Marginocephalia (/mär′jə-nō-sə-făl′ē-ən/ Latin: margin-head) is a clade of ornithischian dinosaurs that is characterized by a bony shelf or margin at the back of the skull. These fringes were likely used for display. There are two clades included in Marginocephalia: the thick-skulled Pachycephalosauria and the horned Ceratopsia. All members of Marginocephalia were primarily herbivores.[citation needed] They basally used gastroliths to aid in digestion of tough plant matter until they convergently evolved tooth batteries in Neoceratopsia (or "new Ceratopsia") and Pachycephalosauria. Marginocephalia first evolved in the Jurassic Period and became more common in the Cretaceous.[1] They are basally small facultative quadrupeds while derived members of the group are large obligate quadrupeds.[2] Primitive marginocephalians are found in Asia, but the group migrated upwards into North America.[3]
Pachycephalosaurs, or "thick-headed reptiles", have primitive features that include basally small sized bodies, obligate bipedalism, and simple teeth with one row in operation at a time that are replaced as they are worn down. As they evolved, pachycephalosaurs evolved much thicker and advanced skull roofs including dome forms with horn-like ornamentation. Some research suggests these domes were used like helmets for protection while head-butting members in intraspecific combat.[4] Some research suggests their necks were not strong enough to support such an impact.[1] Flat-headed pachycephalosaur specimens have been found in Asia, and there is great controversy on the meaning of these flat heads. Recent research suggests the flat heads could be a juvenile state before developing the dome shape in the adult stage. It could also be evidence of sexual dimorphism with the female being more flat-headed.[5][6]
Ceratopsians, or "horned-faces", differ from pachycephalosaurs in the presence of a rostral bone, or beak. They are also known for having a jugal horn and a thin parietal-squamosal shelf that extends back and up into a frill.[7] This frill could have been used for anchoring jaw muscles, as well as for display.[1] The horns were likely used for establishing dominance, or defending territories.[8] It is also possible they were a factor in sexual display and species recognition. One of the basalmost members of this group is Psittacosaurus, which is one of the most species-rich dinosaur genera from Asia. Ceratopsians later evolved into very large quadrupeds with elaborate facial horns such as Triceratops, Styracosaurus, and Centrosaurus. There was no change in richness of species throughout the Cretaceous before the Cretaceous-Paleogene extinction.[9]
Marginocephalians have simple, peg-like teeth surrounded by rhamphotheca, a horny sheath of keratin. The teeth are arranged in batteries for easy replacement and have serrations which may have been useful for cutting up vegetation.[1] Marginocephalia evolved several methods for breaking down vegetation. Pachycephalosaurs had especially large abdomens with broad girths and elongate sacral ribs suggesting the presence of a large stomach. This is presumed to have been useful for breaking down tough vegetation through bacterial fermentation.[10] Another adaptation for advanced vegetation digestion is seen in Ceratopsians, which evolved features to improve their chewing apparatus. Derived ceratopsians have vertical grinding surfaces on their teeth to maximize break-down of tough vegetation.[11] There is also evidence of advanced adductor musculature that extends from a large coronoid process on the mandible up to the ceratopsian frill, which would increase chewing force.[12] Pachycephalosaurs had gastroliths to help in digestion of food, but only primitive ceratopsians, such as Psittacosaurus, have been found to have gastroliths.[13]