New material and systematic re-evaluation of Medusaceratops lokii (Dinosauria, Ceratopsidae) from the Judith River Formation (Campanian, Montana)

Medusaceratops lokii Ryan, Russell, and Hartman, 2010 is an enigmatic taxon of ceratopsid represented by partial parietals from the Mansfield bonebed in the Campanian Judith River Formation, Montana. Originally, all ceratopsid material collected from this bonebed was referred to the centrosaurine ceratopsid Albertaceratops, but subsequently two parietals were designated the types of the chasmosaurine, M. lokii, in part, because they were interpreted to have three epiparietals bilaterally. Here we describe new material from the bonebed that allows a systematic revision of the taxon. A revised reconstruction of the frill, informed by newly discovered parietals, reveals that M. lokii had a broad midline ramus and at least five epiparietals (ep) around the margin of the frill, both traits that are characteristic of Centrosaurinae. From medial to lateral, the epiparietal ornamentation consists of a small, variably procurving epiparietal (ep 1), an anterolaterally curving pachyostotic hook (ep 2), a smaller pachyostoic process (ep 3), and two small triangular epiparietals (ep 4 and 5). A phylogenetic analysis of ceratopsids, which is the first to include Medusaceratops, indicates that M. lokii is a unique, early centrosaurine ceratopsid taxon that is more closely related to Centrosaurini and Pachyrhinosaurini than Nasutoceratopsini. No unequivocal chasmosaurine bones or diagnostic material from any other ceratopsid could be identified from the Mansfield bonebed, suggesting that it represents one of the oldest occurrences of a monodominant accumulation of a centrosaurine ceratopsid on record.