Cooperation under predation risk: experiments on costs and benefits

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Abstract
Two fish that cooperatively inspect a predator may have negotiated the share of the risk that each takes. A test of both the costs of predator inspection dependent on the distance from which the predator is approached and the potential benefits of cooperation was carried out strictly experimentally. We made either singletons or pairs of dead sticklebacks, Gasterosteus aculeatus, approach hungry pike, Esox lucius, by remote control according to an algorithm that mimicked natural inspection. The predation risk of both single inspectors and parallel inspecting pairs increased with closer inspection distances. A member of an inspecting pair had only about half the risk of that of a single inspector. In pairs, a companion diluted the lead fish's risk of being caught, depending on its distance behind the leader. The absolute risk difference between leader and follower was greatest for close inspection distances and decreased further away from the predator. The leader's relative risk increased with its distance ahead of the laggard. However, for a given distance between leader and laggard, the relative risks of the two fish remained similar with distance from the predator. The cost side of the inequalities that define a 'Prisoner's Dilemma' has thus been measured for this system. In a second experiment the 'attack deterrence hypothesis' of predator inspection (i.e. inspection decreases attack probability) was tested. The pike was offered a choice between two sticklebacks, one of which had carried out a predator inspection visit. There was no indication of attack deterrence through predator inspection.
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