[1] Univ S Carolina, Dept Biol Sci, Columbia, SC 29208 USA
[2] Univ S Carolina, Marine Sci Program, Columbia, SC 29208 USA
[3] Columbia Univ, Dept Earth & Environm Sci, New York, NY 10027 USA
[4] Columbia Univ, Lamont Doherty Earth Observ, New York, NY 10027 USA
[5] Texas A&M Univ, Dept Marine Sci & Oceanog, Galveston, TX 77551 USA