1. Ecology, Evolution and Conservation Biology graduate program, Department of Biology, University of Nevada, Reno, Reno NV 89557, USA
2. Department of Integrative Biology, University of California, Berkeley, Berkeley CA 94720, USA
3. Department of Geography, University of Nevada, Reno, Reno NV 89557, USA
4. Institute of Earth Surface Dynamics, University of Lausanne, Lausanne 1015, Switzerland
5. Department of Natural Resources and Environmental Science, University of Nevada, Reno, Reno NV 89557, USA