[1] Univ Laval, Quebec Ocean, Dept Biol, Quebec City, PQ G1V 0A6, Canada
[2] Univ Quebec, Inst Sci Mer Rimouski, Rimouski, PQ G5L 3A1, Canada
[3] Fisheries & Oceans Canada, Inst Ocean Sci, Sidney, BC V8L 4B2, Canada
[4] Fisheries & Oceans Canada, Inst Freshwater, Winnipeg, MB R3T 2N6, Canada
[5] Univ British Columbia, Dept Earth & Oceans Sci, Vancouver, BC V6T 1Z4, Canada