The social cost of shoaling covaries with predation risk in nine-spined stickleback, Pungitius pungitius, populations
Section snippets
Fish Sampling and Rearing
We collected adult sticklebacks from four populations (Fig. 1) before or during the early phase of the reproductive period in 2007. We sampled two ponds (both isolated), Pyöreälampi, Finland, and Bynästjärnen, Sweden, and two marine populations, the Baltic Sea near Helsinki, Finland, and the White Sea in the Levin Navolok Bay, Russia. The four populations are genetically (highly) isolated from each other as revealed by microsatellite markers (T. Shikano, G. Herczeg & J. Merilä; unpublished
Results
The first GLMM revealed a habitat-specific treatment effect on body size in 12-week-old fish (habitat: F1,2 = 0.30, P = 0.638; treatment: F1,376 = 53.86, P < 0.001; habitat*treatment: F1,376 = 49.32, P < 0.001). Fish from ponds were smaller in shoals than when kept individually, while there was no effect of treatment in marine fish. The population effect within habitat was nonsignificant (Z = 1.00, P = 0.159). The first GLM showed a similar pattern at the population level (population: F3,374 = 190.61, P < 0.001;
Discussion
Our results suggest that living in groups can have a considerable cost even when (1) food is not a limiting factor, there are no (2) predators or (3) parasites in the system, and even when (4) the constraints and costs of reproduction are ruled out. Hence, the manifested cost can only be related to sociality itself, resulting purely from contact with conspecifics. Furthermore, we found systematic differences in the presence of such a social cost between different population types: the cost was
Acknowledgments
We thank Victor Berger, Göran Englund, Tuomas Leinonen, Daniel Lussetti and Pirkko Siikamäki for helping us in organizing and executing the field sampling. Special thanks to the Oulanka Research Station and the White Sea Biological Station for their help. We are highly indebted to Esa Ranta and two anonymous referees for their constructive comments and John Loehr for correcting the English. We were supported by the Academy of Finland and Centre for International Mobility.
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