1. aInstitute of Evolutionary Biology, School of Biological Sciences, University of Edinburgh, Ashworth Laboratories, King’s Buildings, West Mains Road, Edinburgh, EH9 3JT, UK
2. bDepartment of Biological Sciences, University of Alabama, 300 Hackberry Lane, Box 870344, SEC Building, Tuscaloosa, AL 35487, USA
3. cDaphne du Maurier Building, Centre for Ecology and Conservation, Biosciences, College of Life and Environmental Sciences, University of Exeter, Cornwall Campus, Treliever Road, Penryn, Cornwall, TR10 9EZ, UK