The origin of the turtle body plan remains one of the great mysteries of reptile evolution. The anatomy of turtles is highly derived, which renders it difficult to establish the relationships of turtles with other groups of reptiles. The oldest known turtle, Proganochelys from the Late Triassic period of Germany, has a fully formed shell and offers no clue as to its origin. Here we describe a new 220-million-year-old turtle from China, somewhat older than Proganochelys, that documents an intermediate step in the evolution of the shell and associated structures. A ventral plastron is fully developed, but the dorsal carapace consists of neural plates only. The dorsal ribs are expanded, and osteoderms are absent. The new species shows that the plastron evolved before the carapace and that the first step of carapace formation is the ossification of the neural plates coupled with a broadening of the ribs. This corresponds to early embryonic stages of carapace formation in extant turtles, and shows that the turtle shell is not derived from a fusion of osteoderms. Phylogenetic analysis places the new species basal to all known turtles, fossil and extant. The marine deposits that yielded the fossils indicate that this primitive turtle inhabited marginal areas of the sea or river deltas.
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.
Sineoamphisbaena hexatabularis Wu et al., 1993 is the earliest known amphisbaenian represented by well-preserved cranial and postcranial material. It reveals a mosaic of generalized lizard-like features and amphisbaenian characters. Most distinctive of the latter are features of cranial consolidation adaptive for a fossorial way of life. Phylogenetic analyses strongly confirm the monophyly of the Amphisbaenia inclusive of S. hexatabularis. The Amphisbaenia is diagnosed by a suite of apomorphic characters. The available evidence suggests a probable Amphisbaenia–Macrocephalosauridae relationship within the Scincomorpha. This is supported primarily by the unique modifications of the palate and temporal region of the skull. It is argued here that the Amphisbaenia evolved in Central Asia during the Cretaceous, in response to the transition from a perennial lacustrine environment to a dry, semiarid eolian environment. The relatively primitive morphology indicates that S. hexatabularis was not permanently subterranean. The further derived modifications of later forms are associated with tunneling in an environment of more compact soils.
A new Triassic archosaurian from China shows a number of aquatic specializations, of which the most striking is the extreme lateral compression of the long tail. Others that may also reflect aquatic adaptations include plate like scapula and coracoid, elongate neck with extremely long and slender ribs, and reduction of osteoderms. In contrast, its pelvic girdle and hind limb have no aquatic modifications. Anatomic features, taphonomy, and local geological data suggest that it may have lived in a coastal-island environment. This lifestyle, convergent with some Jurassic marine crocodyli-forms that lived at least 40 million years later and the saltwater species of extant Crocodylus, contradicts with the prevailing view that Triassic archosaurians were restricted to non-marine ecosystems. Its mosaic anatomy represents a previously unknown ecomorph within primitive archosaurians.
Sauropterygians form the largest and most diverse group of ancient marine reptiles that lived throughout nearly the entire Mesozoic era (from 250 to 65 million years ago). Although thousands of specimens of this group have been collected around the world since the description of the first plesiosaur in 1821 (ref. 3), no direct evidence has been found to determine whether any sauropterygians came on shore to lay eggs (oviparity) like sea turtles, or gave birth in the water to live young (viviparity) as ichthyosaurs and mosasauroids (marine lizards) did. Viviparity has been proposed for plesiosaur, pachypleurosaur and nothosaur sauropterygians, but until now no concrete evidence has been advanced. Here we report two gravid specimens of Keichousaurus hui Young from the Middle Triassic of China. These exquisitely preserved specimens not only provide the first unequivocal evidence of reproductive mode and sexual dimorphism in sauropterygians, but also indicate that viviparity could have been expedited by the evolution of a movable pelvis in pachypleurosaurs. By extension, this has implications for the reproductive pattern of other sauropterygians and Mesozoic marine reptiles that possessed a movable pelvis.
The early evolution of turtles continues to be a contentious issue in vertebrate palaeontology. Recent reports have suggested that they are diapsids, but the position of turtles within Diapsida is controversial and the sequence of acquisition of turtle synapomorphies remains unclear. Here we describe a Triassic turtle from China that has a mixture of derived characters and plesiomorphic features. To our knowledge, it represents the earliest known stem turtle with an edentulous beak and a rigid puboischiadic plate. The discovery of this new form reveals a complex early history of turtles.
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