1. Department of Ecology and Evolution, State University of New York–Stony Brook, Stony Brook, NY 11794‐5245, USA;
2. Present address: Department of Soil, Water & Climate, University of Minnesota, St Paul, MN 55108, USA
3. Department of Ecology and Evolutionary Biology and Department of Earth System Science, University of California, Irvine, Irvine, CA 92697, USA;