Search

Renewable energy is a promising alternative to fossil fuel-based energy, but its development can require a complex set of environmental tradeoffs. A recent increase in solar energy systems, especially large, centralized installations, underscores the urgency of understanding their environmental interactions. Synthesizing literature across numerous disciplines, we review direct and indirect environmental impacts - both beneficial and adverse - of utility-scale solar energy (USSE) development, including impacts on biodiversity, land-use and land-cover change, soils, water resources, and human health. Additionally, we review feedbacks between USSE infrastructure and land-atmosphere interactions and the potential for USSE systems to mitigate climate change. Several characteristics and development strategies of USSE systems have low environmental impacts relative to other energy systems, including other renewables. We show opportunities to increase USSE environmental co-benefits, the permitting and regulatory constraints and opportunities of USSE, and highlight future research directions to better understand the nexus between USSE and the environment. Increasing the environmental compatibility of USSE systems will maximize the efficacy of this key renewable energy source in mitigating climatic and global environmental change. © 2013 Elsevier Ltd.

Monitoring agriculture by remote sensing enables large-scale evaluation of biomass production across space and time. The normalized difference vegetation index (NDVI) is used as a proxy for green biomass. Here, we used satellite-derived NDVI of arable farms in the Netherlands to evaluate changes in biomass following conversion from conventional to organic farming. We compared NDVI and the stability of NDVI across 72 fields on sand and marine clay soils. Thirty-six of these fields had been converted into organic agriculture between 0 and 50 years ago (with 2017 as reference year), while the other 36 were paired control fields where conventional farming continued. We used high-resolution images from the Sentinel-2 satellite to obtain NDVI estimates across 5 years (January 2016-October 2020). Overall, NDVI did not differ between conventional and organic management during the time series, but NDVI stability was significantly higher under organic management. NDVI was lower under organic management in sandy, but not in clay, soils. Organic farms that had been converted less than ~19 years ago had lower NDVI than conventional farms. However, the difference diminished over time and eventually turned positive after ~19 years since the conversion. NDVI, averaged across the 5 years of study, was positively correlated to soil Olsen-P measured from soil samples collected in 2017. We conclude that NDVI in organic fields was more stable than in conventional fields, and that the lower biomass in the early years since the transition to organic agriculture can be overcome with time. Our study also indicates the role of soil P bioavailability for plant biomass production across the examined fields, and the benefit of combining remote sensing with on-site soil measurements to develop a more mechanistic understanding that may help us navigate the transition to a more sustainable type of agriculture., (© 2024 The Author(s). Global Change Biology published by John Wiley & Sons Ltd.)

The importance of phosphorus (P) in regulating ecosystem responses to climate change has fostered P-cycle implementation in land surface models, but their CO 2 effects predictions have not been evaluated against measurements. Here, we perform a data-driven model evaluation where simulations of eight widely used P-enabled models were confronted with observations from a long-term free-air CO 2 enrichment experiment in a mature, P-limited Eucalyptus forest. We show that most models predicted the correct sign and magnitude of the CO 2 effect on ecosystem carbon (C) sequestration, but they generally overestimated the effects on plant C uptake and growth. We identify leaf-to-canopy scaling of photosynthesis, plant tissue stoichiometry, plant belowground C allocation, and the subsequent consequences for plant-microbial interaction as key areas in which models of ecosystem C-P interaction can be improved. Together, this data-model intercomparison reveals data-driven insights into the performance and functionality of P-enabled models and adds to the existing evidence that the global CO 2 -driven carbon sink is overestimated by models.

The rhizosphere influence on the soil microbiome and function of crop wild progenitors (CWPs) remains virtually unknown, despite its relevance to develop microbiome-oriented tools in sustainable agriculture. Here, we quantified the rhizosphere influence-a comparison between rhizosphere and bulk soil samples-on bacterial, fungal, protists and invertebrate communities and on soil multifunctionality across nine CWPs at their sites of origin. Overall, rhizosphere influence was higher for abundant taxa across the four microbial groups and had a positive influence on rhizosphere soil organic C and nutrient contents compared to bulk soils. The rhizosphere influence on abundant soil microbiomes was more important for soil multifunctionality than rare taxa and environmental conditions. Our results are a starting point towards the use of CWPs for rhizosphere engineering in modern crops., (© 2024 The Author(s). Ecology Letters published by John Wiley & Sons Ltd.)

