Vectored jets power arms-first and tail-first turns differently in brief squid with assistance from fins and keeled arms

Squids maneuver to capture prey, elude predators, navigate complex habitats and deny rivals access to mates. Despite the ecological importance of this essential locomotive function, limited quantitative data on turning performance and wake dynamics of squids are available. To better understand the contribution of the jet, fins and arms to turns, the role of orientation (i.e. arms first versus tail first) in maneuvering, and the relationship between jet flow and turning performance, kinematic and 3D velocimetry data were collected in tandem from brief squid, Lolliguncula brevis. The pulsed jet, which can be vectored to direct flows, was the primary driver of most turning behaviors, producing flows with the highest impulse magnitude and angular impulse about the main axis of the turn (yaw) and secondary axes (roll and pitch). The fins and keeled arms played subordinate but important roles in turning performance, contributing to angular impulse, stabilizing the maneuver along multiple axes and/or reducing rotational resistance. Orientation affected turning performance and dynamics, with tail-first turns being associated with greater impulse and angular impulse, longer jet structures, higher jet velocities and greater angular turning velocities than arms-first turns. Conversely, arms-first turns involved shorter, slower jets with less impulse, but these directed short pulses resulted in lower minimum length-specific turning radii. Although the length-to-diameter ratio (L/D) of ejected jet flow was a useful metric for characterizing vortical flow features, it, by itself, was not a reliable predictor of angular velocity or turning radii, which reflects the complexity of the squid multi-propulsor system.