Efficient foraging strategies result in a predator spatially overlapping with its prey, foraging in the most profitable patches, and minimizing the time transiting between patches. Previous studies investigating baleen whale foraging strategies have generally focused on investigating spatial overlap with prey patches, patch profitability or movement within feeding grounds. The present study investigated the fine-scale strategies of movement between individual prey schools and larger prey patches in humpback whales (Megaptera novaeangliae) bubble-feeding on sand lance (Ammodytes spp.) in and around the Stellwagen Bank National Marine Sanctuary, USA. The goal was to investigate the presence of hierarchically nested spatial structures in both sand lance patches and whale bubble-feeding behavior, and to compare the scales and geometry of these patches between predator and prey behavior on each hierarchical scale. Furthermore, the profitability of sand lance schools in feeding areas was investigated.
Using animal-borne tag technology to record underwater movement of whales in combination with surface observations of whale behavior, the locations of bubble-feeding events were identified. Concurrent hydroacoustic measurements of the prey distribution in the water column were used to identify the locations and energetic parameters of sand lance schools around tagged whales. First Passage Time analysis was used to determine the spatial scale of individual bubble-feeding events. Based on spatial proximity, feeding events and prey schools were grouped into larger feeding bouts and prey patches to investigate the presence of hierarchically nested scales.
Up to three hierarchy levels were found in bubble-feeding behavior of nine whales tagged on six days between 2008 and 2012, and up to five hierarchy levels in the sand lance prey field around the tagged whales. There was a significant positive correlation between the lengths of whale bubble-feeding bouts and the lengths of sand lance patches over three common hierarchy levels. On each hierarchy level, the lengths of whale bubble-feeding bouts were significantly smaller than those of sand lance patches. Mean inter-feeding bout distances were significantly positively correlated with mean inter-prey patch distances over two hierarchy levels. Mean distances between feeding events were similar to the mean distances between prey schools. On larger hierarchy levels, mean inter-bout distances were greater than mean distances among prey patches. Mean school height and density tended to be greater in schools recorded inside than outside of feeding bout areas.
The prey field structures found here were likely a result of the specific habitat requirements of sand lance. The results of this study suggest that the tagged whales were able to adapt their foraging movement to the structure of the prey field. By feeding on neighboring schools, whales could minimize the time spent between prey schools. On larger spatial scales, whales did not feed on neighboring prey patches. This could be a result of decreased abilities to find the nearest patch, or because, rather than restricting their foraging movement to neighboring patches, the whales were targeting specific patches. The foraging movement observed in this study led to spatial overlap of the tagged whales with sand lance schools that were characterized by properties rendering them more energetically profitable for bubble-feeding whales.
While hierarchically structured foraging movement has been found in other marine predators, this is the first study that demonstrates this kind of foraging mechanism for baleen whales