[1] Univ Oxford, Dept Zool, Oxford, England
[2] Univ Oxford, Merton Coll, Oxford, England
[3] London Sch Hyg & Trop Med, Ctr Math Modelling Infect Dis, Dept Infect Dis Epidemiol, London, England
[4] Univ Cambridge, Dept Appl Math & Theoret Phys, Cambridge, England
[5] Harvard TH Chan Sch Publ Hlth, Dept Immunol & Infect Dis, Boston, MA USA
[6] Univ East Anglia, Sch Biol Sci, Norwich, Norfolk, England