[1] Univ Oxford, Dept Zool, Math Ecol Res Grp, Oxford OX1 3PS, England
[2] Univ Oxford, Dept Zool, Peter Medawar Bldg Pathogen Res, Oxford, England
[3] Univ Oxford, NIHR Hlth Protect Res Unit Gastrointestinal Infec, Oxford, England
[4] Queen Mary Univ London, Sch Biol & Behav Sci, London, England