[1] Univ Montana, W A Franke Coll Forestry & Conservat, Northern Rocky Mt Sci Ctr, Wildlife Biol Program, Missoula, MT USA
[2] Univ Arizona, Sch Nat Resources & Environm, Tucson, AZ USA
[3] US Fish & Wildlife Serv, Pacific Reg, Portland, OR USA
[4] Washington State Univ, Pullman, WA USA
[5] US Geol Survey, Ft Collins Sci Ctr, Ft Collins, CO USA
[6] US Geol Survey, Cooperat Res Unit, Tucson, AZ USA
[7] Univ Montpellier, EPHE PSL Univ, CEFE, CNRS,IRD, Montpellier, France
[8] Univ Montana, W A Franke Coll Forestry & Conservat, Northern Rocky Mt Sci Ctr, Wildlife Biol Program,US Geol Survey, Missoula, MT 59812 USA