[1] Univ Leeds, Fac Biol Sci, Sch Biol, Leeds LS2 9JT, England
[2] Univ Ryukyus, Grad Sch Engn & Sci, Mol Invertebrate Systemat & Ecol Lab, 1 Senbaru, Nishihara, Okinawa 9030213, Japan
[3] Univ Ryukyus, Trop Biosphere Res Ctr, 1 Senbaru, Nishihara, Okinawa 9030213, Japan
[4] Natl Inst Water & Atmospher Res NIWA, Hamilton, New Zealand
[5] Natl Univ Singapore, Dept Biol Sci, Expt Marine Ecol Lab, 14 Sci Dr 4, Singapore 117558, Singapore
[6] Univ Queensland, Ctr Biodivers & Conservat Sci, Sch Biol Sci, Brisbane, Qld 4072, Australia