[1] Univ Calif Santa Barbara, Dept Ecol Evolut & Marine Biol, Santa Barbara, CA 93106 USA
[2] Univ Calif Santa Barbara, Marine Sci Inst, Santa Barbara, CA 93106 USA
[3] Univ Calif Santa Barbara, Coll Creat Studies, Santa Barbara, CA 93106 USA
[4] Univ Calif Santa Barbara, Marine Sci Inst, US Geol Survey, Western Ecol Res Ctr, Santa Barbara, CA 93106 USA