[1] Univ Hawaii Manoa, Hawaii Inst Marine Biol, Kaneohe, HI 96744 USA
[2] Calif State Univ Northridge, Dept Biol, Northridge, CA 91330 USA
[3] Univ Calif Santa Cruz, Dept Ecol & Evolutionary Biol, Santa Cruz, CA 95064 USA
[4] Univ Hawaii Manoa, Uehiro Ctr Advancement Oceanog, Dept Oceanog, Honolulu, HI 96822 USA