1. Department of Biology, Colorado State University, Fort Collins, CO 80523-1878, USA
2. Department of Physics, Chemistry and Biology, Division of Theoretical Biology, Linköping University, Linköping, Sweden
3. The Zeeman Institute for Systems Biology and Infectious Disease Epidemiology Research, School of Life Sciences and Mathematics Institute, University of Warwick, Coventry, UK
4. USDA APHIS Veterinary Services, Center for Epidemiology and Animal Health, Fort Collins, CO, USA