[1] Pontificia Univ Catolica Chile, Fac Ciencias Biol, Dept Ecol, Santiago, Chile
[2] Pontificia Univ Catolica Chile, Fac Ciencias Biol, Ctr Adv Studies Ecol & Biodivers, Santiago, Chile
[3] Woods Hole Oceanog Inst, Dept Chem, Woods Hole, MA 02543 USA