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Abstract Biodiversity generally enhances ecosystem productivity, but whether such effects persist or intensify during climate extremes is unclear. Here we synthesized data from 75 biodiversity experiments across grasslands and forests to assess how aridity and soil nutrients modulate plant diversity effects under drought and heat extremes. Biodiversity most strongly enhanced productivity under extreme drought in more arid grasslands but had limited effects in forests under such conditions. More arid grasslands showed enhanced complementarity effects under extreme drought, whereas less arid grasslands favoured selection effects driven by productive species. Heat extremes did not produce comparable context-dependent changes in plant diversity effects across ecosystem types or aridity gradients. Soil nutrients did not have any detectable influence under either drought or heat extremes, suggesting that as climatic stress intensifies, hydric limitations override edaphic constraints. Our synthesis identifies when and where plant diversity most strongly enhances productivity, showing that its effects are the greatest under extreme drought in more arid grasslands. 

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. 

Multitrophic interactions can strongly influence the structure and functioning of ecosystems, but how plant diversity influences the direction and predictability of multitrophic interactions across agricultural and natural ecosystems remains unclear. Using 149 field studies across five continents, we found that, on average, increasing plant diversity tended to exert differential top-down and bottom-up effects in croplands versus grasslands and forests. Organic and nonorganic croplands exhibited 846 and 148% higher invertebrate natural enemy-to-herbivore abundance ratios under increased plant diversity, consistent with top-down control patterns where predator gains cause herbivore declines, enhancing crop outcomes. In grasslands and forests, increasing plant diversity was associated with bottom-up effects where enhanced productivity increased both herbivore and predator populations, with the enemy-to-herbivore ratio increasing 4.73% for grasslands and 21.2% for forests. Our findings suggest that biodiversity effects on productivity are not solely explained by direct plant-plant interactions and the resulting biodiversity-productivity relationship. Rather, they reveal patterns consistent with the framework of top-down and bottom-up effects, the relative balance of which may vary depending on ecosystem and management type. The magnitude of the effects of diversified farming on crop pests suggests that crop diversification may be an important avenue for managing crop pests preventatively and thereby enhancing agricultural sustainability. 

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.