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Loss of plant diversity with increased anthropogenic nitrogen (N) deposition in grasslands has occurred globally. In most cases, competitive exclusion driven by preemption of light or space is invoked as a key mechanism. Here, we provide evidence from a 9‐yr N‐addition experiment for an alternative mechanism: differential sensitivity of forbs and grasses to increased soil manganese (Mn) levels. In Inner Mongolia steppes, increasing the N supply shifted plant community composition from grass–forb codominance (primarily Stipa krylovii and Artemisia frigida, respectively) to exclusive dominance by grass, with associated declines in overall species richness. Reduced abundance of forbs was linked to soil acidification that increased mobilization of soil Mn, with a 10‐fold greater accumulation of Mn in forbs than in grasses. The enhanced accumulation of Mn in forbs was correlated with reduced photosynthetic rates and growth, and is consistent with the loss of forb species. Differential accumulation of Mn between forbs and grasses can be linked to fundamental differences between dicots and monocots in the biochemical pathways regulating metal transport. These findings provide a mechanistic explanation for N‐induced species loss in temperate grasslands by linking metal mobilization in soil to differential metal acquisition and impacts on key functional groups in these ecosystems.

Grazing, fire, and climate shape mesic grassland communities. With global change altering all three factors, understanding how grasslands respond to changes in these combined drivers may aid in projecting future changes in grassland ecosystems. We manipulated rainfall and simulated grazing (clipping) in two long-term fire experiments in mesic grasslands in North America (NA) and South Africa (SA). Despite their common drivers, grasslands in NA and SA differ in evolutionary history. Therefore, we expected community structure and production in NA and SA to respond differently to fire, grazing, and drought. Specifically, we hypothesized that NA plant community composition and production would be more responsive than the SA plant communities to changes in the drivers and their interactions, and that despite this expected stability of SA grasslands, drought would be the dominant factor controlling production, but grazing would play the primary role in determining community composition at both sites. Contrary to our hypothesis, NA and SA grasslands generally responded similarly to grazing, drought, and fire. Grazing increased diversity, decreased grass cover and production, and decreased belowground biomass at both sites. Drought alone minimally impacted plant community structure, and we saw similar treatment interactions at the two sites. Drought was not the primary driver of grassland productivity, but instead drought effects were similar to or less than grazing and fire. Even though these grasslands differed in evolutionary history, they responded similarly to our fire, grazing, and climate manipulations. Overall, we found community and ecosystem convergence in NA and SA grasslands. Grazing and fire are as important as climate in controlling mesic grassland ecosystems on both continents.

The structure and function of alpine grassland ecosystems, including their extensive soil carbon stocks, are largely shaped by temperature. The Tibetan Plateau in particular has experienced significant warming over the past 50 y, and this warming trend is projected to intensify in the future. Such climate change will likely alter plant species composition and net primary production (NPP). Here we combined 32 y of observations and monitoring with a manipulative experiment of temperature and precipitation to explore the effects of changing climate on plant community structure and ecosystem function. First, long-term climate warming from 1983 to 2014, which occurred without systematic changes in precipitation, led to higher grass abundance and lower sedge abundance, but did not affect aboveground NPP. Second, an experimental warming experiment conducted over 4 y had no effects on any aspect of NPP, whereas drought manipulation (reducing precipitation by 50%), shifted NPP allocation belowground without affecting total NPP. Third, both experimental warming and drought treatments, supported by a meta-analysis at nine sites across the plateau, increased grass abundance at the expense of biomass of sedges and forbs. This shift in functional group composition led to deeper root systems, which may have enabled plant communities to acquire more water and thus stabilize ecosystem primary production even with a changing climate. Overall, our study demonstrates that shifting plant species composition in response to climate change may have stabilized primary production in this high-elevation ecosystem, but it also caused a shift from aboveground to belowground productivity.

