[1] Pontificia Univ Catolica Chile, Fac Ciencias Biol, Subtidal Ecol Lab, Casilla 114-D, Santiago, Chile
[2] Pontificia Univ Catolica Chile, Fac Ciencias Biol, Ctr Marine Conservat, Estn Costera Invest Marinas, Casilla 114-D, Santiago, Chile
[3] Brown Univ, Dept Ecol & Evolutionary Biol, Providence, RI 02912 USA