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Little is currently known about how climate modulates the relationship between plant diversity and soil organic carbon and the mechanisms involved. Yet, this knowledge is of crucial importance in times of climate change and biodiversity loss. Here, we show that plant diversity is positively correlated with soil carbon content and soil carbon-to-nitrogen ratio across 84 grasslands on six continents that span wide climate gradients. The relationships between plant diversity and soil carbon as well as plant diversity and soil organic matter quality (carbon-to-nitrogen ratio) are particularly strong in warm and arid climates. While plant biomass is positively correlated with soil carbon, plant biomass is not significantly correlated with plant diversity. Our results indicate that plant diversity influences soil carbon storage not via the quantity of organic matter (plant biomass) inputs to soil, but through the quality of organic matter. The study implies that ecosystem management that restores plant diversity likely enhances soil carbon sequestration, particularly in warm and arid climates.

Background and aims The amount of nitrogen (N) derived from symbiotic N2 fixation by legumes in grasslands might be affected by anthropogenic N and phosphorus (P) inputs, but the underlying mechanisms are not known. Methods We evaluated symbiotic N2 fixation in 17 natural and semi-natural grasslands on four continents that are subjected to the same full-factorial N and P addition experiment, using the 15N natural abundance method. Results N as well as combined N and P (NP) addition reduced aboveground legume biomass by 65% and 45%, respectively, compared to the control, whereas P addition had no significant impact. Addition of N and/or P had no significant effect on the symbiotic N2 fixation per unit legume biomass. In consequence, the amount of N fixed annually per grassland area was less than half in the N addition treatments compared to control and P addition, irrespective of whether the dominant legumes were annuals or perennials. Conclusion Our results reveal that N addition mainly impacts symbiotic N2 fixation via reduced biomass of legumes rather than changes in N2 fixation per unit legume biomass. The results show that soil N enrichment by anthropogenic activities significantly reduces N2 fixation in grasslands, and these effects cannot be reversed by additional P amendment.

Background and aims: The amount of nitrogen (N) derived from symbiotic N₂ fixation by legumes in grasslands might be affected by anthropogenic N and phosphorus (P) inputs, but the underlying mechanisms are not known. Methods: We evaluated symbiotic N₂ fixation in 17 natural and semi-natural grasslands on four continents that are subjected to the same full-factorial N and P addition experiment, using the ¹⁵N natural abundance method. Results: N as well as combined N and P (NP) addition reduced aboveground legume biomass by 65% and 45%, respectively, compared to the control, whereas P addition had no significant impact. Addition of N and/or P had no significant effect on the symbiotic N₂ fixation per unit legume biomass. In consequence, the amount of N fixed annually per grassland area was less than half in the N addition treatments compared to control and P addition, irrespective of whether the dominant legumes were annuals or perennials. Conclusion: Our results reveal that N addition mainly impacts symbiotic N₂ fixation via reduced biomass of legumes rather than changes in N₂ fixation per unit legume biomass. The results show that soil N enrichment by anthropogenic activities significantly reduces N₂ fixation in grasslands, and these effects cannot be reversed by additional P amendment.

Droughts can strongly affect grassland productivity and biodiversity, but responses differ widely. Nutrient availability may be a critical factor explaining this variation, but is often ignored in analyses of drought responses. Here, we used a standardized nutrient addition experiment covering 10 European grasslands to test if full-factorial nitrogen, phosphorus, and potassium addition affected plant community responses to inter-annual variation in drought stress and to the extreme summer drought of 2018 in Europe. We found that nutrient addition amplified detrimental drought effects on community aboveground biomass production. Drought effects also differed between functional groups, with a negative effect on graminoid but not forb biomass production. Our results imply that eutrophication in grasslands, which promotes dominance of drought-sensitive graminoids over forbs, amplifies detrimental drought effects. In terms of climate change adaptation, agricultural management would benefit from taking into account differential drought impacts on fertilized versus unfertilized grasslands, which differ in ecosystem services they provide to society.

