1. Centre for Ecology and Hydrology; Environment Centre Wales; Deiniol Road; Bangor; LL57 2UW; UK
2. School of Biological Sciences; Bangor University; Deiniol Road; Bangor; Gwynedd; LL57 2UW; UK
3. Centre for Ecology and Hydrology; Lancaster University; Lancaster; LA1 4AP; UK
4. Soils Research Centre; Department of Geography and Environmental Science; University of Reading; Reading; RG6 6DW; UK