[1] Univ Liverpool, Inst Infect Vet & Ecol Sci, Liverpool, England
[2] Univ British Columbia, Dept Zool, Vancouver, BC, Canada
[3] Univ Calif Irvine, Dept Ecol & Evolutionary Biol, Irvine, CA 92697 USA
[4] Univ Calif Irvine, Ctr Complex Biol Syst, Irvine, CA 92697 USA