[1] Fisheries & Oceans Canada, Pacific Biol Stn, Nanaimo, BC V9T 6N7, Canada
[2] Fisheries & Oceans Canada, Bedford Inst Oceanog, Dartmouth, NS B2Y 4A2, Canada
[3] Univ Guelph, Dept Integrat Biol, Guelph, ON N1G 2W1, Canada
[4] Fisheries & Oceans Canada, Inst Ocean Sci, Sidney, BC V8L 5T5, Canada
[5] Univ British Columbia, Dept Forest & Conservat Sci, Vancouver, BC V6T 1Z4, Canada