1. Ecosystems and Environment Research Programme, Faculty of Biological and Environmental Sciences, University of Helsinki, P.O. Box 65, University of Helsinki FI-00014, Finland
2. Helsinki Institute of Sustainability Science (HELSUS), University of Helsinki, Helsinki, Finland
3. Organismal and Evolutionary Biology Research Programme, Faculty of Biological and Environmental Sciences, University of Helsinki, P.O. Box 65, University of Helsinki FI-00014, Finland
4. Transport Institute, University of Manitoba, 181 Freedman Crescent, Winnipeg, Manitoba R3T 5V4, Canada
5. Department of Mathematics and Statistics, Faculty of Science, University of Helsinki, Helsinki, Finland