The capacity for terrestrial ecosystems to sequester additional carbon (C) with rising CO 2 concentrations depends on soil nutrient availability 1,2 . Previous evidence suggested that mature forests growing on phosphorus (P)-deprived soils had limited capacity to sequester extra biomass under elevated CO 2 (refs. 3-6 ), but uncertainty about ecosystem P cycling and its CO 2 response represents a crucial bottleneck for mechanistic prediction of the land C sink under climate change 7 . Here, by compiling the first comprehensive P budget for a P-limited mature forest exposed to elevated CO 2 , we show a high likelihood that P captured by soil microorganisms constrains ecosystem P recycling and availability for plant uptake. Trees used P efficiently, but microbial pre-emption of mineralized soil P seemed to limit the capacity of trees for increased P uptake and assimilation under elevated CO 2 and, therefore, their capacity to sequester extra C. Plant strategies to stimulate microbial P cycling and plant P uptake, such as increasing rhizosphere C release to soil, will probably be necessary for P-limited forests to increase C capture into new biomass. Our results identify the key mechanisms by which P availability limits CO 2 fertilization of tree growth and will guide the development of Earth system models to predict future long-term C storage., (© 2024. Crown.)

The breakdown of plant material fuels soil functioning and biodiversity. Currently, process understanding of global decomposition patterns and the drivers of such patterns are hampered by the lack of coherent large-scale datasets. We buried 36,000 individual litterbags (tea bags) worldwide and found an overall negative correlation between initial mass-loss rates and stabilization factors of plant-derived carbon, using the Tea Bag Index (TBI). The stabilization factor quantifies the degree to which easy-to-degrade components accumulate during early-stage decomposition (e.g. by environmental limitations). However, agriculture and an interaction between moisture and temperature led to a decoupling between initial mass-loss rates and stabilization, notably in colder locations. Using TBI improved mass-loss estimates of natural litter compared to models that ignored stabilization. Ignoring the transformation of dead plant material to more recalcitrant substances during early-stage decomposition, and the environmental control of this transformation, could overestimate carbon losses during early decomposition in carbon cycle models., (© 2024 The Authors. Ecology Letters published by John Wiley & Sons Ltd.)

Plant-soil biodiversity interactions are fundamental for the functioning of terrestrial ecosystems. Yet, the existence of a set of globally distributed topsoil microbial and small invertebrate organisms consistently associated with land plants (i.e., their consistent soil-borne microbiome), together with the environmental preferences and functional capabilities of these organisms, remains unknown. We conducted a standardized field survey under 150 species of land plants, including 58 species of bryophytes and 92 of vascular plants, across 124 locations from all continents. We found that, despite the immense biodiversity of soil organisms, the land plants evaluated only shared a small fraction (less than 1%) of all microbial and invertebrate taxa that were present across contrasting climatic and soil conditions and vegetation types. These consistent taxa were dominated by generalist decomposers and phagotrophs and their presence was positively correlated with the abundance of functional genes linked to mineralization. Finally, we showed that crossing environmental thresholds in aridity (aridity index of 0.65, i.e., the transition from mesic to dry ecosystems), soil pH (5.5; i.e., the transition from acidic to strongly acidic soils), and carbon (less than 2%, the lower limit of fertile soils) can result in drastic disruptions in the associations between land plants and soil organisms, with potential implications for the delivery of soil ecosystem processes under ongoing global environmental change., (© 2024 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Emerging evidence points out that the responses of soil organic carbon (SOC) to nitrogen (N) addition differ along the soil profile, highlighting the importance of synthesizing results from different soil layers. Here, using a global meta-analysis, we found that N addition significantly enhanced topsoil (0-30 cm) SOC by 3.7% (±1.4%) in forests and grasslands. In contrast, SOC in the subsoil (30-100 cm) initially increased with N addition but decreased over time. The model selection analysis revealed that experimental duration and vegetation type are among the most important predictors across a wide range of climatic, environmental, and edaphic variables. The contrasting responses of SOC to N addition indicate the importance of considering deep soil layers, particularly for long-term continuous N deposition. Finally, the lack of depth-dependent SOC responses to N addition in experimental and modeling frameworks has likely resulted in the overestimation of changes in SOC storage under enhanced N deposition., (© 2024 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Springtails (Collembola) inhabit soils from the Arctic to the Antarctic and comprise an estimated ~32% of all terrestrial arthropods on Earth. Here, we present a global, spatially-explicit database on springtail communities that includes 249,912 occurrences from 44,999 samples and 2,990 sites. These data are mainly raw sample-level records at the species level collected predominantly from private archives of the authors that were quality-controlled and taxonomically-standardised. Despite covering all continents, most of the sample-level data come from the European continent (82.5% of all samples) and represent four habitats: woodlands (57.4%), grasslands (14.0%), agrosystems (13.7%) and scrublands (9.0%). We included sampling by soil layers, and across seasons and years, representing temporal and spatial within-site variation in springtail communities. We also provided data use and sharing guidelines and R code to facilitate the use of the database by other researchers. This data paper describes a static version of the database at the publication date, but the database will be further expanded to include underrepresented regions and linked with trait data., (© 2024. The Author(s).)

