1Centre for Ecology and Conservation, College of Life and Environmental Sciences, University of Exeter, Penryn, Cornwall, TR10 9FE, UK
2Institute of Zoology, Zoological Society of London, Regent's Park, London, NW1 4RY UK
3Environment and Sustainability Institute, Centre for Geography and Environmental Science, University of Exeter, Penryn, Cornwall TR10 9FE, UK