[1] Aix Marseille Univ, Univ Toulon, CNRS INSU, IRD,MIO,UM 110, Marseille, France
[2] Univ Brest, Lab Sci Environm Marin, IUEM, UMR CNRS UBO IRD Ifremer 6539, Plouzane, France
[3] Aix Marseille Univ, Univ Toulon, CNRS INSU, IRD,MIO,UM 110, New Caledonia, New Caledonia
[4] Univ Copenhagen, Marine Biol Sect, Dept Biol, Helsingor, Denmark
[5] Univ Massachusetts Dartmouth, Dept Biol, N Dartmouth, MA USA