The cranial anatomy of the neornithischian dinosaur Thescelosaurus neglectus.

Though the dinosaur Thescelosaurus neglectus was first described in 1913 and is known from the relatively fossiliferous Lance and Hell Creek formations in the Western Interior Basin of North America, the cranial anatomy of this species remains poorly understood. The only cranial material confidently referred to this species are three fragmentary bones preserved with the paratype, hindering attempts to understand the systematic relationships of this taxon within Neornithischia. Here the cranial anatomy of T. neglectus is fully described for the first time based on two specimens that include well-preserved cranial material (NCSM 15728 and TLAM.BA.2014.027.0001). Visual inspection of exposed cranial elements of these specimens is supplemented by detailed CT data from NCSM 15728 that enabled the examination of otherwise unexposed surfaces, facilitating a complete description of the cranial anatomy of this species. The skull of T. neglectus displays a unique combination of plesiomorphic and apomorphic traits. The premaxillary and 'cheek' tooth morphologies are relatively derived, though less so than the condition seen in basal iguanodontians, suggesting that the high tooth count present in the premaxillae, maxillae, and dentaries may be related to the extreme elongation of the skull of this species rather than a retention of the plesiomorphic condition. The morphology of the braincase most closely resembles the iguanodontians Dryosaurus and Dysalotosaurus, especially with regard to the morphology of the prootic. One autapomorphic feature is recognized for the first time, along with several additional cranial features that differentiate this species from the closely related and contemporaneous Thescelosaurus assiniboiensis. Published phylogenetic hypotheses of neornithischian dinosaur relationships often differ in the placement of the North American taxon Parksosaurus, with some recovering a close relationship with Thescelosaurus and others with the South American taxon Gasparinisaura, but never both at the same time. The new morphological observations presented herein, combined with re-examination of the holotype of Parksosaurus, suggest that Parksosaurus shares a closer relationship with Thescelosaurus than with Gasparinisaura, and that many of the features previously cited to support a relationship with the latter taxon are either also present in Thescelosaurus, are artifacts of preservation, or are the result of incomplete preparation and inaccurate interpretation of specimens. Additionally, the overall morphology of the skull and lower jaws of both Thescelosaurus and Parksosaurus also closely resemble the Asian taxa Changchunsaurus and Haya, though the interrelationships of these taxa have yet to be tested in a phylogenetic analysis that includes these new morphological data for T. neglectus.