The diapsid origin of turtles.

The origin of turtles has been a persistent unresolved problem involving unsettled questions in embryology, morphology, and paleontology. New fossil taxa from the early Late Triassic of China (Odontochelys) and the Late Middle Triassic of Germany (Pappochelys) now add to the understanding of (i) the evolutionary origin of the turtle shell, (ii) the ancestral structural pattern of the turtle skull, and (iii) the phylogenetic position of Testudines. As has long been postulated on the basis of molecular data, turtles evolved from diapsid reptiles and are more closely related to extant diapsids than to parareptiles, which had been suggested as stem group by some paleontologists. The turtle cranium with its secondarily closed temporal region represents a derived rather than a primitive condition and the plastron partially evolved through the fusion of gastralia.