[1] Univ Hawaii Manoa, Dept Oceanog, Daniel K Inouye Ctr Microbial Oceanog Res & Educ, Honolulu, HI 96822 USA
[2] Univ Hawaii Manoa, Sch Ocean & Earth Sci & Technol, Sea Grant Coll Program, Honolulu, HI 96822 USA
[3] Univ Calif San Diego, Scripps Inst Oceanog, Marine Biol Res Div, La Jolla, CA 92093 USA
[4] Royal Netherlands Inst Sea Res NIOZ, Dept Marine Microbiol & Biogeochem, Texel, Netherlands