[1] Univ Edinburgh, Roslin Inst, Edinburgh, Midlothian, Scotland
[2] Univ Edinburgh, Royal Dick Sch Vet Studies, Edinburgh EH9 1QH, Midlothian, Scotland
[3] Univ Glasgow, Sch Vet Med, Coll Med Vet & Life Sci, Glasgow, Lanark, Scotland
[4] Inst Anim Hlth, Compton, Berks, England
[5] Fed Res Inst Anim Hlth, Friedrich Loeffler Inst, Inst Novel & Emerging Infect Dis, Riems Isl, Germany
[6] SNBTS, Natl Sci Lab, Edinburgh, Midlothian, Scotland