Abstract
The distribution of individuals is often the outcome of conflicting demands, such as between predator avoidance and reproduction. A factor that has seldom been considered in studies on habitat choice is time-dependent changes in risk-taking. We investigated the distribution of threespine sticklebacks, Gasterosteus aculeatus, over two breeding seasons and found it to change with time towards shallower areas with a more open habitat structure. Shallow and structurally less complex habitats were probably favorable due to a higher reproductive rate, but costly due to an increased risk of predation. Contrary to expectation, changing predation pressure was not a predictor of the shift in habitat use and, thus, not the proximate cue. Instead date was the main predictor. This suggests that increased risk-taking in relation to predation contributed to the habitat shift. The possibility was supported by a laboratory experiment that showed sticklebacks to take larger risks and prefer more predator-exposed areas at the end of the season than at the start of the season. These results demonstrate that temporal changes in risk-taking occur and can influence habitat choice, which points to the importance of considering risk-taking, in addition to predation pressure, when studying the effect of predators on distribution.





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Acknowledgments
We thank Kai Lindström, Mika Kilpi and Jerker Wuorinen for discussions and Dik Heg for comments on the manuscript. Tvärminne Zoological Station and the School of Fisheries in Kirjala provided equipment and facilities for carrying out the study. Financial support was provided from Oscar Öflund Foundation, Nordenskiöldsamfundet and the Academy of Finland to U.C. The experiments comply with the current laws of the country in which they were performed.
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Candolin, U., Voigt, HR. Do changes in risk-taking affect habitat shifts of sticklebacks?. Behav Ecol Sociobiol 55, 42–49 (2003). https://doi.org/10.1007/s00265-003-0678-0
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DOI: https://doi.org/10.1007/s00265-003-0678-0


