[1] Kyoto Univ, Grad Sch Global Environm Studies, Sakyo Ku, Yoshida Honmachi, Kyoto 6068501, Japan
[2] Kyoto Univ, Grad Sch Human & Environm Studies, Sakyo Ku, Yoshida Nihonmatsu Cho, Kyoto 6068501, Japan
[3] Natl Res & Innovat Agcy Indonesia, Res Ctr Biol, Museum Zool Bogoriense, Gd Widyasatwaloka,Jl Raya Jakarta Bogor Km 46, Cibinong, West Java, Indonesia
[4] Univ Texas Arlington, Amphibian & Reptile Divers Res Ctr, Arlington, TX 76010 USA
[5] Univ Texas Arlington, Dept Biol, Arlington, TX 76010 USA