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Abstract Forbs (“wildflowers”) are important contributors to grassland biodiversity but are vulnerable to environmental changes. In a factorial experiment at 94 sites on 6 continents, we test the global generality of several broad predictions: (1) Forb cover and richness decline under nutrient enrichment, particularly nitrogen enrichment. (2) Forb cover and richness increase under herbivory by large mammals. (3) Forb richness and cover are less affected by nutrient enrichment and herbivory in more arid climates, because water limitation reduces the impacts of competition with grasses. (4) Forb families will respond differently to nutrient enrichment and mammalian herbivory due to differences in nutrient requirements. We find strong evidence for the first, partial support for the second, no support for the third, and support for the fourth prediction. Our results underscore that anthropogenic nitrogen addition is a major threat to grassland forbs, but grazing under high herbivore intensity can offset these nutrient effects. 

Abstract Nutrient enrichment impacts grassland plant diversity such as species richness, functional trait composition and diversity, but whether and how these changes affect ecosystem stability in the face of increasing climate extremes remains largely unknown.We quantified the direct and diversity‐mediated effects of nutrient addition (by nitrogen, phosphorus, and potassium) on the stability of above‐ground biomass production in 10 long‐term grassland experimental sites. We measured five facets of stability as the temporal invariability, resistance during and recovery after extreme dry and wet growing seasons.Leaf traits (leaf carbon, nitrogen, phosphorus, potassium, and specific leaf area) were measured under ambient and nutrient addition conditions in the field and were used to construct the leaf economic spectrum (LES). We calculated functional trait composition and diversity of LES and of single leaf traits. We quantified the contribution of intraspecific trait shifts and species replacement to change in functional trait composition as responses to nutrient addition and its implications for ecosystem stability.Nutrient addition decreased functional trait diversity and drove grassland communities to the faster end of the LES primarily through intraspecific trait shifts, suggesting that intraspecific trait shifts should be included for accurately predicting ecosystem stability. Moreover, the change in functional trait diversity of the LES in turn influenced different facets of stability. That said, these diversity‐mediated effects were overall weak and/or overwhelmed by the direct effects of nutrient addition on stability. As a result, nutrient addition did not strongly impact any of the stability facets. These results were generally consistent using individual leaf traits but the dominant pathways differed. Importantly, major influencing pathways differed using average trait values extracted from global trait databases (e.g. TRY).Synthesis. Investigating changes in multiple facets of plant diversity and their impacts on multidimensional stability under global changes such as nutrient enrichment can improve our understanding of the processes and mechanisms maintaining ecosystem stability. 

Experiments comparing diploids with polyploids and in single grassland sites show that nitrogen and/or phosphorus availability influences plant growth and community composition dependent on genome size; specifically, plants with larger genomes grow faster under nutrient enrichments relative to those with smaller genomes. However, it is unknown if these effects are specific to particular site localities with speciifc plant assemblages, climates, and historical contingencies. To determine the generality of genome size-dependent growth responses to nitrogen and phosphorus fertilization, we combined genome size and species abundance data from 27 coordinated grassland nutrient addition experiments in the Nutrient Network that occur in the Northern Hemisphere across a range of climates and grassland communities. We found that after nitrogen treatment, species with larger genomes generally increased more in cover compared to those with smaller genomes, potentially due to a release from nutrient limitation. Responses were strongest for C₃grasses and in less seasonal, low precipitation environments, indicating that genome size effects on water-use-efficiency modulates genome size–nutrient interactions. Cumulatively, the data suggest that genome size is informative and improves predictions of species’ success in grassland communities. 

Abstract Nutrient availability and grazing are known as main drivers of grassland plant diversity, and increased nutrient availability and long‐term cessation of grazing often decrease local‐scale plant diversity. Experimental tests of mechanisms determining plant diversity focus mainly on vascular plants (VP), whereas non‐vascular plants (NVP, here bryophytes) have been ignored. It is therefore not known how the current models based on VPs predict the rates of total (NVP + VP) losses in plant diversity.Here we used plant community data, including VPs and NVPs, from nine sites in Europe and North America and belonging to the Nutrient Network experiment, to test whether neglecting NVPs leads to biased estimates of plant diversity loss rates. The plant communities were subjected to factorial addition of nitrogen (N), phosphorus (P), potassium with micronutrients (K_+μ), as well as a grazing exclusion combined with multi‐nutrient fertilization (NPK_+μ) treatment.We found that nutrient additions reduced both NVP and VP species richness, but the effects on NVP species richness were on average stronger than on VPs: NVP species richness decreased 67%, while VP species richness decreased 28%, causing their combined richness to decrease 38% in response to multi‐nutrient (NPK_+μ) fertilization. Thus, VP diversity alone underestimated total plant diversity loss by 10 percentage points.Although NVP and VP species diversities similarly declined in response to N and NPK_+μfertilizations, the evenness of NVPs increased and that of VPs remained unchanged. NP, NPK_+μfertilization and NPK_+μfertilization combined with grazing exclusion, associated with decreasing light availability at ground level, led to the strongest loss of NVP species or probability of NVP presence. However, grazing did not generally mitigate the fertilization effects.Synthesis. In nine grassland sites in Europe and North America, nutrient addition caused a larger relative decline in non‐vascular plant (NVP) than vascular plant species richness. Hence, not accounting for NVPs can lead to underestimation of losses in plant diversity in response to continued nutrient pollution of grasslands. 

