Analysis of the bite force and mechanical design of the feeding mechanism of the durophagous horn shark Heterodontus francisci.

Three-dimensional static equilibrium analysis of the forces generated by the jaw musculature of the horn shark Heterodontus francisci was used to theoretically estimate the maximum force distributions and loadings on its jaws and suspensorium during biting. Theoretical maximum bite force was then compared with bite forces measured (1) voluntarily in situ, (2) in restrained animals and (3) during electrical stimulation of the jaw adductor musculature of anesthetized sharks. Maximum theoretical bite force ranged from 128 N at the anteriormost cuspidate teeth to 338 N at the posteriormost molariform teeth. The hyomandibula, which connects the posterior margin of the jaws to the base of the chondrocranium, is loaded in tension during biting. Conversely, the ethmoidal articulation between the palatal region of the upper jaw and the chondrocranium is loaded in compression, even during upper jaw protrusion, because H. francisci's upper jaw does not disarticulate from the chondrocranium during prey capture. Maximum in situ bite force averaged 95 N for free-swimming H. francisci, with a maximum of 133 N. Time to maximum force averaged 322 ms and was significantly longer than time away from maximum force (212 ms). Bite force measurements from restrained individuals (187 N) were significantly greater than those from free-swimming individuals (95 N) but were equivalent to those from both theoretical (128 N) and electrically stimulated measurements (132 N). The mean mass-specific bite of H. francisci was greater than that of many other vertebrates and second highest of the cartilaginous fishes that have been studied. Measuring bite force on restrained sharks appears to be the best indicator of maximum bite force. The large bite forces and robust molariform dentition of H. francisci correspond to its consumption of hard prey.