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An important goal of invasion biology is to identify physical and environmental characteristics that may make a region particularly receptive to invasions. The inland waters of California (USA) are highly invaded, particularly by fishes, although there is wide variation in numbers of nonnative fishes across the state's watersheds. Here we examine patterns of fish invasions in California watersheds and their associations with natural environmental characteristics, native fish diversity, and various measures of human habitat disturbance. Our analysis is based on an extensive data set on the distribution of freshwater fishes across California's watersheds and on GIS land‐use coverages for the entire state. We used canonical correspondence analysis to examine associations between environmental characteristics and the distributions of both native and nonnative fish species. We then employed an information–theoretic model‐selection approach to rank hypothesized models derived from the fish invasion literature with regard to how well they predicted numbers of nonnative fishes in California watersheds. Our results indicate that pervasive, anthropogenic, landscape‐level changes (particularly the extent of urban development, small‐scale water diversions, aqueducts, and agriculture) influenced spatial patterns of invasion. In addition, we find that deliberately stocked fishes have different habitat associations, including a strong association with the presence of dams, than other introduced fishes. In our analysis, watersheds with the most native species also contain the most nonnative species. We find no evidence that existing watershed protection helps to prevent fish invasions in California, but we suggest that restoration of natural hydrologic processes may reduce invasion impacts.

Many of California's native populations of freshwater fish are in serious decline, as are freshwater faunas worldwide. Habitat loss and alteration, hydrologic modification, water pollution, and invasions have been identified as major drivers of these losses. Because these potential causes of decline are frequently correlated, it is difficult to separate direct from indirect effects of each factor and to appropriately rank their importance for conservation action. Recently a few authors have questioned the conservation significance of invasions, suggesting that they are "passengers" rather than "drivers" of ecological change. We compiled an extensive, watershed-level data set of fish presence and conservation status, land uses, and hydrologic modifications in California and used an information theoretic approach (Akaike's information criterion, AIC) and path analysis to evaluate competing models of native fish declines. Hydrologic modification (impoundments and diversions), invasions, and proportion of developed land were all predictive of the number of extinct and at-risk native fishes in California watersheds in the AIC analysis. Although nonindigenous fish richness was the best single predictor (after native richness) of fishes of conservation concern, the combined ranking of models containing hydrologic modification variables was slightly higher than that of models containing nonindigenous richness. Nevertheless, the path analysis indicated that the effects of both hydrologic modification and development on fishes of conservation concern were largely indirect, through their positive effects on nonindigenous fish richness. The best-fitting path model was the driver model, which included no direct effects of abiotic disturbance on native fish declines. Our results suggest that, for California freshwater fishes, invasions are the primary direct driver of extinctions and population declines, whereas the most damaging effect of habitat alteration is the tendency of altered habitats to support nonindigenous fishes.

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