[1] Univ Calif Santa Cruz, Dept Environm Studies, Santa Cruz, CA 95064 USA
[2] Univ Calif Santa Cruz, Dept Ecol & Evolutionary Biol, Santa Cruz, CA 95064 USA
[3] San Diego State Univ, Dept Biol, San Diego, CA 92182 USA
[4] Univ Washington, Appl Phys Lab, Polar Sci Ctr, Seattle, WA 98105 USA
[5] Univ Alaska, Sch Fisheries & Ocean Sci, Fairbanks, AK 99701 USA