1. Centre for Ecology and Conservation, School of Biosciences, University of Exeter, Cornwall Campus, Penryn, Cornwall TR10 9EZ, UK
2. Centre for Agri-Environmental Research, School of Agriculture, Policy and Development, University of Reading, Earley Gate, Reading RG6 6AR, UK
3. Faculty of Biological Sciences, University of Leeds, Leeds LS2 9JT, UK