Comparative morphology of the hominin and African ape hyoid bone, a possible marker of the evolution of speech.

This study examines the morphology of the hyoid in three closely related species, Homo sapiens, Pan troglodytes, and Gorilla gorilla. Differences and similarities between the hyoids of these species are characterized and used to interpret the morphology and affi nities of the Dikika A. afarensis, Kebara 2 Neanderthal, and other fossil hominin hyoid bones. Humans and African apes are found to have distinct hyoid morphologies. In humans the maximum width across the distal tips of the articulated greater horns is usually slightly greater than the maximum length (distal greater horn tip to most anterior point of the hyoid body in the midline). A different pattern is usually found in the African ape hyoids, which have much greater maximum lengths. In humans, the hyoid body is also much more anteroposteriorly shallow in proportion to its height and width, and this is true for all age classes. The Dikika australopithecine hyoid body proportions are chimpanzeelike. A discriminant function analysis, using a larger subadult sample from the three extant species than that reported by Alemseged et al. (2006), confirms this finding. The Kebara hyoid dimensions (body alone, and articulated body and greater horns) are almost all within the observed range for human hyoids. Discriminant functions clearly distinguish human from African ape hyoids and classify the Kebara 2 hyoid as human (confirming the finding of Arensburg et al. 1989). Our virtual dissection of a chimpanzee air sac system shows its subhyoid extension into the dorsal hyoid body. Following Alemseged et al. (2006), the expanded bulla characteristic of the African ape and australopithecine hyoid body is therefore interpreted as refl ecting the presence of such a laryngeal air sac extension. Its absence in the human, Neanderthal, and H. heidelbergensis (Atapuerca SH) hyoids implicates the loss of the laryngeal air sacs as a derived Neanderthal and modern human trait, which evolved no later than the middle Pleistocene. If, as has been argued by de Boer (2012), the loss of the air sac helped to enhance perceptual discrimination of speech sounds, then this derived hyoid morphology can be added to the list of fossil markers of the capacity for speech.