1. Centre for Ecology and Conservation, College of Life and Environmental Sciences, University of Exeter, Tremough Campus, Penryn TR10 9EZ, UK
2. Max Planck Institute for Demographic Research, Konrad-Zuse-Strasse 1, 18057 Rostock, Germany
3. Department of Entomology, North Carolina State University, Gardner Hall, Raleigh, NC 27695-7613, USA