1 Environment and Sustainability Institute, College of Life and Environmental Sciences, University of Exeter, Penryn, Cornwall TR10 9FE, UK. Electronic address: s.crowley@exeter.ac.uk.
2 Geography, College of Life and Environmental Sciences, University of Exeter, Exeter EX4 4RJ, UK.
3 Institute of Biological and Environmental Sciences, Zoology Building, Tillydrone Avenue, University of Aberdeen, Aberdeen AB24 2TZ, UK.
4 Environment and Sustainability Institute, College of Life and Environmental Sciences, University of Exeter, Penryn, Cornwall TR10 9FE, UK.