[1] Virginia Polytech Inst & State Univ, Dept Fisheries & Wildlife Sci, Blacksburg, VA 24061 USA
[2] Montana State Univ, Montana Cooperat Fishery Res Unit, Dept Ecol, Bozeman, MT 59717 USA
[3] Univ Georgia, Savannah River Ecol Lab, Aiken, SC 29802 USA