[1] Univ Bretagne Occidentale, CNRS, IFREMER, IRD,LOPS, Brest, France
[2] Council Sci & Ind Res Nat Resources & Env, Cape Town, South Africa
[3] Univ Cape Town, Ma Re Inst, Dept Oceanog, Cape Town, South Africa
[4] Univ Cape Town, Dept Oceanog, Nansen Tutu Ctr Marine Environm Res, Cape Town, South Africa