1. Theoretical Biology Group, University of Groningen, PO Box 14, 9750 AA NN Haren, The Netherlands
2. Institute of Evolutionary Biology, School of Biological Sciences, University of Edinburgh, West Mains Road, Edinburgh EH9 3JT, UK
3. School of Biology, University of St Andrews, Harold Mitchell Building, St Andrews, Fife KY16 9TH, UK
4. Evolutionary Genetics Group, Centre for Ecological and Evolutionary Studies, University of Groningen, PO Box 14, 9750 AA NN Haren, The Netherlands