Cranial Kinesis in the Late Cretaceous Birds Hesperornis and Parahesperornis

A recently discovered skull of the Cretaceous toothed diving bird Hesperornis permits evaluation of previous descriptions of the skull, analysis of cranial kinesis in hesperornithid birds, and discussion of the evolution of avian cranial kinesis. We found no evidence in hesperornithids for mesokinesis, a system that in lizards involves relative movement of the frontals and parietals. "Maxillokinesis," the fore-aft movements of the palato-maxillary arcade, was also rejected. Evidence for cranial kinesis in hesperornithid birds involves the presence of a streptostylic quadrate bone of virtually modern morphology, three types of flexion zones at the posterior end of the upper jaw, and a hiatus in the nasal-interorbital septum. Among the forms of kinesis found in modern birds, prokinesis, in which the upper jaw moves as a unit, was the dominant type of cranial kinesis in hesperornithids as indicated by the rigid construction of the upper jaw, the position of the holorhinal nostril, and the anteriorly truncate mesethmoid bone; rhynchokinesis and amphikinesis were not possible. Prokinesis is considered primitive for at least the group including Hesperornithiformes and later birds. Discovery of prokinesis in Hesperornis represents the first time all of the morphological correlated of prokinesis have been identified in a bird plesiomorphically lacking pterygoid segmentation.