The cervical anatomy of Samotherium, an intermediate-necked giraffid

Giraffidae are represented by many extinct species. The only two extant taxa possess diametrically contrasting cervical morphology, as the okapi is short-necked and the giraffe is exceptionally long-necked. Samotherium major, known from the Late Miocene of Samos in Greece and other Eurasian localities, is a key extinct giraffid; it possesses cervical vertebrae that are intermediate in the evolutionary elongation of the neck. We describe detailed anatomical features of the cervicals of S. major, and compare these characteristics with the vertebrae of the two extant giraffid taxa. Based on qualitative morphological characters and a quantitative analysis of cervical dimensions, we find that the S. major neck is intermediate between that of the okapi and the giraffe. Specifically, the more cranial (C2-C3) vertebrae of S. major represent a mosaic of features shared either with the giraffe or with the okapi. The more caudal (C5-C7) S. major vertebrae, however, appear transitional between the two extant taxa, and hence are more unique. Notably, the C6 of S. major exhibits a partially excavated ventral lamina that is strong cranially but completely absent on the caudal half of the ventral vertebral body, features between those seen in the giraffe and the okapi. Comprehensive anatomical descriptions and measurements of the almost-complete cervical column reveal that S. major is a truly intermediate-necked giraffid. Reconstructions of the neck display our findings.