Microbes inhabiting deep soil layers are known to be different from their counterpart in topsoil yet remain under investigation in terms of their structure, function, and how their diversity is shaped. The microbiome of deep soils (>1 m) is expected to be relatively stable and highly independent from climatic conditions. Much less is known, however, on how these microbial communities vary along climate gradients. Here, we used amplicon sequencing to investigate bacteria, archaea, and fungi along fifteen 18-m depth profiles at 20-50-cm intervals across contrasting aridity conditions in semi-arid forest ecosystems of China's Loess Plateau. Our results showed that bacterial and fungal α diversity and bacterial and archaeal community similarity declined dramatically in topsoil and remained relatively stable in deep soil. Nevertheless, deep soil microbiome still showed the functional potential of N cycling, plant-derived organic matter degradation, resource exchange, and water coordination. The deep soil microbiome had closer taxa-taxa and bacteria-fungi associations and more influence of dispersal limitation than topsoil microbiome. Geographic distance was more influential in deep soil bacteria and archaea than in topsoil. We further showed that aridity was negatively correlated with deep-soil archaeal and fungal richness, archaeal community similarity, relative abundance of plant saprotroph, and bacteria-fungi associations, but increased the relative abundance of aerobic ammonia oxidation, manganese oxidation, and arbuscular mycorrhizal in the deep soils. Root depth, complexity, soil volumetric moisture, and clay play bridging roles in the indirect effects of aridity on microbes in deep soils. Our work indicates that, even microbial communities and nutrient cycling in deep soil are susceptible to changes in water availability, with consequences for understanding the sustainability of dryland ecosystems and the whole-soil in response to aridification. Moreover, we propose that neglecting soil depth may underestimate the role of soil moisture in dryland ecosystems under future climate scenarios., (© 2023 John Wiley & Sons Ltd.)

Increased atmospheric nitrogen (N) deposition affects biodiversity in terrestrial ecosystems. However, we do not know whether the effects of N on above-ground plant β-diversity are coupled with changes occurring in the soil seed bank. We conducted a long-term N-addition experiment in a typical steppe and found that above-ground β-diversity increased and then decreased with increasing N addition, whereas below-ground β-diversity decreased linearly. This suggests decoupled dynamics of plant communities and their soil seed bank under N enrichment. Species substitution determined above- and below-ground β-diversity change via an increasing role of deterministic processes with N addition. These effects were mostly driven by differential responses of the above-ground vegetation and the soil seed bank β-diversities to N-induced changes in environmental heterogeneity, increased soil inorganic N concentrations and soil acidification. Our findings highlight the importance of considering above- and below-ground processes simultaneously for effectively conserving grassland ecosystems under N enrichment., (© 2023 John Wiley & Sons Ltd.)

The decomposition of litter and the supply of nutrients into and from the soil are two fundamental processes through which the above- and belowground world interact. Microbial biodiversity, and especially that of decomposers, plays a key role in these processes by helping litter decomposition. Yet the relative contribution of litter diversity and soil biodiversity in supporting multiple ecosystem services remains virtually unknown. Here we conducted a mesocosm experiment where leaf litter and soil biodiversity were manipulated to investigate their influence on plant productivity, litter decomposition, soil respiration, and enzymatic activity in the littersphere. We showed that both leaf litter diversity and soil microbial diversity (richness and community composition) independently contributed to explain multiple ecosystem functions. Fungal saprobes community composition was especially important for supporting ecosystem multifunctionality (EMF), plant production, litter decomposition, and activity of soil phosphatase when compared with bacteria or other fungal functional groups and litter species richness. Moreover, leaf litter diversity and soil microbial diversity exerted previously undescribed and significantly interactive effects on EMF and multiple individual ecosystem functions, such as litter decomposition and plant production. Together, our work provides experimental evidence supporting the independent and interactive roles of litter and belowground soil biodiversity to maintain ecosystem functions and multiple services., (© 2023 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Human activities cause substantial changes in biodiversity. 1 , 2 Despite ongoing concern about the implications of invertebrate decline, 3 , 4 , 5 , 6 , 7 few empirical studies have examined the ecosystem consequences of invertebrate biomass loss. Here, we test the responses of six ecosystem services informed by 30 above- and belowground ecosystem variables to three levels of aboveground (i.e., vegetation associated) invertebrate community biomass (100%, 36%, and 0% of ambient biomass) in experimental grassland mesocosms in a controlled Ecotron facility. In line with recent reports on invertebrate biomass loss over the last decade, our 36% biomass treatment also represented a decrease in invertebrate abundance (-70%) and richness (-44%). Moreover, we simulated the pronounced change in invertebrate biomass and turnover in community composition across the season. We found that the loss of invertebrate biomass decreases ecosystem multifunctionality, including two critical ecosystem services, aboveground pest control and belowground decomposition, while harvested plant biomass increases, likely because less energy was channeled up the food chain. Moreover, communities and ecosystem functions become decoupled with a lower biomass of invertebrates. Our study shows that invertebrate loss threatens the integrity of grasslands by decoupling ecosystem processes and decreasing ecosystem-service supply., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2023 Elsevier Inc. All rights reserved.)

