Taxonomy, morphology, masticatory function and phylogeny of heterodontosaurid dinosaurs

Heterodontosaurids comprise an important early radiation of small-bodied herbivores that persisted for approximately 100 My from Late Triassic to Early Cretaceous time. Review of available fossils unequivocally establishes Echinodon as a very small-bodied, late-surviving northern heterodontosaurid similar to the other northern genera Fruitadens and Tianyulong. Tianyulong from northern China has unusual skeletal proportions, including a relatively large skull, short forelimb, and long manual digit II. The southern African heterodontosaurid genus Lycorhinus is established as valid, and a new taxon from the same formation is named Pegomastax africanusgen. n., sp. n. Tooth replacement and tooth-to-tooth wear is more common than previously thought among heterodontosaurids, and in Heterodontosaurus the angle of tooth-to-tooth shear is shown to increase markedly during maturation. Long-axis rotation of the lower jaw during occlusion is identified here as the most likely functional mechanism underlying marked tooth wear in mature specimens of Heterodontosaurus. Extensive tooth wear and other evidence suggests that all heterodontosaurids were predominantly or exclusively herbivores. Basal genera such as Echinodon, Fruitadens and Tianyulong with primitive, subtriangular crowns currently are known only from northern landmasses. All other genera except the enigmatic Pisanosaurus have deeper crown proportions and currently are known only from southern landmasses.