1. Organismal and Evolutionary Biology Research Programme, Faculty of Biological and Environmental Sciences, University of Helsinki, Finland
2. Tvärminne Zoological Station, Faculty of Biological and Environmental Sciences, University of Helsinki, Finland
3. Ecology and Genetics Research Unit, Faculty of Science, University of Oulu, Finland
4. Institut für Zoologie und Evolutionsbiologie, Biologie I, Universität Regensburg, Regensburg, Germany