1. Department of Forest and Conservation Sciences, University of British Columbia, Vancouver, British Columbia, Canada V6T 1Z4
2. Department of Watershed Sciences and Ecology Center, Utah State University, Logan, UT 84322, USA
3. School of Biological and Chemical Sciences, Queen Mary University of London, London E1 4NS, UK
4. School of Biology and Ecology, University of Maine, Orono, ME 04469, USA
5. Section of Ecology, Behavior and Evolution, University of California-San Diego, La Jolla, CA 92093, USA