Ontogeny And Variation Of The Pachycephalosaurine Dinosaur Sphaerotholus Buchholtzae, And Its Systematics Within The Genus

The ontogeny and taxonomy of the dome-headed pachycephalosaurs are topics of continued debate. Pachycephalosaurid diversity in the Maastrichtian of North America is particularly controversial, and the validity and composition within the genus Sphaerotholus remains unresolved. While the type species, S. goodwini, is generally considered valid, debate has centred around the validity and taxonomy of S. buchholtzae and S. edmontonensis. Here we employ morphometrics, histology and phylogenetic analysis to resolve these issues.An ontogenetic assessment of S. buchholtzae (N > 20) confirms previously observed ontogenetic morphologies: inflation of the frontoparietal dome, obliteration of tesserate surface texture, blunting of the peripheral nodes and decreasing void space within the dome. While linear bivariate analysis finds S. edmontonensis nested within S. buchholtzae, three-dimensional geometric morphometrics supports S. edmontonensis and S. buchholtzae as distinct species. Phylogenetic analysis recovers a Sphaerotholus lineage with S. goodwini as sister-taxon to a clade formed by S. edmontonensis and S. buchholtzae.The stratigraphic, phylogenetic, morphometric and ontogenetic data support the validity of both S. edmontonensis and S. buchholtzae, and their placement within the genus Sphaerotholus. The morphological similarities of S. edmontonensis to immature S. buchholtzae, and the slightly older geological age of S. edmontonensis, suggest that S. edmontonensis and S. buchholtzae may be part of an anagenetic lineage.