[1] Univ Ryukyus, Dept Chem Biol & Marine Sci, Okinawa 9030213, Japan
[2] Univ Ryukyus, Sesoko Stn, Trop Biosphere Res Ctr, Okinawa 9070227, Japan
[3] Univ Tokyo, Grad Sch Arts & Sci, Dept Life Sci, Meguro Ku, Tokyo 1538902, Japan
[4] James Cook Univ N Queensland, Comparat Genom Ctr, Townsville, Qld 4811, Australia