A new possible megalosauroid theropod from the Middle Jurassic Xintiangou Formation of Chongqing, People's Republic of China and its implication for early tetanuran evolution

Tetanurae is a special group of theropod dinosaurs that originated by the late Early Jurassic. It includes several early-diverging groups of generally large-bodied predators (megalosauroids, allosauroids, tyrannosauroid coelurosaurs) as well as morphologically disparate small-bodied coelurosaurs, including birds. Aspects of the evolutionary history of tetanurans remain contested, including the topology of their deep phylogenetic divergences (among Megalosauroidea, Allosauroidea and Coelurosauria). We report a new theropod, Yunyangosaurus puanensis gen. et sp. nov., based on a fragmentary specimen recovered from the Middle Jurassic Xintiangou Formation of Chongqing, southwestern China. It shares several features uniquely with some megalosauroids (the clade of megalosaurids + spinosaurids + piatnitzkysaurids), such as prominent rims around the anterior articular surfaces of cervical centra and bifurcated anterior dorsal neural spines (present in piatnitzkysaurids). Nevertheless, it also shows several features that are rare or absent among megalosauroids and more crownward tetanurans, including prominent spinopostyzgopophyseal laminae (also present in non-tetanurans and metriacanthosaurid allosauroids), flat anterior articular surfaces of the cervical centra (also present in piatnitzkysaurids and some earlier-diverging tetanurans), and the presence of a posterior pneumatic foramen or fossa (absent in most tetanurans, but sporadically present in some cervical vertebrae of piatnitzkysaurids). Yunyangosaurus therefore presents a combination of derived and apparently primitive character states that are not seen in other theropods. This suggests that patterns of morphological evolution associated with deep tetanuran divergences were more complex than currently recognized, with implications for understanding the character evolution in theropods.