1. Department of Zoology, Oxford University, South Parks Road, Oxford OX1 3PS, UK
2. Biosciences, University of Exeter, Cornwall Campus, Penryn TR10 9EZ, UK
3. Institute of Evolutionary Biology, University of Edinburgh, Edinburgh EH9 3JT, UK
4. Centre for Immunity, Infection and Evolution, School of Biological Sciences, Ashworth Laboratories, University of Edinburgh, Edinburgh EH9 3JT, UK