1. Organismal and Evolutionary Biology Research Programme, Faculty of Biological and Environmental Sciences, University of Helsinki, Helsinki, Finland
2. Tvärminne Zoological Station, University of Helsinki, Hanko, Finland
3. Finnish Museum of Natural History, University of Helsinki, Helsinki, Finland
4. Finnish Environment Institute, Helsinki, Finland
5. Department of Environmental and Biological Sciences, University of Eastern Finland, Kuopio, Finland
6. Ecology and genetics research unit, University of Oulu, Oulu, Finland