[1] Univ Liverpool, Inst Vet Sci, Leahurst Campus, Neston CH64 7TE, England
[2] Univ Liverpool, Dept Epidemiol & Populat Hlth, Inst Infect & Global Hlth, Leahurst Campus, Neston CH64 7TE, England
[3] NIHR Hlth Protect Res Unit Emerging & Zoonot Infe, Liverpool L69 7BE, Merseyside, England
[4] Royal Dick Sch Vet Studies, Easter Bush Campus, Edinburgh EH25 9RG, Midlothian, Scotland