Cranial morphology of the late Oligocene Patagonian notohippid Rhynchippus equinus Ameghino, 1897 (Mammalia, Notoungulata) with emphases in basicranial and auditory region

Abstract

"Notohippidae" is a probably paraphyletic family of medium sized notoungulates with complete dentition and early tendency to hypsodonty. They have been recorded from early Eocene to early Miocene, being particularly diverse by the late Oligocene. Although Rhynchippus equinus Ameghino is one of the most frequent notohippids in the fossil record, there are scarce data about cranial osteology other than the classical descriptions which date back to the early last century. In this context, we describe the exceptionally preserved specimen MPEF PV 695 (based on CT scanning technique and 3D reconstruction) with the aim of improving our knowledge of the species, especially regarding auditory region (petrosal, tympanic and surrounding elements), sphenoidal and occipital complexes. Besides a modular description of the whole skull, osteological correlates identified on the basicranium are used to infer some soft-tissue elements, especially those associated with vessels that supply the head, mainly intracranially. One of the most informative elements was the petrosal bone, whose general morphology matches that expected for a toxodont. The endocranial surface, together with the surrounding parietal, basisphenoid, occipital, and squamosal, enabled us to propose the location and communication of main venous sinuses of the lateral head wall (temporal, inferior and sigmoid sinuses), whereas the tympanic aspect and the identification of a posterior carotid artery canal provided strong evidence in support of an intratympanic course of the internal carotid artery, a controversial issue among notoungulates. Regarding the arrangement of tympanic and paratympanic spaces, the preservation of the specimen allowed us to appreciate the three connected spaces that constitute a heavily pneumatized middle ear; the epitympanic sinus, the tympanic cavity itself, and the ventral expansion of the tympanic cavity through the notably inflated bullae. We hope this study stimulates further inquires and provides potentially informative data for future research involving other representatives of the order.