Flipper strokes can predict energy expenditure and locomotion costs in free-ranging northern and Antarctic fur seals

Flipper strokes have been proposed as proxies to estimate the energy expended by marine vertebrates while foraging at sea, but this has never been validated on free-ranging otariids (fur seals and sea lions). Our goal was to investigate how well flipper strokes correlate with energy expenditure in 33 foraging northern and Antarctic fur seals equipped with accelerometers, GPS, and time-depth recorders. We concomitantly measured field metabolic rates with the doubly-labelled water method and derived activity-specific energy expenditures using fine-scale time-activity budgets for each seal. Flipper strokes were detected while diving or surface transiting using dynamic acceleration. Despite some inter-species differences in flipper stroke dynamics or frequencies, both species of fur seals spent 3.79 +/- 0.39 J/kg per stroke and had a cost of transport of similar to 1.6-1.9 J/kg/m while diving. Also, flipper stroke counts were good predictors of energy spent while diving (R-2 = 0.76) and to a lesser extent while transiting (R-2 = 0.63). However, flipper stroke count was a poor predictor overall of total energy spent during a full foraging trip (R-2 = 0.50). Amplitude of flipper strokes (i.e., acceleration amplitude x number of strokes) predicted total energy expenditure (R-2 = 0.63) better than flipper stroke counts, but was not as accurate as other acceleration-based proxies, i.e. Overall Dynamic Body Acceleration.