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Ecosystems are experiencing changing global patterns of mean annual precipitation (MAP) and enrichment with multiple nutrients that potentially colimit plant biomass production. In grasslands, mean aboveground plant biomass is closely related to MAP, but how this relationship changes after enrichment with multiple nutrients remains unclear. We hypothesized the global biomass–MAP relationship becomes steeper with an increasing number of added nutrients, with increases in steepness corresponding to the form of interaction among added nutrients and with increased mediation by changes in plant community diversity. We measured aboveground plant biomass production and species diversity in 71 grasslands on six continents representing the global span of grassland MAP, diversity, management, and soils. We fertilized all sites with nitrogen, phosphorus, and potassium with micronutrients in all combinations to identify which nutrients limited biomass at each site. As hypothesized, fertilizing with one, two, or three nutrients progressively steepened the global biomass–MAP relationship. The magnitude of the increase in steepness corresponded to whether sites were not limited by nitrogen or phosphorus, were limited by either one, or were colimited by both in additive, or synergistic forms. Unexpectedly, we found only weak evidence for mediation of biomass–MAP relationships by plant community diversity because relationships of species richness, evenness, and beta diversity to MAP and to biomass were weak or opposing. Site-level properties including baseline biomass production, soils, and management explained little variation in biomass–MAP relationships. These findings reveal multiple nutrient colimitation as a defining feature of the global grassland biomass–MAP relationship. 

Abstract The Arctic is warming four times faster than the global average¹and plant communities are responding through shifts in species abundance, composition and distribution^2–4. However, the direction and magnitude of local changes in plant diversity in the Arctic have not been quantified. Using a compilation of 42,234 records of 490 vascular plant species from 2,174 plots across the Arctic, here we quantified temporal changes in species richness and composition through repeat surveys between 1981 and 2022. We also identified the geographical, climatic and biotic drivers behind these changes. We found greater species richness at lower latitudes and warmer sites, but no indication that, on average, species richness had changed directionally over time. However, species turnover was widespread, with 59% of plots gaining and/or losing species. Proportions of species gains and losses were greater where temperatures had increased the most. Shrub expansion, particularly of erect shrubs, was associated with greater species losses and decreasing species richness. Despite changes in plant composition, Arctic plant communities did not become more similar to each other, suggesting no biotic homogenization so far. Overall, Arctic plant communities changed in richness and composition in different directions, with temperature and plant–plant interactions emerging as the main drivers of change. Our findings demonstrate how climate and biotic drivers can act in concert to alter plant composition, which could precede future biodiversity changes that are likely to affect ecosystem function, wildlife habitats and the livelihoods of Arctic peoples^5,6. 

Vegetation change of the Arctic tundra due to global warming is a well-known process, but the implication for the belowground microbial communities, key in nutrient cycling and decomposition, is poorly understood. We characterized the fungal and bacterial abundances in litter and soil layers across 16 warming experimental sites at 12 circumpolar locations. We investigated the relationship between microbial abundances and nitrogen (N) and carbon (C) isotopic signatures, indicating shifts in microbial processes with warming. Microbial abundances were 2–3 orders of magnitude larger in litter than in soil. Local, site-dependent responses of microbial abundances were variable, and no general effect of warming was detected. The only generalizable trend across sites was a dependence between the warming response ratios and C:N ratio in controls, highlighting a legacy of the vegetation on the microbial response to warming. We detected a positive effect of warming on the litter mass and δ 15 N, which was linked to bacterial abundance under warmed conditions. This effect was stronger in experimental sites dominated by deciduous shrubs, suggesting an altered bacterial N-cycling with increased temperatures, mediated by the vegetation, and with possible consequences on ecosystem feedbacks to climate change. 

