1. Division of Evolution, Infection and Genomics, School of Biological Sciences, Faculty of Biology, Medicine and Health, The University of Manchester, Manchester, UK
2. Pembroke College, University of Cambridge, Cambridge, UK
3. Present address: MRC Centre for Molecular Bacteriology and Infection, Imperial College London, London, UK
4. Department of Earth and Environmental Sciences, School of Natural Sciences, Faculty of Science and Engineering, The University of Manchester, Manchester, UK