[1] Univ Tubingen, Inst Geosci, D-72076 Tubingen, Germany
[2] European Inst Marine Studies, F-29280 Plouzane, France
[3] UFR Sci, Lab Bioindicateurs Actuels & Fossiles, F-49045 Angers, France
[4] Univ Southampton, Natl Oceanog Ctr, Southampton SO14 3ZH, Hants, England
[5] Royal Netherlands Inst Sea Res, Dept Marine Organ Biogeochem, NL-1790 AB Den Burg, Netherlands