1. Department of Animal and Plant Sciences, University of Sheffield, Sheffield S10 2TN, UK;
2. These authors contributed equally to this work;
3. Present address: School of Biological Sciences, Cruickshank Building, St Machar Drive, University of Aberdeen, Aberdeen AB24 3UU, UK
4. Department of Biology, University of York, PO Box 373, York YO10 5YW, UK;
5. School of Biology, Sir Harold Mitchell Building, University of St Andrews, St Andrews, Fife KY16 9TH, UK;
6. Unit of Comparative Plant Ecology, Department of Animal and Plant Sciences, University of Sheffield, Sheffield S10 2TN, UK;