Evidence Supporting Predation of 4-m Marine Reptile by Triassic Megapredator.

Air-breathing marine predators have been essential components of the marine ecosystem since the Triassic. Many of them are considered the apex predators but without direct evidence-dietary inferences are usually based on circumstantial evidence, such as tooth shape. Here we report a fossil that likely represents the oldest evidence for predation on megafauna, i.e., animals equal to or larger than humans, by marine tetrapods-a thalattosaur (∼4 m in total length) in the stomach of a Middle Triassic ichthyosaur (∼5 m). The predator has grasping teeth yet swallowed the body trunk of the prey in one to several pieces. There were many more Mesozoic marine reptiles with similar grasping teeth, so megafaunal predation was likely more widespread than presently conceived. Megafaunal predation probably started nearly simultaneously in multiple lineages of marine reptiles in the Illyrian (about 242-243 million years ago).

Repeated evolution of durophagy during ichthyosaur radiation after mass extinction indicated by hidden dentition.

Marine tetrapods quickly diversified and were established as marine top predators after the end-Permian Mass extinction (EPME). Ichthyosaurs were the forerunner of this rapid radiation but the main drivers of the diversification are poorly understood. Cartorhynchus lenticarpus is a basal ichthyosauriform with the least degree of aquatic adaptation, holding a key to identifying such a driver. The unique specimen appeared edentulous based on what was exposed but a CT scanning revealed that the species indeed had rounded teeth that are nearly perpendicular to the jaw rami, and thus completely concealed in lateral view. There are three dental rows per jaw ramus, and the root lacks infoldings of the dentine typical of ichthyopterygians. The well-developed and worn molariform dentition with three tooth rows supports the previous inference that the specimen is not of a juvenile. The premaxilla and the corresponding part of the dentary are edentulous. Molariform dentition evolved three to five times independently within Ichthyosauriformes in the Early and Middle Triassic. Convergent exploitation of hard-shelled invertebrates by different subclades of ichthyosauriforms likely fueled the rapid taxonomic diversification of the group after EPME.

The new ichthyosauriform Chaohusaurus brevifemoralis (Reptilia, Ichthyosauromorpha) from Majiashan, Chaohu, Anhui Province, China.

A new species of ichthyosauriform is recognized based on 20 specimens, including nearly complete skeletons, and named Chaohusaurus brevifemoralis. A part of the specimens was previously identified as Chaohusaurus chaoxianensis and is herein reassigned to the new species. The new species differs from existing species of Chaohusaurus in a suite of features, such as the bifurcation of the caudal peak neural spine and a short femur relative to trunk length. The specimens include both complete and partially disarticulated skulls, allowing rigorous scrutiny of cranial sutures. For example, the squamosal does not participate in the margin of the upper temporal fenestra despite previous interpretations. Also, the frontal unequivocally forms a part of the anterior margin of the upper temporal fenestra, forming the most medial part of the anterior terrace. The skull of the holotype largely retains three-dimensionality with the scleral rings approximately in situ, revealing that the eyeball was uncovered in two different directions, that is, laterally and slightly dorsally through the main part of the orbit, and dorsally through the medial extension of the orbit into the skull roof. This skull construction is likely a basal feature of Ichthyosauromorpha. Phylogenetic analyses place the new species as a sister taxon of Chaohusaurus chaoxianensis.

Early Triassic marine reptile representing the oldest record of unusually small eyes in reptiles indicating non-visual prey detection.

The end-Permian mass extinction (EPME) led to reorganization of marine predatory communities, through introduction of air-breathing top predators, such as marine reptiles. We report two new specimens of one such marine reptile, Eretmorhipis carrolldongi, from the Lower Triassic of Hubei, China, revealing superficial convergence with the modern duckbilled platypus (Ornithorhynchus anatinus), a monotreme mammal. Apparent similarities include exceptionally small eyes relative to the body, snout ending with crura with a large internasal space, housing a bone reminiscent of os paradoxum, a mysterious bone of platypus, and external grooves along the crura. The specimens also have a rigid body with triangular bony blades protruding from the back. The small eyes likely played reduced roles during foraging in this animal, as with extant amniotes (group containing mammals and reptiles) with similarly small eyes. Mechanoreceptors on the bill of the animal were probably used for prey detection instead. The specimens represent the oldest record of amniotes with extremely reduced visual capacity, utilizing non-visual cues for prey detection. The discovery reveals that the ecological diversity of marine predators was already high in the late Early Triassic, and challenges the traditional view that the ecological diversification of marine reptiles was delayed following the EPME.

Separating sexual dimorphism from other morphological variation in a specimen complex of fossil marine reptiles (Reptilia, Ichthyosauriformes, Chaohusaurus).

