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 nonlinear
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 finbeat
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
