1. Department of Earth Sciences, The Open University, Milton Keynes MK7 6AA, UK.
2. National Environment Research Council Centre for Ecology and Hydrology, Monks Wood, Huntingdon PE17 2LS, UK.
3. Department of Biology, Villanova University, Villanova, PA 19085, USA.
4. Department of Biological Sciences, The Open University, Milton Keynes MK7 6AA, UK.