1. Laboratoire de Biométrie et Biologie Evolutive; UMR 5558; CNRS; Université Lyon 1; Université de Lyon; Campus de la Doua 69622 Villeurbanne France
2. University Lyon 1; Lyon France
3. Centre d'Ecologie et des Sciences de la Conservation; UMR 7204; Muséum National d'Histoire Naturelle; 61 rue Buffon CP 53 75005 Paris France
4. Department of Evolutionary Genetics; Max Planck Institute of Evolutionary Biology; August-Thienemann-Str. 2 24306 Plön Germany
5. Department of Life and Environmental Sciences; Faculty of Science and Technology; Bournemouth University; Christchurch House Talbot Campus Poole Dorset BH12 5BB UK