1. Montana Cooperative Fishery Research Unit, Department of Ecology, Montana State University, Bozeman, Montana, USA
2. U.S. Geological Survey, Montana Cooperative Fishery Research Unit, Department of Ecology, Montana State University, Bozeman, Montana, USA
3. Department of Interior, U.S. National Park Service, Yellowstone Center for Resources, Native Fish Conservation Program, Yellowstone National Park, Wyoming, USA