Nitrogen addition, not heterogeneity, alters the relationship between invasion and native decline in California grasslands.

The presence of invasive species reduces the growth and performance of native species; however, the linear or non-linear relationships between invasive abundance and native population declines are less often studied. We examine how the amount and spatial distribution of experimental N deposition influences the relationship between non-native, invasive annual grass abundance (Bromus hordeaceus and Bromus diandrus) and a dominant, native perennial grass species (Stipa pulchra) in California. We hypothesized that native populations would decline as invasion increased, and that high nitrogen availability would cause native species to decline at lower invasion levels. We predicted that the rate of population decline would be slower in heterogeneous, compared to homogeneous, environments. We employed a field experiment that manipulated the amount and spatial heterogeneity of N addition across a range of invasive/native-dominated communities. There were strong negative and non-linear associations between level of invasion and S. pulchra proportional change (PC). Stipa pulchra PC was more negative and seedling survival was lower when N was added, and the negative effects of N addition on PC became larger in the final year of the study when S. pulchra had the largest declines. There was not strong evidence showing reduced competition in heterogeneous, compared to homogeneous, N treatments. Soil moisture was similar between S. pulchra and B. hordeaceus plots under ambient N, but B. hordeaceus under added N reduced soil moisture. Under N addition, Bromus spp. take up N earlier, reduce soil moisture, and create dry conditions in which S. pulchra declines.