1. Division of Environmental and Evolutionary Biology, University of GlasgowGraham Kerr Building, Glasgow G12 8QQ, UK
2. Department of Ecology and Evolution, University of LausanneBiology Building, 1015 Lausanne, Switzerland
3. School of Biological Sciences, University of BristolBristol BS8 1UG, UK
4. Laboratoire Evolution et Diversité BiologiqueBâtiment 4R3, UMR 5174 CNRS, Université Paul Sabatier, 118 route de Narbonne, 31062 Toulouse Cedex 9, France