1. Institute of Evolutionary Biology, University of Edinburgh, Edinburgh EH9 3JT, UK.
2. NERC Centre for Ecology and Hydrology, Bush Estate, Penicuik, Midlothian EH26 0QB, UK.
3. Environmental and Evolutionary Biology, Institute of Biomedical and Life Sciences, University of Glasgow, Glasgow G12 8QQ, UK.
4. British Antarctic Survey, Natural Environment Research Council, High Cross, Madingley Road, Cambridge CB3 0ET, UK.