Avoidance or escape? Discriminating between two hypotheses for the function of schooling in threespine sticklebacks.

In many species, there are antipredator benefits of grouping with conspecifics. For example, animals often aggregate to better avoid potential predators (the 'avoidance hypothesis'). Animals also often group together in direct response to predators to facilitate escape (the 'escape hypothesis'). The avoidance hypothesis predicts that animals with previous experience with predation risk will aggregate more than animals without experience with predation risk. In contrast, the escape hypothesis predicts that immediate exposure to predation risk causes animals to aggregate. We simultaneously tested these two nonexclusive hypotheses in threespine sticklebacks, Gasterosteus aculeatus. Schooling behaviour (time spent schooling, latency to school and time schooling in the middle of the school) was quantified with a mobile model school. Fish that had been chased by a model predator in the past schooled more, started schooling faster and spent a marginally greater proportion of time schooling in the middle of the school than fish that had not been chased. In contrast, there was no difference in the schooling behaviour of fish that were immediately exposed to either a model pike or a control, stick stimulus. A second experiment confirmed that fish perceived the model pike and stick differently: fish froze more often in the presence of the model pike, oriented to it more often and spent less time with the model pike than they did with the stick. These results provide strong support for the avoidance hypothesis.