1. The Zeeman Institute for Systems Biology and Infectious Disease Epidemiology Research, School of Life Sciences and Mathematics Institute, University of Warwick, Coventry CV4 7AL, UK
2. Mathematics for Real World Systems Centre for Doctoral Training, Mathematics Institute, University of Warwick, Coventry CV4 7AL, UK
3. Odum School of Ecology, University of Georgia, Athens, GA, USA
4. Center for the Ecology of Infectious Diseases, University of Georgia, Athens, GA, USA