[1] Univ Edinburgh, Ctr Infect Dis, Edinburgh EH9 3JT, Midlothian, Scotland
[2] Penn State Univ, Ctr Infect Dis Dynam, Dept Biol, Mueller Lab 208, University Pk, PA 16802 USA
[3] Penn State Univ, Ctr Infect Dis Dynam, Dept Entomol, Mueller Lab 208, University Pk, PA 16802 USA
[4] Univ New England, Ctr Anim Hlth & Welf, Sch Environm & Rural Sci, Armidale, NSW 2351, Australia
[5] Yale Univ, Sch Med, Dept Epidemiol & Publ Hlth, New Haven, CT 06510 USA