1. Department of Biology, Stanford University, Stanford, CA 94305, USA
2. Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ 08544, USA
3. US Geological Survey, Southwest Biological Science Center, Canyonlands Research Station, Moab, UT 84532, USA
4. Department of Ecosystem and Conservation Sciences, University of Montana, Missoula, MT 59812, USA