1. Department of Genetics, Evolution, and Environment, University College London , Gower Street, London WC1E 6BT , UK
2. CoMPLEX, University College London , Gower Street, London WC1E 6BT , UK
3. CNRS, UMR7144 Adaptation et Diversité en Milieu Marin, Station Biologique de Roscoff , Roscoff 29680 , France
4. Sorbonne Universités, UPMC Université Paris VI , Roscoff 29680 , France
5. School of Biological Sciences, University of East Anglia , Norwich Research Park, Norwich NR4 7TJ , UK