Abstract Eutrophication usually impacts grassland biodiversity, community composition, and biomass production, but its impact on the stability of these community aspects is unclear. One challenge is that stability has many facets that can be tightly correlated (low dimensionality) or highly disparate (high dimensionality). Using standardized experiments in 55 grassland sites from a globally distributed experiment (NutNet), we quantify the effects of nutrient addition on five facets of stability (temporal invariability, resistance during dry and wet growing seasons, recovery after dry and wet growing seasons), measured on three community aspects (aboveground biomass, community composition, and species richness). Nutrient addition reduces the temporal invariability and resistance of species richness and community composition during dry and wet growing seasons, but does not affect those of biomass. Different stability measures are largely uncorrelated under both ambient and eutrophic conditions, indicating consistently high dimensionality. Harnessing the dimensionality of ecological stability provides insights for predicting grassland responses to global environmental change. 

Abstract Plant diversity effects on community productivity often increase over time. Whether the strengthening of diversity effects is caused by temporal shifts in species-level overyielding (i.e., higher species-level productivity in diverse communities compared with monocultures) remains unclear. Here, using data from 65 grassland and forest biodiversity experiments, we show that the temporal strength of diversity effects at the community scale is underpinned by temporal changes in the species that yield. These temporal trends of species-level overyielding are shaped by plant ecological strategies, which can be quantitatively delimited by functional traits. In grasslands, the temporal strengthening of biodiversity effects on community productivity was associated with increasing biomass overyielding of resource-conservative species increasing over time, and with overyielding of species characterized by fast resource acquisition either decreasing or increasing. In forests, temporal trends in species overyielding differ when considering above- versus belowground resource acquisition strategies. Overyielding in stem growth decreased for species with high light capture capacity but increased for those with high soil resource acquisition capacity. Our results imply that a diversity of species with different, and potentially complementary, ecological strategies is beneficial for maintaining community productivity over time in both grassland and forest ecosystems. 

Abstract Dominance often indicates one or a few species being best suited for resource capture and retention in a given environment. Press perturbations that change availability of limiting resources can restructure competitive hierarchies, allowing new species to capture or retain resources and leaving once dominant species fated to decline. However, dominant species may maintain high abundances even when their new environments no longer favour them due to stochastic processes associated with their high abundance, impeding deterministic processes that would otherwise diminish them.Here, we quantify the persistence of dominance by tracking the rate of decline in dominant species at 90 globally distributed grassland sites under experimentally elevated soil nutrient supply and reduced vertebrate consumer pressure.We found that chronic experimental nutrient addition and vertebrate exclusion caused certain subsets of species to lose dominance more quickly than in control plots. In control plots, perennial species and species with high initial cover maintained dominance for longer than annual species and those with low initial cover respectively. In fertilized plots, species with high initial cover maintained dominance at similar rates to control plots, while those with lower initial cover lost dominance even faster than similar species in controls. High initial cover increased the estimated time to dominance loss more strongly in plots with vertebrate exclosures than in controls. Vertebrate exclosures caused a slight decrease in the persistence of dominance for perennials, while fertilization brought perennials' rate of dominance loss in line with those of annuals. Annual species lost dominance at similar rates regardless of treatments.Synthesis.Collectively, these results point to a strong role of a species' historical abundance in maintaining dominance following environmental perturbations. Because dominant species play an outsized role in driving ecosystem processes, their ability to remain dominant—regardless of environmental conditions—is critical to anticipating expected rates of change in the structure and function of grasslands. Species that maintain dominance while no longer competitively favoured following press perturbations due to their historical abundances may result in community compositions that do not maximize resource capture, a key process of system responses to global change.