1 Department of Biology, University of Oxford, Oxford, UK.
2 School of Natural and Environmental Sciences, Newcastle University, Newcastle-upon-Tyne, UK.
3 Mixed Research Unit (UMR 5558) "Biometry & Evolutionary Biology", University Claude Bernard Lyon 1, Campus de la Doua, Bâtiment Mendel, Villeurbanne Cedex, France.
4 School of Biological Sciences, University of Aberdeen, Aberdeen, UK.
5 Department of Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, Michigan, USA.