1. New Zealand Institute for Advanced Study and Allan Wilson Center for Molecular Ecology and Evolution, Massey University, Auckland 0745, New Zealand;
2. Max Planck Institute for Evolutionary Biology, 24306 Plön, Germany
3. Institut de Biologie de l'Ecole Normale Supérieure, UMR CNRS 8197 INSERM 1024, F-75005 Paris, France;