Digital skull anatomy of the Oligocene North American tortoise Stylemys nebrascensis with taxonomic comments on the species and comparisons with extant testudinids of the Gopherus–Manouria clade

The anatomy of North American tortoises is poorly understood, despite a rich fossil record from the Eocene and younger strata. Stylemys nebrascensis is a particularly noteworthy turtle in this regard, as hundreds of specimens are known from Oligocene deposits, and as this species is one of the earliest fossil turtles to have been described in the scientific literature. Since its initial description based on a shell, many specimens with more complete material have been referred to Stylemys nebrascensis. Here, we review and confirm the referral of an important historic specimen to Stylemys nebrascensis, which includes shell, non-shell postcranial, and skull material. This allows us to document unique skull features of Stylemys nebrascensis (e.g., an unusual ‘poststapedial canal’ that connects the posterior skull surface with the cavum acustico-jugulare) and to refer another well-preserved skull to the species. Based on computed-tomography scanning of these two skulls, we provide a detailed description of the cranial and mandibular osteology of Stylemys nebrascensis. Stylemys nebrascensis has a combination of plesiomorphic skull characteristics (e.g., retention of a medial jugal process) and derived traits shared with extant gopher tortoises (e.g., median premaxillary ridge) that suggest it may be a stem-representative of the gopher tortoise lineage. This supports the hypothesis that extant and fossil tortoises from North America form a geographically restricted clade that split from Asian relatives during the Paleogene.