How to capture fish in a school? Effect of successive predator attacks on seabird feeding success.

Prey aggregations, such as fish schools, attract numerous predators. This typically leads to the formation of multispecific groups of predators. These aggregations can be seen both as a place of increased competition and as a place of possible facilitation between predators. Consequently, the functional role of such predator-prey aggregation is uncertain, and its effect on individual feeding success is virtually unknown. Using underwater film footage of different predators feeding on fish schools during the sardine run in South Africa, we directly measured the in situ feeding success of individual Cape gannets Morus capensis in different foraging situations. We determined the types of Cape gannet attacks (direct plunge dive or plunge dive followed by underwater pursuit) and we measured the occurrences and timing of attacks from the different species (mostly Cape gannets and long-beaked common dolphins Delphinus capensis). We also estimated the size of the targeted fish schools. These observations were complemented with a simulation model to evaluate the cumulative effect of successive predator attacks on the prey aggregation structure. The probability to capture a fish in one feeding attempt by Cape gannets averaged 0·28. It was lower when gannets engaged in underwater prey pursuit after the plunge compared to direct plunge (0·13 vs. 0·36). We found no effect of the number of prey on gannets' feeding success. However, the timing and frequency of attacks influenced strongly and positively the feeding success of individuals. The probability to capture a fish was the lowest (0·16) when no attack occurred in the few seconds (1-15 s) prior to a dive and the highest (˜0·4, i.e. more than twice) when one or two attacks occurred during this time window. The simulation model showed that a prey aggregation disorganized just after an attack and that the maximum of disturbance was obtained a few seconds after the initiation of the successive attacks. Our study suggests that, in multispecies predator assemblages, the cumulative effect (through disorganization of school cohesiveness) of the multiple species attacking a prey aggregation may increase the feeding success of each individual. Therefore, facilitation between predators is likely to overcome competition in these multispecific assemblages.