[1] Univ Edinburgh, Inst Evolutionary Biol, Edinburgh, Midlothian, Scotland
[2] Penn State Univ, Ctr Infect Dis Dynam, University Pk, PA 16802 USA
[3] Univ London Imperial Coll Sci Technol & Med, Dept Infect Dis Epidemiol, London, England
[4] Leiden Univ, Med Ctr, Leiden Malaria Res Grp, NL-2300 RA Leiden, Netherlands
[5] Univ Edinburgh, Ctr Immun Infect & Evolut, Edinburgh, Midlothian, Scotland