Up for grabs: prey herding by penguins facilitates shallow foraging by volant seabirds

Visual and olfactory signals are commonly used by seabirds to locate prey in the horizontal domain, but foraging success depends on prey depth and the seabird's ability to access it. Facilitation by diving seabirds has long been hypothesized as a mechanism to elevate deep prey to regions more accessible to volant seabirds, but this has never been demonstrated empirically. Footage from animal-borne video loggers deployed on African penguins was analysed to establish if volant seabird encounters involved active cuing by seabirds on penguins to obtain prey and, during mutual prey encounters, if interactions were driven by the vertical displacement of prey by penguins. Independent of prey biomass estimates, we found a strong inverse relationship between penguin group size, a proxy for visibility, and the time elapsed from the start of penguins' dive bouts to their first encounter with other seabirds. Most mutual prey encounters (7 of 10) involved schooling prey elevated from depths greater than 33 m by penguins and only pursued by other seabird species once prey was herded into shallow waters. This is likely to enhance foraging efficiency in volant seabird species. As such, penguins may be integral to important processes that influence the structure and integrity of marine communities.