[1] Univ Corsica, Equipe Ecosyst Littoraux, FRES 3041, UMR CNRS SPE 6134, F-20250 Corte, France
[2] Aix Marseille Univ, F-13288 Marseille, France
[3] Toulon Univ, Mediterranean Inst Oceanog MIO, CNRS, IRD, F-13288 Marseille, France
[4] Spanish Council Sci Res, Ctr Adv Studies Blanes, Access Cala St Francesc, Blanes 17300, Spain
[5] Edith Cowan Univ, Ctr Marine Ecosyst Res, Perth, WA 6027, Australia