1. Faculty of Biological and Environmental Sciences, Organismal and Evolutionary Biology Research Programme , P.O.BOX 65, FI 00014 , Helsinki University, Finland
2. Tvärminne Zoological station , J.A. Palménintie 260, FI 10900 Hanko , Finland
3. Faculty of Science, Ecology and Genetics Research Unit , FI 90014, University of Oulu , Finland