[1] Univ Victoria, Sch Earth & Ocean Sci, Bob Wright Ctr A405, Victoria, BC V8W 3V6, Canada
[2] Fisheries & Oceans Canada, Inst Freshwater, Winnipeg, MB R3T 2N6, Canada
[3] Univ Manitoba, Dept Geog & Environm, Winnipeg, MB R3T 2N2, Canada
[4] Mem Univ Newfoundland, Ctr Ocean Sci, St John, NF A1C 5S7, Canada
[5] Univ Laval, Dept Biol, Quebec City, PQ G1K 7P4, Canada