1. Institute for Immunology and Infection Research, School of Biological Sciences, University of Edinburgh, Edinburgh EH9 3FL, UK
2. Unité PhEED, Laboratoire de Santé Animale, Site de Normandie, ANSES, RD675, 1443012 Goustranville, France
3. Institute of Evolutionary Biology, Ashworth Laboratories, School of Biological Sciences, University of Edinburgh, Edinburgh EH9 3JT, UK