Spatial Group Structure as Potential Mechanism to Maintain Cooperation in Fish Shoals of Unrelated Individuals

In public goods games, group members share the benefits created through individual investments. If benefits are shared equally, individual contributions readily become altruistic, and hence, a social dilemma arises in which group interests and individual interests oppose each other. However, contributions to public goods can be self-serving if each investor gains a disproportionate benefit from its own contribution. This scenario may hold for our study system, the interactions of shoaling-unrelated scalefin anthias Pseudanthias squamipinnis and the ectoparasitic blenny Plagiotremus tapeinosoma. The blenny bites off pieces of skin, mucus and scales from anthias that in return may chase the blenny. Chasing the blenny represents a public good as it makes the parasite change victim species for its next attack. Laboratory experiments using artificial Plexiglas hosts suggest that one reason why individuals contribute to the public good is that the blenny may specialise on non-punishing free-riders. Here, we investigated how far a spatial structure within the shoal and limited space use by the blenny may contribute to punishment being self-serving. Field observation reveals that anthias indeed live in spatially structured groups and that blennies have preferred areas for attacks. Thus, some anthias individuals are consistently more exposed to blennies than others and hence may gain disproportional benefits from their punishment. In conclusion, spatial structure may contribute to the maintenance of punishment in blennyanthias interactions even when groups are large.