1. Department of Integrative Biology, University of California, Berkeley, Berkeley, CA 94720
2. Medical Research Council–University of Glasgow Centre for Virus Research, Glasgow G61 1QH, United Kingdom
3. Institute of Biodiversity, Animal Health and Comparative Medicine, College of Medical, Veterinary and Life Sciences, University of Glasgow, Glasgow G12 8QQ, United Kingdom
4. Centre for Ecology and Conservation, University of Exeter, Exeter TR10 9FE, United Kingdom
5. Department of Ecology and Evolution, University of Chicago, Chicago, IL 60637