[1] Univ Glasgow, Boyd Orr Ctr Populat & Ecosyst Hlth, Inst Biodivers Anim Hlth & Comparat Med, Coll Med Vet & Life Sci, Glasgow, Lanark, Scotland
[2] Albert Einstein Coll Med, Dept Syst & Computat Biol, Bronx, NY 10461 USA
[3] Univ Edinburgh, Sch Biol Sci, Inst Evolut, Ctr Immun Infect & Evolut, Edinburgh, Midlothian, Scotland
[4] Univ Edinburgh, Sch Biol Sci, Inst Immunol, Ctr Immun Infect & Evolut, Edinburgh, Midlothian, Scotland
[5] Univ Edinburgh, Sch Biol Sci, Inst Infect Res, Ctr Immun Infect & Evolut, Edinburgh, Midlothian, Scotland
[6] Burnet Inst, Melbourne, Vic, Australia
[7] Albert Einstein Coll Med, Dept Microbiol & Immunol, Bronx, NY 10461 USA