Larger, more frequent wildfires in arid and semi‐arid ecosystems have been associated with invasion by non‐native annual grasses, yet a complete understanding of fine fuel development and subsequent wildfire trends is lacking. We investigated the complex relationships among weather, fine fuels, and fire in the Great Basin, USA. We first modeled the annual and time‐lagged effects of precipitation and temperature on herbaceous vegetation cover and litter accumulation over a 26‐year period in the northern Great Basin. We then modeled how these fine fuels and weather patterns influence subsequent wildfires. We found that cheatgrass cover increased in years with higher precipitation and especially when one of the previous 3 years also was particularly wet. Cover of non‐native forbs and native herbs also increased in wet years, but only after several dry years. The area burned by wildfire in a given year was mostly associated with native herb and non‐native forb cover, whereas cheatgrass mainly influenced area burned in the form of litter derived from previous years’ growth. Consequently, multiyear weather patterns, including precipitation in the previous 1–3 years, was a strong predictor of wildfire in a given year because of the time needed to develop these fine fuel loads. The strong relationship between precipitation and wildfire allowed us to expand our inference to 10,162 wildfires across the entire Great Basin over a 35‐year period from 1980 to 2014. Our results suggest that the region's precipitation pattern of consecutive wet years followed by consecutive dry years results in a cycle of fuel accumulation followed by weather conditions that increase the probability of wildfire events in the year when the cycle transitions from wet to dry. These patterns varied regionally but were strong enough to allow us to model annual wildfire risk across the Great Basin based on precipitation alone. We investigated the complex interactions of weather, fine fuels, and wildfires in the Great Basin. We found that both native and non‐native grasses and forbs influence the number of fires and area burned suggesting that cheatgrass is not the sole driver of the grass–fire cycle in this region. We also found that consecutive wet years and consecutive dry years result in a cycle of fuel accumulation followed by weather conditions that increase the probability of wildfire events in the year when the cycle transitions from wet to dry, a pattern that could be mapped and used to increase fire preparedness and plan fuel reduction.

Enhanced deposition of atmospheric nitrogen (N) has profound impacts on ecosystem processes such as above‐ground productivity and community structure in grasslands across the globe. But how N deposition affects below‐ground processes of grasslands is less well known. Here, we evaluated the effects of chronic N amendment at a relatively low rate (20 kg ha⁻¹ year⁻¹) on root traits (root productivity, root biomass, root/shoot ratio) in Inner Mongolia steppes by rhizotron and ingrowth core and soil monolith techniques at levels of individual species, functional groups and ecosystem. For 8 years, N amendment suppressed above‐ground net primary production (ANPP), photosynthetic rates and root biomass of forbs, but enhanced ANPP and root biomass of grasses. This led to an overall reduction in below‐ground productivity of the grassland by 24–33%, while ANPP remained unchanged. Nitrogen amendment acidified soil and subsequently increased extractable soil manganese (Mn) concentration. Nitrogen amendment increased foliar Mn concentrations in forb, but not grass species, leading to a significant inhibition of photosynthetic rates in forb species. Synthesis. These findings highlight the importance of the differentiating responses of plant functional groups to long‐term N deposition and the important consequences of these responses for below‐ground productivity and long‐term soil C sequestration.

Fertile soils have been an essential resource for humanity for 10,000 y, but the ecological mechanisms involved in the creation and restoration of fertile soils, and especially the role of plant diversity, are poorly understood. Here we use results of a long-term, unfertilized plant biodiversity experiment to determine whether biodiversity, especially plant functional biodiversity, impacted the regeneration of fertility on a degraded sandy soil. After 23 y, plots containing 16 perennial grassland plant species had, relative to monocultures of these same species, ∼30 to 90% greater increases in soil nitrogen, potassium, calcium, magnesium, cation exchange capacity, and carbon and had ∼150 to 370% greater amounts of N, K, Ca, and Mg in plant biomass. Our results suggest that biodiversity, likely in combination with the increased plant productivity caused by higher biodiversity, led to greater soil fertility. Moreover, plots with high plant functional diversity, those containing grasses, legumes, and forbs, accumulated significantly greater N, K, Ca, and Mg in the total nutrient pool (plant biomass and soil) than did plots containing just one of these three functional groups. Plant species in these functional groups had trade-offs between their tissue N content, tissue K content, and root mass, suggesting why species from all three functional groups were essential for regenerating soil fertility. Our findings suggest that efforts to regenerate soil C stores and soil fertility may be aided by creative uses of plant diversity.

• Understanding how plant species influence soil nutrient cycling is a major theme in terrestrial ecosystem ecology. However, the prevailing paradigm has mostly focused on litter decomposition, while rhizosphere effects on soil organic matter (SOM) decomposition have attracted little attention. • Using a dual 13C/15N labeling approach in a ‘common garden’ glasshouse experiment, we investigated how the economic strategies of 12 grassland plant species (graminoids, forbs and legumes) drive soil nitrogen (N) cycling via rhizosphere processes, and how this in turn affects plant N acquisition and growth. • Acquisitive species with higher photosynthesis, carbon rhizodeposition and N uptake than conservative species induced a stronger acceleration of soil N cycling through rhizosphere priming of SOM decomposition. This allowed them to take up larger amounts of N and allocate it above ground to promote photosynthesis, thereby sustaining their faster growth. The N₂-fixation ability of legumes enhanced rhizosphere priming by promoting photosynthesis and rhizodeposition. • Our study demonstrates that the economic strategies of plant species regulate a plant–soil carbon–nitrogen feedback operating through the rhizosphere. These findings provide novel mechanistic insights into how plant species with contrasting economic strategies sustain their nutrition and growth through regulating the cycling of nutrients by soil microbes in their rhizosphere.

