[1] Univ Calif Santa Barbara, Inst Marine Sci, Coastal Res Ctr, Santa Barbara, CA 93106 USA
[2] Univ Calif Santa Barbara, Dept Ecol Evolut & Marine Biol, Santa Barbara, CA 93106 USA
[3] Calif State Univ Northridge, Dept Biol, Northridge, CA 91330 USA
[4] Univ Calif Santa Cruz, Dept Ecol & Evolutionary Biol, Santa Cruz, CA 95064 USA