In many aquatic taxa, formation traveling can reduce the energetic expenditure of locomotion by exploiting the vorticity trails shed by neighbors or through drafting. Cetaceans, especially odontocetes, often swim in groups; nevertheless, the possibility that whales gain energetic benefits from swimming in formation remains poorly studied, apart from mother-calf pairs. Between June and September in 2019 and 2021, we recorded aerial videos of Southern Resident killer whales (Orcinus orca) in the Salish Sea (USA) travelling in groups. We estimated whale tailbeat and breathing frequencies as proxies of the relative energetic costs of swimming, and tested the effect of swimming speed, relative positioning (e.g., leaders, whales in the middle of groups, or followers), sex and estimated size on these observed proxies. Our results reveal a complex relationship between physical characteristics, relative positioning, and energetic proxies. Intervals between respiration lasted longer in large-sized trailing individuals, but the overall breathing frequency was similar for all whales regardless of their position. The tailbeat frequency was mainly associated to whale sex, size, and swimming speed; in addition, tailbeat frequency showed a decreasing trend as the number of individuals in the formation increased. We found moderate evidence that position-based energetic effects may be present in the formation swimming of killer whales, and it is likely that additional factors such as social ties and hierarchies, play a key role in determining individual positioning in travelling groups.
Significance
Swimming in formation has been extensively studied in fish and other aquatic animals and has been documented to provide energetic advantages. Our understanding of the potential energetic benefits of wild cetacean formation swimming has been constrained by the difficulties of studying the movement of whale groups from traditional observation platforms. In recent years, non-invasive observations of cetaceans using unoccupied aerial systems have significantly improved the observation of these species in the wild, providing an exciting opportunity to better understand their behaviors and habits. Our results show a tendency for formation swimming to affect two energetic proxies (tailbeat frequency and the duration of underwater intervals between surfacing events). The results of this study set the stage for further research to identify the multiple determinants affecting killer whale formation swimming which go beyond purely energetic advantages, e.g. social relationships.