1. T. H. Huxley School of the Environment, Earth Sciences and Engineering, Imperial College of Science, Technology and Medicine, Silwood Park, Ascot, Berks SL5 7PY, UK;
2. Present address: Department of Environmental Science, University of Bradford, West Yorkshire BD7 1DP, UK; Present address: Department of Agricultural and Environmental Science, University of Newcastle upon Tyne, NE1 7RU, UK