1. Research Centre for Ecological Change, Organismal and Evolutionary Biology Research Programme, Faculty of Biological and Environmental Sciences University of Helsinki Helsinki Finland
2. Finnish Environment Institute Jyväskylä Finland
3. Finnish Environment Institute Helsinki Finland
4. Centre for Biodiversity Dynamics, Department of Biology Norwegian University of Science and Technology Trondheim Norway
5. Department of Biological and Environmental Science University of Jyväskylä Jyväskylä Finland