Evolution and mechanics of long jaws in butterflyfishes (family Chaetodontidae)

We analyzed the functional morphology and evolution of the long jaws found in several butterflyfishes. We used a conservative reanalysis of an existing morphological dataset to generate a phylogeny that guided our selection of seven short- and long-jawed taxa in which to investigate the functional anatomy of the head and jaws: Chaetodon xanthurus, Prognathodes falcifer (formerly Chaetodon falcifer), Chelmon rostratus, Heniochus acuminatus, Johnrandallia nigrirostris, Forcipiger flavissimus, and F. longirostris. We used manipulations of fresh, preserved, and cleared and stained specimens to develop mechanical diagrams of how the jaws might be protruded or depressed. Species differed based on the number of joints within the suspensorium. We used high-speed video analysis of five of the seven species (C. xanthurus, Chel. rostratus, H. acuminatus, F. flavissimus, and F. longirostris) to test our predictions based on the mechanical diagrams: two suspensorial joints should facilitate purely anteriorly directed protrusion of the lower jaw, one joint should allow less anterior protrusion and result in more depression of the lower jaw, and no joints in the suspensorium should constrain the lower jaw to simple ventral rotation around the jaw joint, as seen in generalized perciform fishes. We found that the longest-jawed species, F. longirostris, was able to protrude its jaws in a predominantly anterior direction and further than any other species. This was achieved with little input from cranial elevation, the principal input for other known lower jaw protruders, and is hypothesized to be facilitated by separate modifications to the sternohyoideus mechanism and to the adductor arcus palatini muscle. In F. longirostris the adductor arcus palatini muscle has fibers oriented anteroposteriorly rather than medial-laterally, as seen in most other perciforms and in the other butterflyfish studied. These fibers are oriented such that they could rotate the ventral portion of the quadrate anteriorly, thus projecting the lower jaw anteriorly. The intermediate species lack modification of the adductor arcus palatini and do not protrude their jaws as far (in the case of F. flavissimus) or in a purely anterior fashion (in the case of Chel. rostratus). The short-jawed species both exhibit only ventral rotation of the lower jaw, despite the fact that H. acuminatus is closely related to Forcipiger.