Droughts can strongly affect grassland productivity and biodiversity, but responses differ widely. Nutrient availability may be a critical factor explaining this variation, but is often ignored in analyses of drought responses. Here, we used a standardized nutrient addition experiment covering 10 European grasslands to test if full‐factorial NPK‐addition affected plant community responses to inter‐annual variation in drought stress and to the extreme summer drought of 2018 in Europe. We found that nutrient addition amplified detrimental drought effects on community above‐ground biomass production. Drought effects also differed between functional groups, with a negative effect on graminoid but not forb biomass production. Our results imply that eutrophication in grasslands, which promotes dominance of drought‐sensitive graminoids over forbs, amplifies detrimental drought effects. In terms of climate change adaptation, agricultural management would benefit from taking into account differential drought impacts on fertilized versus unfertilized grasslands, which differ in ecosystem services they provide to society.

Plant growth is often co‐limited by nitrogen (N) and phosphorus (P). Plants might use one element to acquire another (i.e. trading N for P and P for N), which potentially explains synergistic growth responses to NP addition. We studied a 66‐year‐old grassland experiment in South Africa that consists of four levels of N addition with and without P addition. We investigated the response of aboveground net primary production (ANPP) to N and P addition over the last 66 years. Further, we tested whether phosphatase activity and plant P uptake depend on N availability, and vice versa, whether non‐symbiotic N2 fixation and plant N uptake depend on P availability. We expected that the interaction of both elements promote processes of nutrient acquisition and contribute to synergistic plant growth effects in response to NP addition. We found synergistic N and P co‐limitation of ANPP for the period from 1951 to 2017 but the response to N and P addition diminished over time. In 2017, aboveground P stocks, relative rRNA operon abundance of arbuscular mycorrhizal fungi and soil organic P storage increased with N fertilization rate when N was added with P compared to the treatment in which only N was added. Further, N addition increased phosphatase activity, which indicates that plants used N to acquire P from organic sources. In contrast, aboveground N stocks and non‐symbiotic N2 fixation did not change significantly due to P addition. Taken together, our results indicate that trading N for P likely contributes to synergistic plant‐growth response. Plants used added N to mobilize and take up P from organic sources, inducing stronger recycling of P and making the plant community less sensitive to external nutrient inputs. The latter could explain why indications of synergistic co‐limitation diminished over time, which is usually overlooked in short‐term nutrient addition experiments.

10Microorganisms mediate nutrient cycling in soils, and thus it is assumed that they largely control responses of terrestrial ecosystems to anthropogenic nutrient inputs. Therefore, it is important to understand how increased nitrogen (N) and phosphorus (P) availabilities, first, affect soil prokaryotic and fungal community composition and second, if and how changes in the community composition affect soil element cycling. We measured soil microbial communities and soil element cycling processes along a nine-year old experimental N-addition gradient partially crossed with a P-addition treatment in a temperate grassland. Nitrogen addition affected microbial community composition, and prokaryotic communities were less sensitive to N addition than fungal communities. P addition only marginally affected microbial community composition, indicating that P is less selective than N for microbial taxa in this grassland. Soil pH and total organic carbon (C) concentration were the main factors associated with prokaryotic community composition, while the dissolved organic C-to-dissolved N ratio was the predominant driver of fungal community composition. Against our expectation, plant biomass and plant community structure only explained a small proportion of the microbial community composition. Although microbial community composition changed with nutrient addition, microbial biomass concentrations and respiration rates did not change, indicating functional redundancy of the microbial community. Microbial respiration, net N mineralization, and non-symbiotic N2 fixation were more strongly controlled by abiotic factors than by plant biomass, plant community structure or microbial community, showing that community shifts under increasing nutrient inputs may not necessarily be reflected in element cycling rates. This study suggests that atmospheric N deposition may impact the composition of fungi more than of prokaryotes and that nutrient inputs act directly on element-cycling rates

