1. Centre for Ecology and Hydrology (CEH), Maclean Building, Benson Lane, Crowmarsh Gifford, Wallingford Oxfordshire, OX10 8BB, UK
2. Bournemouth University, Centre for Conservation, Ecology & Environmental Change, Christchurch House, Talbot Campus, Poole Dorset, BH12 5BB, UK
3. Department of Geography, School of Geography, Archaeology and Earth Resources, University of Exeter, Amory Building, Rennes Drive, Exeter EX4 4RJ, UK