An emergent multiple predator effect may enhance biotic resistance in a stream fish assemblage

While two cyprinid fishes introduced from nearby drainages have become widespread and abundant in the Eel River of northwestern California, a third nonindigenous cyprinid has remained largely confined to less than or equal to25 km of one major tributary. (the Van Duzen River) for at least 15 years. The downstream limit of this species, speckled dace, does not appear to correspond with any thresholds or steep gradients in abiotic conditions, but it lies near the upstream limits of three other fishes: coastrange sculpin, prickly sculpin, and nonindigenous Sacramento pikeminnow. We conducted a laboratory stream experiment to explore the potential for emergent multiple predator effects to influence biotic resistance in this situation. Sculpins in combination with Sacramento pikeminnow caused greater mortality of speckled dace than predicted based on their separate effects. In contrast to speckled dace, 99% of sculpin survived trials with Sacramento pikeminnow, in part because sculpin usually occupied benthic cover units while Sacramento pikeminnow occupied the water column. A 10-fold difference in benthic cover availability did not detectably influence biotic interactions in the experiment. The distribution of speckled dace in the Eel River drainage may be limited by two predator taxa with very different patterns of habitat use and a shortage of alternative habitats.