1. Department of Zoology, Edward Grey Institute, University of Oxford, South Parks Road, Oxford OX1 3PS, UK
2. Centre d'Etudes Biologiques de Chizé, CNRS-UPR1934, Villiers en Bois, France
3. Theoretical Biology, and Behavioural Ecology and Self-organization, University of Groningen, PO Box 11103, 9700 CC Groningen, The Netherlands