Anatomical description and digital reconstruction of the skull of Jeholosaurus shangyuanensis (Dinosauria, Ornithopoda) from China

Ornithopod dinosaurs appeared during the Middle Jurassic, but it was in the Lower Cretaceous they started their successful evolutionary history. Different phylogenies describing the evolutionary relationships of Ornithopoda are mostly based on cranial features, however there is a lack of well-preserved and complete skulls for the basal member of the clade, hampering our knowledge on the mode and tempo of these herbivorous dinosaurs. Here we describe YLSNHM 01942, a well-preserved skull of a juvenile neornithischian from the Liaoning Province of China. The specimen was scanned with a μCT scan, and all the elements were segmented and extrapolated for description. The specimen shows a ventral deformation due to the compression of the sediment, and a few rostral elements were artificially added. The specimen is attributed to the basal ornithopod Jeholosaurus shangyuanensis because of the presence of a large foramen in the quadratojugal, however it lacks the nodular ornamentation on the postorbital and jugal, herein interpreted as an ontogenetic feature. This, together with the disarticulation degree of the cranial elements, suggest YLSNHM 01942 represents a juvenile Jeholosaurus. The endosseous labyrinth is tentatively reconstructed, although the disarticulation of the neurocranial bones hampers its complete reconstruction. Thanks to the analysis of previously undescribed inner neurocranial bones (such as the prootics, the exoccipital/ophistotic, basioccipital, and basisphenoid), we improve the previous phylogenetical scoring for J. shangyuanensis, and perform a phylogenetical analysis adding the basal ornithopod Changmiania liaoningensis and the recently re-evaluated Ajkaceratops kozmai. The phylogenetical analysis reports a well-supported base of Ornithopoda, with C. liaoningensis as the most basal ornithopod, and a resolved topology for Nanosaurus agilis, Changchunsaurus parvus, Haya griva, Yandusaurus hongheensis and J. shangyuanensis.