[1] Univ Wisconsin, Dept Geog, Platteville, WI 53818 USA
[2] Univ Idaho, Coll Nat Resources, Dept Fish & Wildlife Sci, Moscow, ID USA
[3] US Geol Survey, Patuxent Wildlife Res Ctr, Laurel, MD USA
[4] Univ Idaho, US Geol Survey, Idaho Cooperat Fish & Wildlife Res Unit, Dept Fish & Wildlife Sci,Coll Nat Resources, Moscow, ID USA