A New Species of Prolibytherium (Ruminantia,Mammalia) from Pakistan,and the Functional Implications of an Atypical Atlanto-Occipital Morphology

We describe a new species of Prolibytherium, P. fusus, sp. nov., from the lower Miocene of Pakistan, thus extending the genus to Asia. Prolibytherium is otherwise known only from Libya. This species differs from Prolibytherium magnieri in several basioccipital and atlanto-occipital morphologies. Namely, the posterior basioccipital tuberosities are continuous at the midline and lack the elevated transverse ridge seen in P. magnieri, and the notch formed between the lateral occipital condyles and paraoccipital process is lower. Both species of Prolibytherium have a characteristic ventrally fused occipital condyle at the midline, with a notably fuller circumferential articular surface. Prolibytherium magnieri also has thickened dorsal and ventral arches of the atlas. These specimens also possess a longitudinal groove for the Eustachian tube extending from the alisphenoid canal to the bullae, and a second deep grove isolating the basisphenoid bone from the temporal bone. These, plus several other atlanto-occipital morphologies strengthen the cervical support of the head. This is especially important for Prolibytherium, as the taxon possesses massive aliform cranial appendages. We relate the approximation of the occipital condyles to a convergent state in two giraffids (Giraffokeryx punjabiensis and Schansitherium tafeli), each of which possesses multiple pairs of ossicones, presumably necessitating a strengthened atlanto-occipital joint.