1. Gatty Marine Laboratory, School of Biology, Division of Environmental and Evolutionary Biology, University of St Andrews, St Andrews, Scotland, KY16 8LB, UK
2. Centro Austral de Investigaciones Cientificas (CADIC), Consejo Nacional de Investigaciones Cientificas y Tecnicas (CONICET), CC92, Ushuaia, 9410, Tierra del Fuego, Argentina
3. British Antarctic Survey, High Cross, Madingley Road, Cambridge, CB3 OET,UK