[1] Univ Edinburgh, Sch Biol Sci, Edinburgh EH9 3JT, Midlothian, Scotland
[2] Scottish Agr Coll, Penicuik EH26 0PH, Midlothian, Scotland
[3] Vet Labs Agcy Weybridge, Ctr Epidemiol & Risk Anal, Weybridge KT15 3NB, Surrey, England
[4] Univ Coll Dublin, Ctr Vet Epidemiol & Risk Anal, Sch Agr Food Sci & Vet Med, Dublin 4, Ireland
[5] Agr Hse, Dept Agr Fisheries & Food, Dublin 2, Ireland
[6] Roslin Bioctr, Roslin Inst, Roslin EH25 9PS, Midlothian, Scotland