The Early Triassic Chaohu Fauna from Anhui Province, China, contains the oldest record of Mesozoic marine reptiles, such as Cartorhynchus and Sclerocormus. Most specimens from the fauna belong to the ichthyosauriform Chaohusaurus, more specifically resembling C. chaoxianensis. However, a wide range of morphological variation exists within about 40 skeletons that have been prepared, likely reflecting mixed signals from both sexual and taxonomic differences. We test whether the sexual and taxonomic signals are separable based on quantification, aided by the knowledge of sexual dimorphism in extant marine tetrapods. There are two different suites of dimorphism that divide the specimens differently from each other yet consistently within each suite, resulting in four morphotypes in combination, likely representing two sexes of two taxa. Presumed males have larger 'organ of prehension' sensu Darwin, specifically limbs in the present case, for a given body length. This sexing criterion is supported by the only specimen of a gravid female, which belongs to the morphotype with short limbs. Males also have larger skulls for the trunk length compared to females. This study demonstrates that sexual and taxonomic signals are separable in fossil reptiles, with a sufficient sample size and careful analyses.

Pre- versus post-mass extinction divergence of Mesozoic marine reptiles dictated by time-scale dependence of evolutionary rates.

The fossil record of a major clade often starts after a mass extinction even though evolutionary rates, molecular or morphological, suggest its pre-extinction emergence (e.g. squamates, placentals and teleosts). The discrepancy is larger for older clades, and the presence of a time-scale-dependent methodological bias has been suggested, yet it has been difficult to avoid the bias using Bayesian phylogenetic methods. This paradox raises the question of whether ecological vacancies, such as those after mass extinctions, prompt the radiations. We addressed this problem by using a unique temporal characteristic of the morphological data and a high-resolution stratigraphic record, for the oldest clade of Mesozoic marine reptiles, Ichthyosauromorpha. The evolutionary rate was fastest during the first few million years of ichthyosauromorph evolution and became progressively slower over time, eventually becoming six times slower. Using the later slower rates, estimates of divergence time become excessively older. The fast, initial rate suggests the emergence of ichthyosauromorphs after the end-Permian mass extinction, matching an independent result from high-resolution stratigraphic confidence intervals. These reptiles probably invaded the sea as a new ecosystem was formed after the end-Permian mass extinction. Lack of information on early evolution biased Bayesian clock rates.

Eccentricity and obliquity paced carbon cycling in the Early Triassic and implications for post-extinction ecosystem recovery.

The timing of marine ecosystem recovery following the End Permian Mass Extinction (EPME) remains poorly constrained given the lack of radiometric ages. Here we develop a high-resolution carbonate carbon isotope (δ(13)Ccarb) record for 3.20 million years of the Olenekian in South China that defines the astronomical time-scale for the critical interval of major evolutionary and oceanic events in the Spathian. δ(13)Ccarb documents eccentricity modulation of carbon cycling through the period and a strong obliquity signal. A shift in phasing between short and long eccentricity modulation, and amplification of obliquity, is nearly coincident with a 2% decrease in seawater δ(13)CDIC, the last of a longer-term stepped decrease through the Spathian. The mid-Spathian shift in seawater δ(13)CDIC to typical thermocline values is interpreted to record a major oceanic reorganization with global climate amelioration. Coincidence of the phasing shift with the first occurrence of marine reptiles (248.81 Ma), suggests that their invasion into the sea and the onset of a complex ecosystem were facilitated by restoration of deep ocean ventilation linked mechanistically to a change in the response of the oceanic carbon reservoir to astronomical forcing. Together these records place the first constraints on the duration of the post-extinction recovery to 3.35 myr.

A large aberrant stem ichthyosauriform indicating early rise and demise of ichthyosauromorphs in the wake of the end-Permian extinction.

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.

Adult sex ratio, sexual dimorphism and sexual selection in a Mesozoic reptile.

The evolutionary history of sexual selection in the geologic past is poorly documented based on quantification, largely because of difficulty in sexing fossil specimens. Even such essential ecological parameters as adult sex ratio (ASR) and sexual size dimorphism (SSD) are rarely quantified, despite their implications for sexual selection. To enable their estimation, we propose a method for unbiased sex identification based on sexual shape dimorphism, using size-independent principal components of phenotypic data. We applied the method to test sexual selection in Keichousaurus hui, a Middle Triassic (about 237 Ma) sauropterygian with an unusually large sample size for a fossil reptile. Keichousaurus hui exhibited SSD biased towards males, as in the majority of extant reptiles, to a minor degree (sexual dimorphism index -0.087). The ASR is about 60% females, suggesting higher mortality of males over females. Both values support sexual selection of males in this species. The method may be applied to other fossil species. We also used the Gompertz allometric equation to study the sexual shape dimorphism of K. hui and found that two sexes had largely homogeneous phenotypes at birth except in the humeral width, contrary to previous suggestions derived from the standard allometric equation.

A New Specimen of Carroll's Mystery Hupehsuchian from the Lower Triassic of China.

