1. School of Biosciences, University of Exeter, Geoffrey Pope Building, Stocker Road, Exeter EX4 4QD, UK
2. Defence Science and Technology Laboratory, Porton Down, Salisbury SP4 0JQ, UK
3. London School of Hygiene and Tropical Medicine, Department of Infectious and Tropical Diseases, Keppel Street, London WC1E 7HT, UK