Kinematics of diving Atlantic puffins (Fratercula arctica L.): evidence for an active upstroke.

To examine the propulsion mechanism of diving Atlantic puffins (Fratercula arctica), their three-dimensional kinematics was investigated by digital analysis of sequential video images of dorsal and lateral views. During the dives of this wing-propelled bird, the wings are partly folded, with the handwings directed backwards. The wings go through an oscillating motion in which the joint between the radius-ulna and the hand bones leads the motion, with the wing tip following. There is a large rotary motion of the wings during the stroke, with the wings being pronated at the beginning of the downstroke and supinated at the end of the downstroke/beginning of the upstroke. Calculated instantaneous velocities and accelerations of the bodies of the birds show that, during the downstroke, the birds accelerate upwards and forwards. During the upstroke, the birds accelerate downwards and, in some sequences analysed, also forwards, but in most cases the birds decelerate. In all the upstrokes analysed, the forward/backward acceleration shows the same pattern, with a reduced deceleration or even a forward acceleration during 'mid' upstroke indicating the production of a forward force, thrust. Our results show that the Atlantic puffin can use an active upstroke during diving, in contradiction to previous data. Furthermore, we suggest that the partly folded wings of diving puffins might act as efficient aft-swept wingtips, reducing the induced drag and increasing the lift-to-drag ratio. A movie is available on-line.