1. Pacific Salmon Ecology and Conservation Laboratory, Department of Forest and Conservation Sciences, University of British Columbia, Vancouver, British Columbia, BC V6T 1Z4, Canada
2. Raincoast Conservation Foundation, Sidney, BC V8G 1P2, Canada
3. Department of Biology, University of Victoria, Victoria, BC V8W 2Y2, Canada
4. Pacific Biological Station, Fisheries and Oceans Canada, Nanaimo, BC V9T 6N7, Canada