1. Department of Zoology, University of Manitoba, Winnipeg MB, R3T 2N2, Canada, Univeristy of Victoria/Freshwater Institute/Fisheries and Oceans Canada, 501 University Cres., Winnipeg MB, R3T 2N6, Canada, and Department of Environment and Geography, University of Manitoba, 500 University Cres., Winnipeg MB, R3T 2N2, Canada