In forests, where the supply of bioavailable phosphorus (P) from easily weatherable primary minerals is small, plants are thought to recycle P efficiently by uptake of P released from decomposing forest floor material. Yet a share of the P is leached into the subsoil, where it is strongly adsorbed onto the reactive surfaces of pedogenic Fe and Al oxides. This raises the question of whether P leached into subsoil is also recycled. To investigate the mobilization of P bound to hydrous Fe oxides, we conducted a mesocosm experiment in a greenhouse. Beech saplings were grown for 14 months in subsoil material (Bw horizon from the P-poor Lüss beech forest) with added goethite-P adsorption complexes, in either inorganic (orthophosphate) or organic (phytate) form. Four types of control mesocosms were run: soil only and soil mixed with either dissolved orthophosphate or dissolved phytate or goethite. At the end of the experiment, neither total P mass in trees nor P contents in leaves differed between the treatments. According to leaf nutrient contents, plant growth was strongly limited by P in all treatments. Yet total P mass in trees did not increase over the course of the experiment. Thus, despite its P demand, beech was not able to acquire P from goethite surfaces within two vegetation periods. Also P added in dissolved form to the soil before transplanting as well as native soil P were not available. This suggests that, once inorganic and organic P is bound to pedogenic metal oxides in mineral soil, it is not or hardly recycled, which can be an explanation for field data demonstrating quantitatively significant stocks of P in the subsoil of P-deficient forests.

Background: The dynamics of phosphorus (P) in the environment is important for regulating nutrient cycles in natural and managed ecosystems and an integral part in assessing biological resilience against environmental change. Organic P (Po) compounds play key roles in biological and ecosystems function in the terrestrial environment being critical to cell function, growth and reproduction. Scope: We asked a group of experts to consider the global issues associated with Po in the terrestrial environment, methodological strengths and weaknesses, benefits to be gained from understanding the Po cycle, and to set priorities for Po research. Conclusions: We identified seven key opportunities for Po research including: the need for integrated, quality controlled and functionally based methodologies; assessment of stoichiometry with other elements in organic matter; understanding the dynamics of Po in natural and managed systems; the role of microorganisms in controlling Po cycles; the implications of nanoparticles in the environment and the need for better modelling and communication of the research. Each priority is discussed and a statement of intent for the Po research community is made that highlights there are key contributions to be made toward understanding biogeochemical cycles, dynamics and function of natural ecosystems and the management of agricultural systems.

The article “Organic phosphorus in the terrestrial environment: a perspective on the state of the art and future priorities”, written by Timothy S George et al., was originally published with incorrect affiliation information for one of the co-authors, E. Klumpp.

Nitrogen (N) and phosphorus (P) in topsoils are critical for plant nutrition. Relatively little is known about the spatial patterns of N and P in the organic layer of mountainous landscapes. Therefore, the spatial distributions of N and P in both the organic layer and the A horizon were analyzed using a light detection and ranging (LiDAR) digital elevation model and vegetation metrics. The objective of the study was to analyze the effect of vegetation and topography on the spatial patterns of N and P in a small watershed covered by forest in South Korea. Soil samples were collected using the conditioned latin hypercube method. LiDAR vegetation metrics, the normalized difference vegetation index (NDVI), and terrain parameters were derived as predictors. Spatial explicit predictions of N/P ratios were obtained using a random forest with uncertainty analysis. We tested different strategies of model validation (repeated 2-fold to 20-fold and leave-one-out cross validation). Repeated 10-fold cross validation was selected for model validation due to the comparatively high accuracy and low variance of prediction. Surface curvature was the best predictor of P contents in the organic layer and in the A horizon, while LiDAR vegetation metrics and NDVI were important predictors of N in the organic layer. N/P ratios increased with surface curvature and were higher on the convex upper slope than on the concave lower slope. This was due to P enrichment of the soil on the lower slope and a more even spatial distribution of N. Our digital soil maps showed that the topsoils on the upper slopes contained relatively little P. These findings are critical for understanding N and P dynamics in mountainous ecosystems.