[1] Univ Delaware, Coll Marine Studies, Lewes, DE 19958 USA
[2] Univ Calif Santa Barbara, Inst Marine Sci, Santa Barbara, CA 93106 USA
[3] Univ Calif Santa Barbara, Dept Ecol Evolut & Marine Biol, Santa Barbara, CA 93106 USA
[4] Univ Calif Santa Cruz, Inst Marine Sci, Santa Cruz, CA 95064 USA