1. Ecological Genetics Research Unit, Organismal and Evolutionary Biology Research Programme, Faculty of Biological and Environmental Sciences, University of Helsinki, FI-00014, Finland
2. CSIRO Agriculture and Food, GPO Box 1600, Canberra, ACT 2601, Australia
3. Research Unit of Mathematical Sciences, University of Oulu, FI-90014, Finland
4. Area of Ecology and Biodiversity, School of Biological Sciences, The University of Hong Kong, Hong Kong SAR