. Mixosaurus, characterized by a relatively short and wide humerus, includes M. panxianensis sp. nov., M. cornalianus, and M. kuhnschnyderi. Synapomorphies of Phalarodon, which contains P. fraasi, P. callawayi, and P. atavus, are a narial shelf and the absence of a maxillary dental groove.

A new pachypleurosaur (Reptilia: Sauropterygia) from the lower Middle Triassic of southwestern China and the phylogenetic relationships of Chinese pachypleurosaurs

ABSTRACT A new genus and species of Pachypleurosauria, Dianopachysaurus dingi, gen. et sp. nov., from a recently discovered Middle Triassic Lagerstätte in the Anisian Guanling Formation of Yunnan Province is described. The monophyly of Pachypleurosauria, including all potential Chinese pachypleurosaurs in this study, is supported by the rigorous cladistic analysis conducted here. Phylogenetic relationships of Chinese pachypleurosaurs are clarified. Wumengosaurus occupies the most basal position within Pachypleurosauria. Keichousaurus and Dianopachysaurus form the monophyletic Keichousauridae that is the sister group of all European pachypleurosaurs. Our cladistic analysis also supports a previously proposed paleobiogeographic hypothesis of an eastern Tethyan origin of pachypleurosaurs and their migration into the western Tethyan faunal province. The existence of a long ghost lineage as required by the paleobiogeographic hypothesis is greatly shortened by the discovery of Dianopachysaurus.

A new species of Xinpusaurus (Thalattosauria) from the Upper Triassic of China

Abstract A new species of Xinpusaurus Yin in Yin et al., 2000 (Thalattosauria, Thalattosauridae), is described from the Wayao Member of the Falang Formation (Tuvalian, Carnian, Upper Triassic) of Guanling County in Guizhou, southwestern China. It is the third thalattosaur species known from the Guanling faunal assemblage of marine reptiles. The type and only specimen consists of an entire skeleton, including a complete skull. It differs from the type species, X. suni, by its smaller adult size, the larger nasal, the more slender angular and low retroarticular process, the shape of the cervical neural spines, the number of presacral vertebrae, the large size of the scapula, the shape of the radius, the presence of a well ossified carpus, the more slender femur, the smaller hindfin, the presence of a complete row of distal tarsal ossifications, and the proportions of the metatarsals. It is consequently referred to a new species, Xinpusaurus kohi. A new analysis of thalattosaur interrelationships based on 30 cranial and postcranial characters corroborates a sister-group relationship between Xinpusaurus and Nectosaurus from the Carnian of California.

A New Middle Triassic Eosauropterygian (Reptilia, Sauropterygia) from Southwestern China

Abstract A new eosauropterygian genus and species is described from the middle Anisian (Middle Triassic) of Panxian (Guizhou Province, southwestern China). The new taxon is characterized by an unique specialization of the jaws that form an elongate pointed rostrum furnished with numerous small, monocuspid and vertically oriented teeth with a basally expanded crown and pointed tip, which is quite unlike any other sauropterygian known. Other diagnostic characters include: frontals paired; parietal unpaired; pineal foramen located centrally in broad parietal skull table; two carpal ossification; ilium with distinct preacetabular process at the base of the dorsally extending iliac blade; pubis plate-like and of rounded contours; two tarsal ossifications. The curved and distally expanded humerus, the reversed topological relationship of the clavicle and scapula, and the presence of three sacral ribs indicate sauropterygian affinities of the new taxon, whereas the ‘butterfly-shaped’ or ‘cruciform’ facet for the neural arch on the dorsal centrum surface indicate its eosauropterygian status.

A gigantic nothosaur (Reptilia: Sauropterygia) from the Middle Triassic of SW China and its implication for the Triassic biotic recovery

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.

The enigmatic marine reptile nanchangosaurus from the lower triassic of Hubei, China and the phylogenetic affinities of Hupehsuchia.

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.

Absence of Suction Feeding Ichthyosaurs and Its Implications for Triassic Mesopelagic Paleoecology

Mesozoic marine reptiles and modern marine mammals are often considered ecological analogs, but the extent of their similarity is largely unknown. Particularly important is the presence/absence of deep-diving suction feeders among Mesozoic marine reptiles because this would indicate the establishment of mesopelagic cephalopod and fish communities in the Mesozoic. A recent study suggested that diverse suction feeders, resembling the extant beaked whales, evolved among ichthyosaurs in the Triassic. However, this hypothesis has not been tested quantitatively. We examined four osteological features of jawed vertebrates that are closely linked to the mechanism of suction feeding, namely hyoid corpus ossification/calcification, hyobranchial apparatus robustness, mandibular bluntness, and mandibular pressure concentration index. Measurements were taken from 18 species of Triassic and Early Jurassic ichthyosaurs, including the presumed suction feeders. Statistical comparisons with extant sharks and marine mammals of known diets suggest that ichthyosaurian hyobranchial bones are significantly more slender than in suction-feeding sharks or cetaceans but similar to those of ram-feeding sharks. Most importantly, an ossified hyoid corpus to which hyoid retractor muscles attach is unknown in all but one ichthyosaur, whereas a strong integration of the ossified corpus and cornua of the hyobranchial apparatus has been identified in the literature as an important feature of suction feeders. Also, ichthyosaurian mandibles do not narrow rapidly to allow high suction pressure concentration within the oral cavity, unlike in beaked whales or sperm whales. In conclusion, it is most likely that Triassic and Early Jurassic ichthyosaurs were ‘ram-feeders’, without any beaked-whale-like suction feeder among them. When combined with the inferred inability for dim-light vision in relevant Triassic ichthyosaurs, the fossil record of ichthyosaurs does not suggest the establishment of modern-style mesopelagic animal communities in the Triassic. This new interpretation matches the fossil record of coleoids, which indicates the absence of soft-bodied deepwater species in the Triassic.