Upstroke-based acceleration and head stabilization are the norm for the wing-propelled swimming of alcid seabirds

Alcids, a family of seabirds including murres, guillemots and puffins, exhibit the greatest mass-specific dive depths and durations of any birds or mammals. These impressive diving capabilities have motivated numerous studies on the biomechanics of alcid swimming and diving, with one objective being to compare stroke-acceleration patterns of swimming alcids with those of penguins, where upstroke and downstroke are used for horizontal acceleration. Studies of free-ranging, descending alcids have found that alcids accelerate in the direction of travel during both their upstroke and downstroke, but only at depths <20 m, whereas studies of alcids swimming horizontally report upstroke-based acceleration to be rare (<= 16% of upstrokes). We hypothesized that swimming trajectory, via its interaction with buoyancy, determines the magnitude of acceleration produced during the upstroke. Thus, we studied the stroke-acceleration relationships of five species of alcid swimming freely at the Alaska SeaLife Center using videography and kinematic analysis. Contrary to our prediction, we found that upstroke-based acceleration is very common (87% of upstrokes) during both descending and horizontal swimming. We reveal that head-damping - wherein an animal extends and retracts its head to offset periodic accelerations - is common in swimming alcids, underscoring the importance of head stabilization during avian locomotion.