[1] Univ Utrecht, Dept Earth Sci, NL-3584 CD Utrecht, Netherlands
[2] Royal Netherlands Inst Sea Res, Dept Marine Organ Biogeochem, NL-1790 AB Den Burg, Netherlands
[3] Royal Netherlands Inst Sea Res, Dept Biol Oceanog, NL-1790 AB Den Burg, Netherlands
[4] Wageningen Univ, Microbiol Lab, NL-6703 HB Wageningen, Netherlands