Plants respond to resource stress by changing multiple aspects of their biomass allocation, morphology, physiology and architecture. To date, we lack an integrated view of the relative importance of these plastic responses in alleviating resource stress and of the consistency/variability of these responses among species. We subjected nine species (legumes, forbs and graminoids) to nitrogen and/or light shortages and measured 11 above-ground and below-ground trait adjustments critical in the alleviation of these stresses (plus several underlying traits). Nine traits out of 11 showed adjustments that improved plants’ potential capacity to acquire the limiting resource at a given time. Above ground, aspects of plasticity in allocation, morphology, physiology and architecture all appeared important in improving light capture, whereas below ground, plasticity in allocation and physiology were most critical to improving nitrogen acquisition. Six traits out of 11 showed substantial heterogeneity in species plasticity, with little structuration of these differences within trait covariation syndromes. Such comprehensive assessment of the complex nature of phenotypic responses of plants to multiple stress factors, and the comparison of plant responses across multiple species, makes a clear case for the high (but largely overlooked) diversity of potential plastic responses of plants, and for the need to explore the potential rules structuring them.

Background and aims Precipitation and nitrogen (N) deposition are predicted to increase in northern China. The present paper aimed to better understand how different dominant species in semi-arid grasslands in this region vary in their litter decomposition and nutrient release responses to increases in precipitation and N deposition. Methods Above-ground litter of three dominant species (two grasses, Agropyron cristatum and Stipa krylovii , and one forb, Artemisia frigida ) was collected from areas without experimental treatments in a semi-arid grassland in Inner Mongolia. Litter decomposition was studied over three years to determine the effects of water and N addition on litter decomposition rate and nutrient dynamics. Results Litter mass loss and nutrient release were faster for the forb species than for the two grasses during decomposition. Both water and N addition increased litter mass loss of the grass A. cristatum , while the treatments showed no impacts on that of the forb A. frigida . Supplemental N had time-dependent, positive effects on litter mass loss of the grass S. krylovii . During the three-year decomposition study, the release of N from litter was inhibited by N addition for the three species, and it was promoted by water addition for the two grasses. Across all treatments, N and potassium (K) were released from the litter of all three species, whereas calcium (Ca) was accumulated. Phosphorus (P) and magnesium (Mg) were released from the forb litter but accumulated in the grass litter after three years of decomposition. Conclusions Our findings revealed that the litter decomposition response to water and N supplementation differed among dominant plant species in a semi-arid grassland, indicating that changes in dominant plant species induced by projected increases in precipitation and N deposition are likely to affect litter decomposition, nutrient cycling, and further biogeochemical cycles in this grassland. The asynchronous nutrient release of different species’ litter found in the present study highlights the complexity of nutrient replenishment from litter decomposition in the temperate steppe under scenarios of enhancing precipitation and N deposition.

1. Management of high-nature-value (HNV) grasslands follows agri-environmental schemes across large areas of Europe. Long-term agreements and restrictions of fertilizers cause soil nutrient impoverishment, but remarkably this quite often does not reduce biomass production. Therefore, we tested how species-rich vegetation copes with nutrient impoverishment under the most frequently used treatments, that is summer mowing and livestock grazing. 2. During 2011–2012 we studied, simultaneously, plant species composition, soil and biomass chemical properties in two equally designed experiments where mowing, grazing or leaving fallow have been applied since 2004. We asked whether soil-based (Corg : Ntot, plant-available P and K) and plant-based measures (N : P, N : K, K : P ratios and N-, P-, K-nutrition indices) indicate the same pattern of nutrient limitation as the observed productivity gradient. 3. Seven years of management application resulted in the lowest plant-available P under grazing and the lowest plant-available K under mowing, but neither grazed nor mown plots produced less biomass than fallow ones. Grazing supported dominance of grasses while mowing that of non-leguminous forbs. 4. Projection of nutrition indices to a common framework with nutrient ratios suggests that critical thresholds for diagnosis of nutrient limitation are a function of N deficiency. At biomass production of 2 t ha 1 a N-nutrition index of 50 yielded threshold N : P = 140; hence, all our treatments with N : P of 99–125 should be N limited. 5. Inspecting the productivity gradient separately for each management, we found only soil Corg : Ntot negatively related to biomass production in mown plots indicating N limitation. However in grazed plots, positive association of biomass production with plant-available P and negative with biomass N : P and N : K suggested PK co-limitation. 6. Synthesis and applications. Mowing and grazing induced different patterns of soil nutrient impoverishment and nutrient limitation, but they did not reduce biomass production of highnature-value grasslands. Non-leguminous forbs prevailing under mowing precluded shortage of P, while grasses dominating under grazing efficiently captured N. We recommend designing agri-environmental measures that will encourage alternating mowing and grazing. This should promote coexistence of multiple forbs and grasses, balance nutrient limitation and ensure stable biomass production under future low-input scenarios.