title: Structure and function of the horn shark (Heterodontus francisci) cranium through ontogeny: development of a hard prey specialist.
abstract: The horn sharks (Heterodontidae: Chondrichthyes) represent one of four independent evolutions of durophagy in the cartilaginous fishes. We used high-resolution computed tomography (CT scanning) to visualize and quantify the mineralized tissue of an ontogenetic series of horn sharks. CT scanning of neonatal through adult California horn sharks (Heterodontus francisci) confirmed that this technique is effective for examining mineralized tissue in even small (<10 mm) specimens. The jaw joint is among the first areas to become mineralized and is the most heavily mineralized area in the cranium of a neonatal horn shark. The hyoid is also well mineralized, although the poorly mineralized molariform teeth indicate that the neonatal animal may be a suction feeder on softer prey. The symphysis of the jaws never mineralizes, in sharp contrast to the condition in the hard prey-crushing stingrays. Digitally reslicing the CT scans along the jaws allowed measurement of the second moment of area (Ina). Assuming that the jaws are made of the same material at all ages, Ina is an indicator of the flexural stiffness of the jaws. In all sizes of shark the lower jaws were stiffer than the upper and the stiffness increased in the area of the molariform teeth. The central region of the jaws, where the rami meet, support cuspidate grasping teeth and has the lowest Ina. The spotted eagle ray (Aetobatus narinari), a hard prey-crushing stingray, shows a different pattern of flexural stiffness, with the peak at the central part of the jaws where the prey is reduced between flattened tooth plates. Although the eagle ray jaws have a higher Ina than the horn shark, they are also far more heavily mineralized. When the relative amounts of mineralization are taken into account, horn sharks do better with what mineral they have than does the eagle ray. With a tight jaw joint and loose mandibular symphysis, as well as nearly opposite patterns of stiffness in the jaws, it is clear that two of the clades of hard prey specialists use very different methods for cracking the hard prey problem.
authors.affiliation: Ecology and Evolutionary Biology, University of California, Irvine, California 92697, USA. asummers@uci.edu
authors.forename: Adam P
authors.initials: AP
authors.lastname: Summers
authors.forename: Richard A
authors.initials: RA
authors.lastname: Ketcham
authors.forename: Timothy
authors.initials: T
authors.lastname: Rowe
mesh_terms: D000818:Animals; D000831:Animals, Newborn; D005075:Biological Evolution; D001732:Bite Force; D002113:Calcification, Physiologic / Q000502:physiology; D005247:Feeding Behavior / Q000502:physiology; D005260:Female; D006928:Hyoid Bone / Q000033:anatomy & histology / Q000000981:diagnostic imaging / Q000254:growth & development; D007568:Jaw / Q000033:anatomy & histology* / Q000000981:diagnostic imaging; D008297:Male; D008409:Mastication / Q000502:physiology; D008445:Maxillofacial Development / Q000502:physiology*; D010802:Phylogeny; D012754:Sharks / Q000033:anatomy & histology* / Q000254:growth & development*; D012860:Skates, Fish / Q000033:anatomy & histology / Q000254:growth & development; D012886:Skull / Q000033:anatomy & histology* / Q000000981:diagnostic imaging / Q000254:growth & development*; D013704:Temporomandibular Joint / Q000033:anatomy & histology / Q000000981:diagnostic imaging / Q000254:growth & development; D014057:Tomography, X-Ray Computed; D014070:Tooth / Q000033:anatomy & histology / Q000254:growth & development