[1] Univ Utrecht, Fac Geosci, Dept Earth Sci, NL-3584 CD Utrecht, Netherlands
[2] Helmholtz Ctr Polar & Marine Res, Alfred Wegener Inst, Marine Biogeosci, D-27570 Bremerhaven, Germany
[3] Netherlands Inst Ecol NIOO KNAW, Dept Aquat Ecol, NL-6708 PB Wageningen, Netherlands
[4] Royal Netherlands Inst Sea Res NIOZ, NL-1797 SZ T Horntje, Texel, Netherlands