[1] Univ Sheffield, Dept Anim & Plant Sci, Sheffield S10 2TN, S Yorkshire, England
[2] Univ Edinburgh, Ctr Immunol Infect & Evolut, Inst Evolutionary Biol, Ashworth Labs, Edinburgh EH9 3JT, Midlothian, Scotland
[3] Univ Edinburgh, Ctr Immunol Infect & Evolut, Inst Immunol, Ashworth Labs, Edinburgh EH9 3JT, Midlothian, Scotland
[4] Univ Edinburgh, Ctr Immunol Infect & Evolut, Inst Infect Res, Ashworth Labs, Edinburgh EH9 3JT, Midlothian, Scotland
[5] Univ Liverpool, Inst Integrat Biol, Liverpool L69 7ZB, Merseyside, England
[6] Univ Zurich, Inst Evolutionary Biol & Environm Studies, CH-8057 Zurich, Switzerland