[1] Bangor Univ, Ctr Appl Marine Sci, Menai Bridge, Gwynedd, Wales
[2] Bangor Univ, Sch Ocean Sci, Menai Bridge, Gwynedd, Wales
[3] CSIRO Oceans & Atmosphere, Castray Esplanade Battery Point, Hobart, Tas, Australia
[4] Alfred Wegener Inst, Helmholz Ctr Polar & Marine Res, Biol Anstalt Helgoland, Helgoland, Germany