1. School of Human and Environmental Sciences, University of Reading, Reading, RG6 6AB, UK
2. School of Geography and the Environment, University of Oxford, Oxford, OX1 3QY, UK
3. Centre for Ecology and Hydrology, Maclean Building, Crowmarsh Gifford, Wallingford, OX10 8BB, UK
4. Environment Agency, Reading, RG2 0SF, UK