About the Authors

Martin R. Miller

Affiliation Centre for Infectious Diseases, Institute of Immunology and Infection Research, University of Edinburgh, Edinburgh, United Kingdom

Lars Råberg

Affiliation Department of Animal Ecology, Lund University, Lund, Sweden

Andrew F. Read

Affiliation Center for Infectious Disease Dynamics and Departments of Biology and Entomology, Pennsylvania State University, University Park, Pennsylvania, United States of America

Nicholas J. Savill

nick.savill@ed.ac.uk

Affiliation Centre for Infectious Diseases, Institute of Immunology and Infection Research, University of Edinburgh, Edinburgh, United Kingdom

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Conceived and designed the experiments: LR AFR. Performed the experiments: LR. Analyzed the data: MRM NJS. Contributed reagents/materials/analysis tools: LR. Wrote the paper: MRM LR AFR NJS. Conceived and designed the mathematical models: NJS MRM. Conceived and designed the statistical methods: NJS.