The Cosumnes River is the largest stream without a major dam on its mainstem in the Sacramento–San Joaquin drainage, central California, U.S.A. We studied its fishes over a 3-year period to answer the following questions: (1) Was the native fish fauna still present? (2) Why were alien fishes so abundant in a river system with a 'natural' flow regime, which elsewhere has been shown to favor native fishes? (3) Were there assemblages of fishes that reflected environmental differences created by the underlying geology? (4) Were there features of the watershed that consistently favored native fishes or that could be managed to favor native fishes? Of the 25 species collected, 17 were alien species; 14 species (five native) were abundant or widely distributed enough to use in detailed analyses. Of the native species, only rainbow trout, Oncorhynchus mykiss, still occupied much of its native range in headwater streams. Other native species have been extirpated or persisted mainly above barriers to alien invasions. The most widely distributed alien species was redeye bass, Micropterus coosae, previously unknown from the river, whose abundance was associated with low-numbers of native species. Other aliens were found primarily in low-land habitats on the valley floor or foothills. Canonical Correspondence Analysis indicated that both native and alien species located on environmental gradients determined largely by elevation, temperature, flow, and emergent vegetation, but the associations with these variables were not strong. While most alien fishes were found in lowland sections of river flowing through agricultural regions, the general relationships between species abundance and landscape-level variables were weak. Assemblages of fishes were poorly defined mixtures of native and alien species. The strikingly distinct geological regions of the basin no longer supported distinct fish assemblages. Species distributions were highly individualistic, reflecting dynamic patterns of introductions, invasions, and local extinctions, as well as physiological tolerances and life history patterns. Most native fishes are likely to persist in the Cosumnes River only if summer flows are increased and if populations above natural barriers are protected from further invasions by alien species, especially redeye bass. General conclusions from this study include: (1) altered habitats can support native species under some circumstances; (2) new fish assemblages with characteristics of 'natural' communities are likely to develop in invaded systems; (3) restoring flow regimes to favor native fishes may require restoring minimum summer flows as well as high channel-forming flows. However, reversing or even reducing, the impact of the predatory redeye bass, pre-adapted for California streams, is probably not possible. [ABSTRACT FROM AUTHOR]