1. Department of Zoology, University of Oxford, South Parks Road, Oxford OX1 3PS, UK
2. Department of Biomedical Sciences, University of Oxford, South Parks Road, Oxford OX1 3UD, UK
3. Division of Veterinary Pathology, Infection and Immunity, School of Clinical Veterinary Science, University of Bristol, Langford BS40 5DU, UK
4. Department of Animal and Poultry Science, University of Guelph, Ontario, N1G 2W1, Canada