Watershed, reach, and riparian influences on stream fish assemblages in the Northern Lakes and Forest Ecoregion, USA

We analyzed data from 79 watersheds in an undegraded U.S. ecoregion to identify key environmental factors that explained stream fish assemblage patterns and to evaluate the relative influence of environmental factors operating at different spatial scales. A few key factors from the watershed, reach, and riparian scale explained a significant amount of the variance in fish attributes. Three environment-fish associations were evident. Top carnivores and intolerant cold-water fishes were associated with relatively narrow, deep, high-gradient, cold streams with strong groundwater inputs. Tolerant cyprinids occurred in small streams with low groundwater input, low dissolved oxygen, and abundant macrophytes. A diverse assemblage (Cyprinidae, Catostomidae, Centrarchidae, Percidae) existed in warm, wide streams in watersheds dominated by lacustrine sand and gravel geology and abundant wetlands and lakes. Overall, reach-scale variables directly explained the most, watershed-scale variables less, and riparian-scale variables the least variation in fish attributes. Watershed and riparian variables had indirect connections with fishes through their direct influence on reach variables. In conjunction with findings from more degraded regions, we conclude that the relative influence of reach-scale variables on fishes are greatest in undegraded areas and that direct effects of watershed-scale variables are increasingly important as human modifications of the landscape increase.