A new specimen of an enigmatic hupehsuchian genus is reported. The genus was first recognized by Robert L. Carroll and Zhi-ming Dong in 1991, who refrained from naming it because of the poor quality of the only specimen known at the time. After more than two decades, we finally report a second specimen of this genus, which remained unprepared until recently. The new specimen preserves most of the skeleton except the skull, allowing us to erect a new genus and species, Eretmorhipis carrolldongi. The new species shares many characters with Parahupehsuchus longus, including the strange axial skeleton that forms a bony body tube. However, the body tube is short in the new species, being limited to the pectoral region. The vertebral count and limb morphology considerably differ between the new species and P. longus. The forelimb of E. carrolldongi is markedly larger than its hind limb as in Hupehsuchus nanchangensis but unlike in P. longus. The new species is unique among hupehsuchians in a list of features. It has manual and pedal digits that spread radially, forming manus and pes that are almost as wide as long. The third-layer elements of the dermal armor are unusually large, spanning four vertebral segments, yet there are substantial gaps among them. With the addition of the unique paddle, it is now clear that Hupehsuchia had diverse forelimb morphologies spanning from paddles to flippers, unlike ichthyopterygians that were taxonomically more diverse yet only had flippers.

Lunge feeding in early marine reptiles and fast evolution of marine tetrapod feeding guilds.

Traditional wisdom holds that biotic recovery from the end-Permian extinction was slow and gradual, and was not complete until the Middle Triassic. Here, we report that the evolution of marine predator feeding guilds, and their trophic structure, proceeded faster. Marine reptile lineages with unique feeding adaptations emerged during the Early Triassic (about 248 million years ago), including the enigmatic Hupehsuchus that possessed an unusually slender mandible. A new specimen of this genus reveals a well-preserved palate and mandible, which suggest that it was a rare lunge feeder as also occurs in rorqual whales and pelicans. The diversity of feeding strategies among Triassic marine tetrapods reached their peak in the Early Triassic, soon after their first appearance in the fossil record. The diet of these early marine tetrapods most likely included soft-bodied animals that are not preserved as fossils. Early marine tetrapods most likely introduced a new trophic mechanism to redistribute nutrients to the top 10 m of the sea, where the primary productivity is highest. Therefore, a simple recovery to a Permian-like trophic structure does not explain the biotic changes seen after the Early Triassic.

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.

A small short-necked hupehsuchian from the lower Triassic of Hubei Province, China.

Hupehsuchia is a group of enigmatic Triassic marine reptiles that is known exclusively from two counties in Hubei Province, China. One of the common features of the group was a modestly long neck with nine to ten cervical vertebrae. We report a new species of Hupehsuchia, Eohupehsuchus brevicollis gen. et sp. nov., which for the first time shows a short neck in this group, with six cervicals. The configuration of the skull roof in Eohupehsuchus is also unique among Hupehsuchia, with narrow frontals and posteriorly shifted parietals, warranting recognition of a new species. The taxon superficially resembles Nanchangosaurus in retaining hupehsuchian plesiomorphies, such as low neural spines and small body size. However, its limbs are well-developed, unlike in Nanchangosaurus, although the latter genus is marginally larger in body length. Thus, the individual is unlikely to be immature. Also, Eohupehsuchus shares a suite of synapomorphies with Hupehsuchus, including the second and third layers of dermal ossicles above the dorsal neural spines. A phylogenetic analysis suggests that the new species is not the most basal hupehsuchian despite its short neck, and instead forms the sister taxon of Hupehsuchidae. Until recently, Hupehsuchia contained only two monotypic genera. Now there are at least four genera among Hupehsuchia, and the undescribed diversity is even higher. The left forelimb of the only specimen is incomplete, ending with broken phalanges distally. The breakage could only have occurred pre-burial. The individual may have been attacked by a predator and escaped, given that scavenging is unlikely.

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.

A carapace-like bony 'body tube' in an early triassic marine reptile and the onset of marine tetrapod predation.

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.

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.

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.

{µ-2-[4-(1,3-Benzothia-zol-2-yl)phen-yl]-2-aza-propane-1,3-dithiol-ato-κS,S':S,S'}bis-[tricarbonyl-iron(I)].

The title compound, [Fe(2)(C(15)H(12)N(2)S(3))(CO)(6)], was prepared as an aza-dithiol-atodiiron model for the active site of [FeFe]-hydrogenase. The Fe(2)S(2) core adopts a butterfly shape, with each metal having a pseudo square-pyramidal geometry. The N-substituted aza-dithiol-ate is µ(2)-κ(4)S,S':S,S'-coordinated to the Fe(CO)(3) moieties to form two fused six-membered rings with different conformations. The sum of the C-N-C angles around the N atom [356.85 (15)°] indicates a flattening of the trigonal-pyramidal geometry about the N atom and an increase in the degree of sp(2)-hybridization.