1. Edward Grey Institute, Department of Zoology, University of Oxford, South Parks Road, Oxford OX1 3PS, UK.
2. Centre d'Ecologie Fonctionelle et Evolutive, Unité Mixte de Recherche CNRS 5175, 1919 Route de Mende, 34293 Montpellier Cedex 5, France.
3. Institute of Evolutionary Biology, School of Biological Sciences, University of Edinburgh, Edinburgh EH9 3JT, UK.