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ABSTRACT With many species interacting in nature, determining which interactions describe community dynamics is nontrivial. By applying a computational modeling approach to an extensive field survey, we assessed the importance of interactions from plants (both inter‐ and intra‐specific), pollinators and insect herbivores on plant performance (i.e., viable seed production). We compared the inclusion of interaction effects as aggregate guild‐level terms versus terms specific to taxonomic groups. We found that a continuum from positive to negative interactions, containing mostly guild‐level effects and a few strong taxonomic‐specific effects, was sufficient to describe plant performance. While interactions with herbivores and intraspecific plants varied from weakly negative to weakly positive, heterospecific plants mainly promoted competition and pollinators facilitated plants. The consistency of these empirical findings over 3 years suggests that including the guild‐level effects and a few taxonomic‐specific groups rather than all pairwise and high‐order interactions, can be sufficient for accurately describing species variation in plant performance across natural communities. 

Abstract While most studies of species coexistence focus on the mechanisms that maintain coexistence, it is equally important to understand the mechanisms that structure failed coexistence. For example, California annual grasslands are heavily invaded ecosystems, where non‐native annuals have largely dominated and replaced native communities. These systems are also highly variable, with a high degree of rainfall seasonality and interannual rainfall variability—a quality implicated in the coexistence of functionally distinct species. Yet, despite the apparent strength of this variation, coexistence between native and non‐native annuals in this system has faltered.To test how variation‐dependent coexistence mechanisms modulate failed coexistence, we implemented a competition experiment between two previously common native forbs and three now‐dominant non‐native annual grasses spanning a conservative‐acquisitive range of traits. We grew individuals from each species under varying densities of all other species as competitors, under either wetter or drier early season rainfall treatments. Using subsequent seed production, we parameterized competition models, assessed the potential for coexistence among species pairs and quantified the relative influence of variation‐dependent coexistence mechanisms.As expected, we found little potential for coexistence. Competition was dominated by the non‐native grassAvena fatua, while native forbs were unable to invade non‐native grasses. Mutual competitive exclusion was common across almost all species and often contingent on rainfall, suggesting rainfall‐mediated priority effects. Among variation‐dependent mechanisms, the temporal storage effect had a moderate stabilizing effect for four of five species when averaged across competitors, while relative nonlinearity in competition was largely destabilizing, except for the most conservative non‐native grass, which benefited from a competitive release under dry conditions.Synthesis: Our findings suggest that rainfall variability does little to mitigate the fitness differences that underlie widespread annual grass invasion in California, but that it influences coexistence dynamics among the now‐dominant non‐native grasses. 

Abstract Declines in grassland diversity in response to nutrient addition are a general consequence of global change. This decline in species richness may be driven by multiple underlying processes operating at different time‐scales. Nutrient addition can reduce diversity by enhancing the rate of local extinction via competitive exclusion, or by reducing the rate of colonization by constraining the pool of species able to colonize under new conditions. Partitioning net change into extinction and colonization rates will better delineate the long‐term effect of global change in grasslands.We synthesized changes in richness in response to experimental fertilization with nitrogen, phosphorus and potassium with micronutrients across 30 grasslands. We quantified changes in local richness, colonization, and extinction over 8–10 years of nutrient addition, and compared these rates against control conditions to isolate the effect of nutrient addition from background dynamics.Total richness at steady state in the control plots was the sum of equal, relatively high rates of local colonization and extinction. On aggregate, 30%–35% of initial species were lost and the same proportion of new species were gained at least once over a decade. Absolute turnover increased with site‐level richness but was proportionately greater at lower‐richness sites relative to starting richness. Loss of total richness with nutrient addition, especially N in combination with P or K, was driven by enhanced rates of extinction with a smaller contribution from reduced colonization. Enhanced extinction and reduced colonization were disproportionately among native species, perennials, and forbs. Reduced colonization plateaued after the first few (<5) years after nutrient addition, while enhanced extinction continued throughout the first decade.Synthesis. Our results indicate a high rate of colonizations and extinctions underlying the richness of ambient communities and that nutrient enhancement drives overall declines in diversity primarily by exclusion of previously established species. Moreover, enhanced extinction continues over long time‐scales, suggesting continuous, long‐term community responses and a need for long‐term study to fully realize the extinction impact of increased nutrients on grassland composition.