1. Department of Evolution, Ecology and Behaviour, Institute of Infection, Veterinary and Ecological Sciences, University of Liverpool, Crown Street, Liverpool, L69 7ZB, UK
2. College of Life and Environmental Science, University of Exeter, Penryn, Cornwall, TR10 9FE, UK
3. Department of Microbiology, Moyne Institute for Preventive Medicine, School of Genetics and Microbiology, Trinity College Dublin, Dublin 2, Republic of Ireland