1. Institute of Infection, Veterinary and Ecological Sciences, University of Liverpool, Liverpool L69 7ZB, UK
2. Institute of Ecology and Evolution, School of Biological Sciences, University of Edinburgh, Edinburgh EH9 3FL, UK
3. School of Biological Sciences, Cardiff University, Cardiff CF10 3AX, UK
4. Spatial Epidemiology Lab (SpELL), Université Libre de Bruxelles, B-1050 Bruxelles, Belgium