[1] Univ Ryukyus, Grad Sch Engn & Sci, Mol Invertebrate Systemat & Ecol Lab, 1 Senbaru, Nishihara, Okinawa 9030123, Japan
[2] Naturalis Biodivers Ctr, Darwinweg 2, NL-2333 CR Leiden, Netherlands
[3] Univ Ryukyus, Trop Biosphere Res Ctr, 1 Senbaru, Nishihara, Okinawa 9030123, Japan