1. Montana Conservation Science Institute5200 Upper Miller Creek Road, Missoula, MT 59803, USA
2. Division of Biological Sciences, University of MontanaMissoula, MT 59812, USA
3. Department of Mathematics, University of MontanaMissoula, MT 59812, USA
4. Laboratoire d'Ecologie Alpine, Génomique des Populations et Biodiversité, CNRS UMR5553, Université Joseph FourierBP 53, 38041 Grenoble Cedex 9, France