1. Queen Elizabeth House, University of Oxford, 21 St Giles, Oxford, OX1 3LA, UK,
2. NERC Centre for Ecology and Hydrology, Hill of Brathens, Banchory, Aberdeenshire, AB31 4BW, UK,
3. Department of Biological Sciences, University of Durham, Science Laboratories, South Road, Durham, DH1 3LE, UK