Boxfishes (Teleostei: Ostraciidae) as a model system for fishes swimming with many fins: kinematics.

Swimming movements in boxfishes were much more complex and varied than classical descriptions indicated. At low to moderate rectilinear swimming speeds (<5 TL s(-1), where TL is total body length), they were entirely median- and paired-fin swimmers, apparently using their caudal fins for steering. The pectoral and median paired fins generate both the thrust needed for forward motion and the continuously varied, interacting forces required for the maintenance of rectilinearity. It was only at higher swimming speeds (above 5 TL s(-1)), when burst-and-coast swimming was used, that they became primarily body and caudal-fin swimmers. Despite their unwieldy appearance and often asynchronous fin beats, boxfish swam in a stable manner. Swimming boxfish used three gaits. Fin-beat asymmetry and a relatively non-linear swimming trajectory characterized the first gait (0--1 TL s(-1)). The beginning of the second gait (1--3 TL s(-1)) was characterized by varying fin-beat frequencies and amplitudes as well as synchrony in pectoral fin motions. The remainder of the second gait (3--5 TL s(-1)) was characterized by constant fin-beat amplitudes, varying fin-beat frequencies and increasing pectoral fin-beat asynchrony. The third gait (>5 TL s(-1)) was characterized by the use of a caudal burst-and-coast variant. Adduction was always faster than abduction in the pectoral fins. There were no measurable refractory periods between successive phases of the fin movement cycles. Dorsal and anal fin movements were synchronized at speeds greater than 2.5 TL s(-1), but were often out of phase with pectoral fin movements.