1. Institute of Infection, Immunity and Inflammation, Centre for Biomolecular Sciences, University Park, University of NottinghamNottingham NG7 2RD, UK
2. Institute of Evolutionary Biology, School of Biological Sciences, University of EdinburghKing's Buildings, Edinburgh EH9 3JT, UK
3. Department of Biology, Queen's UniversityKingston, Ontario K7L 3N6, Canada
4. Department of Mathematics and Statistics, Queen's UniversityKingston, Ontario K7L 3N6, Canada