1. Department of Computer Science, University of Oxford, Oxford OX1 3QD, UK
2. Mathematical Institute, University of Oxford, Oxford OX2 6GG, UK
3. Roche Pharmaceutical Research and Early Development, Pharmaceutical Sciences, Roche Innovation Center Basel, Basel CH-4070, Switzerland
4. Mathematics Institute, University of Warwick, Coventry CV4 7AL, UK
5. Zeeman Institute for Systems Biology and Infectious Disease Epidemiology Research, University of Warwick, Coventry CV4 7AL, UK