[1] Univ Cambridge, Dept Genet, Cambridge CB2 3EH, England
[2] Univ St Andrews, Sch Biol, St Andrews KY19 9ST, Fife, Scotland
[3] Univ Jyvaskyla, Dept Biol & Environm Sci, Jyvaskyla, Finland
[4] Univ Edinburgh, Ctr Immun Infect & Evolut, Edinburgh EH9 3JT, Midlothian, Scotland
[5] Univ Edinburgh, Inst Evolutionary Biol, Edinburgh EH9 3JT, Midlothian, Scotland