Springhares, flying and flightless scaly-tailed squirrels (Anomaluromorpha, Rodentia) are the squirrely mouse: comparative anatomy of the masticatory musculature and its implications on the evolution of hystricomorphy in rodents.

Anomaluromorpha is a particularly puzzling suborder of Rodentia. Endemic to Africa, this clade includes the extant genera Idiurus, Anomalurus, Zenkerella, and Pedetes. These rodents present an hystricomorphous condition of the skull, characterized by a large infraorbital foramen, which evolved independently within the mouse-related clade over a span of approximately 57 million years. They exhibit a high disparity in craniomandibular and dental morphology that has kept their phylogenetic affinities disputed for a long time. Given the past significance of masticatory morphotypes in establishing the classification of Rodentia, we propose to explore variations in the masticatory apparatus of Anomaluromorpha in order to evaluate whether its related features can offer additional data for systematics and contribute to our understanding of the complexity of hystricomorphy. In order to do so, we used traditional dissection and diffusible iodine-based contrast-enhanced computed tomography (diceCT) to accurately describe and compare the anatomy of the specimens. We found that the muscle morphology displays clear differentiation among each anomaluromorph taxonomic unit. Specifically, the masseteric complex of Anomaluromorpha exhibits distinctive synapomorphies such as the infraorbital part of the zygomaticomandibularis muscle being separated into a rostral and orbital part and an absence of a posterior part of the zygomaticomandibularis. Additionally, the orbital portion of the infraorbital part originates from a well-marked ridge and fossa at the level of its area of origin on the anteromedial wall of the orbital cavity, a feature that is absent in other members of the mouse-related clade. This evident bony feature, among others, is strongly associated with muscular anatomy and can contribute to ascertaining the taxonomic status of extinct representatives of the clade. Finally, we showed that the hystricomorphy of Anomaluromorpha largely differs from those of Ctenohystrica and Dipodoidea and that the definition of this morphotype is complex and cannot be reduced simply to the size of the opening of the infraorbital foramen.