[1] Univ Hawaii Manoa, Hawaii Inst Marine Biol, Kaneohe, HI 96744 USA
[2] Univ Hawaii Manoa, Dept Zool, Honolulu, HI 96822 USA
[3] San Francisco State Univ, Dept Biol, Tiburon, CA 94920 USA
[4] San Francisco State Univ, Romberg Tiburon Ctr Environm Studies, Tiburon, CA 94920 USA
[5] US Geol Survey, Hawaii Cooperat Fishery Res Unit, Honolulu, HI 96822 USA