The facial integument of centrosaurine ceratopsids: morphological and histological correlates of novel skin structures.

The horned dinosaur Pachyhinosaurus possesses rugose bony bosses across the skull roof in lieu of the projecting bony horn cores seen in most ceratopsians. This elaboration of typical ceratopsian ornaments provides an opportunity to test hypotheses of ceratopsian facial skin morphology and function. We analyze bone morphology and histology associated with several classes of skin features in extant amniotes using a classification tree analysis. We isolate key osteological and histological correlates for unpreserved skin structures, including both a pattern of pitting and resorption characteristic of muskox (Ovibos) frontal horn boss, and a pattern of metaplastic ossification characteristic of rhinoceros nasal horn boss. We also describe correlates for other skin features, such as epidermal scales and horn sheaths. Dermatocranial elements from centrosaurine ceratopsians are then examined for the same osteological and histological correlates. From this comparison we propose that the rugose bosses that replace horn cores in many centrosaurine dinosaurs, most notably Achelousaurus and Pachyrhinosaurus, were covered by a thick pad of cornified skin derived from the caudodorsal side of the primitive horn sheath comparable to the horny boss of extant muskoxen (Ovibos). We examine extant taxa with skin morphologies similar to Pachyrhinosaurus for consistent adaptive relationships between structure and behavior. We determine that high-energy headbutting is consistently associated with the acquisition of thick cornified pads, seen in muskoxen as well as helmeted hornbills [Buceros (=Rhinoplax) vigil] and African buffalo (Syncerus). The association of the bony ornaments of Pachyrhinosaurus with risky agonistic behaviors casts doubt on the role of species recognition as a primary selection pressure driving the diversity of all ceratopsian horns. We conclude that social selection (a broad form of intraspecific competition) is a more appropriate explanation for the diversity of centrosaurine ceratopsian ornaments in the Late Cretaceous.