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Squids use a pulsed jet and fin movements to swim both arms-first (forward) and tail-first (backward). Given the complexity of the squid multi-propulsor system, 3D velocimetry techniques are required for the comprehensive study of wake dynamics. Defocusing digital particle tracking velocimetry, a volumetric velocimetry technique, and high-speed videography were used to study arms-first and tail-first swimming of brief squid Lolliguncula brevis over a broad range of speeds [0-10 dorsal mantle lengths (DML) s(-1)] in a swim tunnel. Although there was considerable complexity in the wakes of these multi-propulsor swimmers, 3D vortex rings and their derivatives were prominent reoccurring features during both tail-first and arms-first swimming, with the greatest jet and fin flow complexity occurring at intermediate speeds (1.5-3.0 DML s(-1)). The jet generally produced the majority of thrust during rectilinear swimming, increasing in relative importance with speed, and the fins provided no thrust at speeds &gt;4.5 DML s(-1). For both swimming orientations, the fins sometimes acted as stabilizers, producing negative thrust (drag), and consistently provided lift at low/intermediate speeds (&lt;2.0 DML s(-1)) to counteract negative buoyancy. Propulsive efficiency (η) increased with speed irrespective of swimming orientation, and η for swimming sequences with clear isolated jet vortex rings was significantly greater (η=78.6±7.6%, mean±s.d.) than that for swimming sequences with clear elongated regions of concentrated jet vorticity (η=67.9±19.2%). This study reveals the complexity of 3D vortex wake flows produced by nekton with hydrodynamically distinct propulsors.

Volumetric flow imaging reveals the importance of vortex ring formation in squid swimming tail-first and arms-first