[1] Cent Publ Hlth Lab, Lab Enter Pathogens, Specialist & Reference Microbiol Div, Hlth Protect Agcy, London NW9 5HT, England
[2] Scottish Agr Coll, Vet Sci Div, Inverness IV2 4JZ, Scotland
[3] Univ Edinburgh, Ctr Trop & Vet Med, Roslin EH25 9RG, Midlothian, Scotland
[4] Univ London Imperial Coll Sci & Technol, Ctr Mol Microbiol & Immunol, London SW7 2AZ, England