Sciuromorphy outside rodents reveals an ecomorphological convergence between squirrels and extinct South American ungulates

Notoungulates were a diverse group of South American ungulates that included the rodent-like typotherians. However, they are typically compared with other ungulates and interpreted as grazers. Here we present the first detailed reconstruction of the masticatory muscles of the pachyrukhine typotherians Paedotherium and Tremacyllus. An outstanding feature is the presence of a true sciuromorph condition, defined by an anterior portion of the deep masseter muscle originating from a wide zygomatic plate that reaches the rostrum, a trait traceable since the Oligocene pachyrukhines. Consequently, pachyrukhines are the first case of sciuromorph non-rodent mammals. This morphology would have allowed them to explore ecological niches unavailable for the exclusively hystricomorph coexisting rodents. This innovative acquisition seems to be synchronous in Pachyrukhinae and sciuromorph rodents and related to hard-food consumption. We postulate the expansion of nut and cone trees during the major environmental changes at Eocene−Oligocene transition as a potential trigger for this convergence.