[1] Univ Concepcion, Dept Oceanog, Concepcion, Chile
[2] Univ Concepcion, Ctr Invest Oceanog Pacifico Sudoriental COPAS, Concepcion, Chile
[3] Univ Concepcion, Dept Oceanog, Programa Postgrad Oceanog, Concepcion, Chile
[4] Univ Paris 06, Roscoff, France
[5] CNRS, Biol Stn, UMR 7144, Roscoff, France