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First published online July 25, 2005
Journal of Experimental Biology 208, 2939-2949 (2005)
Published by The Company of Biologists 2005
doi: 10.1242/jeb.01733

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Aquatic wing flapping at low Reynolds numbers: swimming kinematics of the Antarctic pteropod, Clione antarctica

Brendan J. Borrell^1,*, Jeremy A. Goldbogen¹ and Robert Dudley^1,2

¹ Department of Integrative Biology, University of California, Berkeley, CA 94720, USA
² Smithsonian Tropical Research Institute, P.O. Box 2072, Balboa, Republic of Panama

^* Author for correspondence (e-mail: bborrell{at}berkeley.edu)

Accepted 6 June 2005

We studied swimming kinematics of the Antarctic pteropod, Clione antarctica, to investigate how propulsive forces are generated by flexible oscillating appendages operating at low Reynolds numbers (10<Re<100). We filmed ten ascending individuals at 125 frames s^–1 from two orthogonal views, and reconstructed three-dimensional coordinates of the wing tip and body. Each half-stroke of flapping consisted of distinct power and recovery phases, which were of approximately equal duration in both the upstroke and the downstroke. As pteropods ascended, the body traced a sawtooth path when viewed laterally. The magnitude of these oscillations decreased with body mass, and larger animals (operating at Re>25) exhibited gliding during the recovery phase of each half-stroke. Maximum translational and rotational accelerations of the body occurred at the initiation of each power phase, suggesting that rotational circulation, the acceleration reaction, and wake recapture may all potentially contribute to vertical force production. Individual contributions of these mechanisms cannot, however, be assessed from these kinematic data alone. During recovery phases of each half-stroke, C. antarctica minimized adverse drag forces by orienting the wings parallel to flow and by moving them along the body surface, possibly taking advantage of boundary layer effects. Vertical force production was altered through changes in the hydrodynamic angle of attack of the wing that augmented drag during the power phase of each half-stroke. At higher translational velocities of the body, the inclination of the power phase also became more nearly vertical. These results indicate that, in addition to serotonin-mediated modulation of wingbeat frequency reported previously in Clione, geometric alteration of wingbeat kinematics offers a precise means of controlling swimming forces.

Key words: Clione antarctica, drag, flapping flight, kinematics, lift, pteropod, swimming

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