1. Institute of Evolutionary Biology, School of Biological Sciences, University of Edinburgh, Edinburgh EH9 3FL, UK
2. Institute of Immunology and Infection Research, School of Biological Sciences, University of Edinburgh, Edinburgh EH9 3FL, UK
3. Faculty of Biological Sciences, School of Biology, University of Leeds, Leeds LS2 9JT, UK
4. Moredun Research Institute, Pentlands Science Park, Bush Loan, Midlothian EH26 0PZ, UK