[1] Univ Calif San Diego, Scripps Inst Oceanog, San Diego, CA 92103 USA
[2] St John Plant Sci Lab, Dept Trop Plant & Soil Sci, Honolulu, HI USA
[3] Univ Hawaii Manoa, Dept Oceanog, Daniel K Inouye Ctr Microbial Oceanog Res & Educ, Honolulu, HI 96822 USA
[4] Univ Hawaii Manoa, Sea Grant Coll Program, Honolulu, HI 96822 USA