1. Institutes of Evolution, Immunology, and Infection Research, School of Biological Sciences, Ashworth Laboratories, University of Edinburgh, Edinburgh EH9 3JT Scotland, UK.
2. CABI Bioscience at Department of Agricultural Sciences, Imperial College London, Wye Campus, Wye, Kent, TN25 5AH, UK.
3. Division of Biology and NERC Centre for Population Biology, Imperial College London, Wye Campus, Wye, Kent, TN25 5AH, UK.