[1] Univ Tokyo, Dept Gen Syst Studies, Meguro Ku, Tokyo 1538902, Japan
[2] Univ Queensland, Sch Biol Sci, Brisbane 4072, Australia
[3] Nagasaki Univ, Inst Trop Med, Nagasaki 8528523, Japan
[4] Res Inst Humanity & Nat, Kita ku, Kyoto 6038047, Japan
[5] Kyoto Univ, Ctr Ecol Res, Otsu, Shiga 5202113, Japan
[6] Natl Inst Genet, Ctr Frontier Res, Mishima, Shizuoka 4518540, Japan
[7] Univ Tokyo, Grad Sch Agr & Life Sci, Bunkyo Ku, Tokyo 1130032, Japan