1. Institute of Biotechnology, University of Cambridge, Tennis Court Road, Cambridge CB2 1QT, UK, Institute of Biomedical & Life Sciences, University of Glasgow, Glasgow G12 8QQ, UK, NERC Centre for Ecology & Hydrology Oxford, Mansfield Road, Oxford OX1 3SR, UK, and CNAP, Department of Biology, University of York, York YO10 5YW, UK