[1] Univ British Columbia, Dept Zool, Vancouver, BC V6T 1Z4, Canada
[2] Univ British Columbia, Biodivers Res Ctr, Vancouver, BC V6T 1Z4, Canada
[3] Bedford Inst Oceanog, Populat Ecol Div, Dept Fisheries & Oceans, Dartmouth, NS B2Y 4A2, Canada
[4] Univ British Columbia, Dept Bot, Vancouver, BC V6T 1Z4, Canada