Craniomandibular osteology of Manidens condorensis (Ornithischia: Heterodontosauridae) from the upper Lower Jurassic of Argentina

Heterodontosauridae is a clade that appears early in the ornithischian fossil record, and includes small-bodied, highly specialized species characterized by an unusual heterodont dentition. Although known from relatively few taxa, the early representation of the clade and unsolved phylogenetic relationships within heterodontosaurids and among early ornithischians implies that novel information has a marked effect on broader phylogenetic hypotheses and our understanding of early diversification patterns within Ornithischia. This paper describes the cranial osteology of the heterodontosaurid Manidens condorensis based on computed micro-tomographic scans of MPEF-PV 3211 and MPEF-PV 3809. This enabled more detailed descriptions of previously recognized bones, corrections to the literature, and the identification of undescribed elements. We present a new skull reconstruction and propose an emended diagnosis in light of novel anatomical information. Areas of jaw muscle attachment were identified and compared with Heterodontosaurus and Lesothosaurus, and mandibular function among heterodontosaurids is discussed. Our results indicate that diverse skull morphologies and functions existed among Early Jurassic ornithischians, with Manidens being intermediate between the plesiomorphic cranial shape and function associated with a generalist diet in ornithischians such as Tianyulong and Lesothosaurus, and the more derived cranial construction specialized for herbivory identified in heterodontosaurids from South Africa such as Heterodontosaurus.