1. Institute of Ecology and Evolution, School of Biological Sciences, University of Edinburgh, Edinburgh EH9 3FL, UK
2. School of Biosciences, University of Sheffield, Sheffield S10 2TN, UK
3. Moredun Research Institute, Pentlands Science Park, Bush Loan, Penicuik EH26 0PZ, UK
4. Institute for Research in Immunology and Cancer, Université de Montréal, Montréal, Canada H3C 3J7