[1] Hokkaido Univ, Grad Sch Environm Sci, Kita Ku, Sapporo, Hokkaido 0600810, Japan
[2] Hokkaido Univ, Global Ctr Excellence Program Integrated Field En, Sapporo, Hokkaido 0600810, Japan
[3] Univ Nacl, Escuela Med Vet, Programa Invest Enfermedades Trop, Heredia, Costa Rica
[4] Nagasaki Univ, Inst Trop Med NEKKEN, Nagasaki, Japan
[5] Nagasaki Univ, Global Ctr Excellence Program Trop & Emergent Inf, Nagasaki, Japan