Swimming mechanics and behavior of the shallow-water brief squidLolliguncula brevis

Although squid are among the most versatile swimmers and rely on a unique locomotor system, little is known about the swimming mechanics and behavior of most squid, especially those that swim at low speeds in inshore waters. Shallow-water brief squid Lolliguncula brevis, ranging in size from 1.8 to 8.9 cm in dorsal mantle length (DML), were placed in flumes and videotaped, and the data were analyzed using motion-analysis equipment. Flow visualization and force measurement experiments were also performed in water tunnels. Mean critical swimming speeds (U(crit)) ranged from 15.3 to 22.8 cm s(-1), and mean transition speeds (U(t); the speed above which squid swim exclusively in a tail-first orientation) varied from 9.0 to 15.3 cm s(-1). At low speeds, negatively buoyant brief squid generated lift and/or improved stability by positioning the mantle and arms at high angles of attack, directing high-speed jets downwards (angles >50 degrees ) and using fin activity. To reduce drag at high speeds, the squid decreased angles of attack and swam tail-first. Fin motion, which could not be characterized exclusively as drag- or lift-based propulsion, was used over 50-95 % of the sustained speed range and provided as much as 83.8 % of the vertical and 55.1 % of the horizontal thrust. Small squid (<3.0 cm DML) used different swimming strategies from those of larger squid, possibly to maximize thrust benefits from vortex ring formation. Furthermore, brief squid employed various unsteady behaviors, such as manipulating funnel diameter during jetting, altering arm position and swimming in different orientations, to boost swimming performance. These results demonstrate that locomotion in slow-swimming squid is complex, involving intricate spatial and temporal interactions between the mantle, fins, arms and funnel.