The relative contribution of bryophytes to plant diversity, primary productivity, and ecosystem functioning increases towards colder climates. Bryophytes respond to environmental changes at the species level, but because bryophyte species are relatively difficult to identify, they are often lumped into one functional group. Consequently, bryophyte function remains poorly resolved. Here, we explore how higher resolution of bryophyte functional diversity can be encouraged and implemented in tundra ecological studies. We briefly review previous bryophyte functional classifications and the roles of bryophytes in tundra ecosystems and their susceptibility to environmental change. Based on shoot morphology and colony organization, we then propose twelve easily distinguishable bryophyte functional groups. To illustrate how bryophyte functional groups can help elucidate variation in bryophyte effects and responses, we compiled existing data on water holding capacity, a key bryophyte trait. Although plant functional groups can mask potentially high interspecific and intraspecific variability, we found better separation of bryophyte functional group means compared with previous grouping systems regarding water holding capacity. This suggests that our bryophyte functional groups truly represent variation in the functional roles of bryophytes in tundra ecosystems. Lastly, we provide recommendations to improve the monitoring of bryophyte community changes in tundra study sites. 

Global change drivers, such as anthropogenic nutrient inputs, are increasing globally. Nutrient deposition simultaneously alters plant biodiversity, species composition and ecosystem processes like aboveground biomass production. These changes are underpinned by species extinction, colonisation and shifting relative abundance. Here, we use the Price equation to quantify and link the contributions of species that are lost, gained or that persist to change in aboveground biomass in 59 experimental grassland sites. Under ambient (control) conditions, compositional and biomass turnover was high, and losses (i.e. local extinctions) were balanced by gains (i.e. colonisation). Under fertilisation, the decline in species richness resulted from increased species loss and decreases in species gained. Biomass increase under fertilisation resulted mostly from species that persist and to a lesser extent from species gained. Drivers of ecological change can interact relatively independently with diversity, composition and ecosystem processes and functions such as aboveground biomass due to the individual contributions of species lost, gained or persisting. 

Abstract Given the current rates of climate change, with associated shifts in herbivore population densities, understanding the role of different herbivores in ecosystem functioning is critical for predicting ecosystem responses. Here, we examined how migratory geese and resident, non‐migratory reindeer—two dominating yet functionally contrasting herbivores—control vegetation and ecosystem processes in rapidly warming Arctic tundra.We collected vegetation and ecosystem carbon (C) flux data at peak plant growing season in the two longest running, fully replicated herbivore removal experiments found in high‐Arctic Svalbard. Experiments had been set up independently in wet habitat utilised by barnacle geeseBranta leucopsisin summer and in moist‐to‐dry habitat utilised by wild reindeerRangifer tarandus platyrhynchusyear‐round.Excluding geese induced vegetation state transitions from heavily grazed, moss‐dominated (only 4 g m⁻²of live above‐ground vascular plant biomass) to ungrazed, graminoid‐dominated (60 g m⁻²after 4‐year exclusion) and horsetail‐dominated (150 g m⁻²after 15‐year exclusion) tundra. This caused large increases in vegetation C and nitrogen (N) pools, dead biomass and moss‐layer depth. Alterations in plant N concentration and CN ratio suggest overall slower plant community nutrient dynamics in the short‐term (4‐year) absence of geese. Long‐term (15‐year) goose removal quadrupled net ecosystem C sequestration (NEE) by increasing ecosystem photosynthesis more than ecosystem respiration (ER).Excluding reindeer for 21 years also produced detectable increases in live above‐ground vascular plant biomass (from 50 to 80 g m⁻²; without promoting vegetation state shifts), as well as in vegetation C and N pools, dead biomass, moss‐layer depth and ER. Yet, reindeer removal did not alter the chemistry of plants and soil or NEE.Synthesis. Although both herbivores were key drivers of ecosystem structure and function, the control exerted by geese in their main habitat (wet tundra) was much more pronounced than that exerted by reindeer in their main habitat (moist‐to‐dry tundra). Importantly, these herbivore effects are scale dependent, because geese are more spatially concentrated and thereby affect a smaller portion of the tundra landscape compared to reindeer. Our results highlight the substantial heterogeneity in how herbivores shape tundra vegetation and ecosystem processes, with implications for ongoing environmental change.