Soil life supports the functioning and biodiversity of terrestrial ecosystems. Springtails (Collembola) are among the most abundant soil arthropods regulating soil fertility and flow of energy through above- and belowground food webs. However, the global distribution of springtail diversity and density, and how these relate to energy fluxes remains unknown. Here, using a global dataset representing 2470 sites, we estimate the total soil springtail biomass at 27.5 megatons carbon, which is threefold higher than wild terrestrial vertebrates, and record peak densities up to 2 million individuals per square meter in the tundra. Despite a 20-fold biomass difference between the tundra and the tropics, springtail energy use (community metabolism) remains similar across the latitudinal gradient, owing to the changes in temperature with latitude. Neither springtail density nor community metabolism is predicted by local species richness, which is high in the tropics, but comparably high in some temperate forests and even tundra. Changes in springtail activity may emerge from latitudinal gradients in temperature, predation and resource limitation in soil communities. Contrasting relationships of biomass, diversity and activity of springtail communities with temperature suggest that climate warming will alter fundamental soil biodiversity metrics in different directions, potentially restructuring terrestrial food webs and affecting soil functioning., (© 2023. The Author(s).)

Biological soil crusts (BSC) are associations of different macro and microorganisms and aggregated soil particles located on the surface of soils in many different habitats. BSC harbour a diverse and complex community of ciliates and testate amoebae. These phagotrophic protists play an important role in C and N recycling in soil ecosystems but have not been frequently studied in BSC. In this context, the effects of three increasing N inputs on ciliates and testate amoebae in crusts from a semi-arid Mediterranean ecosystem were evaluated. A field experiment with artificial N-deposition was designed to mimic the effects caused by anthropogenic N depositions. The results have shown that the protist populations of these semi-arid Mediterranean environments have lower species richness than other soil environments. The increase in N produces a net loss of diversity in the populations studied and shifts in the community structure. It has also been shown that some ciliates and testate amoebae, due to their population responses to increased N concentrations, could potentially be used as bio-indicators of N contamination in these BSCs., (Copyright © 2022 The Author(s). Published by Elsevier GmbH.. All rights reserved.)

Soil micronutrients are capital for the delivery of ecosystem functioning and food provision worldwide. Yet, despite their importance, the global biogeography and ecological drivers of soil micronutrients remain virtually unknown, limiting our capacity to anticipate abrupt unexpected changes in soil micronutrients in the face of climate change. Here, we analyzed >1300 topsoil samples to examine the global distribution of six metallic micronutrients (Cu, Fe, Mn, Zn, Co and Ni) across all continents, climates and vegetation types. We found that warmer arid and tropical ecosystems, present in the least developed countries, sustain the lowest contents of multiple soil micronutrients. We further provide evidence that temperature increases may potentially result in abrupt and simultaneous reductions in the content of multiple soil micronutrients when a temperature threshold of 12-14°C is crossed, which may be occurring on 3% of the planet over the next century. Altogether, our findings provide fundamental understanding of the global distribution of soil micronutrients, with direct implications for the maintenance of ecosystem functioning, rangeland management and food production in the warmest and poorest regions of the planet., (© 2022 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Current and continuing climate change in the Anthropocene epoch requires sustainable agricultural practices. Additionally, due to changing consumer preferences, organic approaches to cultivation are gaining popularity. The global market for organic grapes, grape products, and wine is growing. Biostimulant and biocontrol products are often applied in organic vineyards and can reduce the synthetic fertilizer, pesticide, and fungicide requirements of a vineyard. Plant growth promotion following application is also observed under a variety of challenging conditions associated with global warming. This paper reviews different groups of biostimulants and their effects on viticulture, including microorganisms, protein hydrolysates, humic acids, pyrogenic materials, and seaweed extracts. Of special interest are biostimulants with utility in protecting plants against the effects of climate change, including drought and heat stress. While many beneficial effects have been reported following the application of these materials, most studies lack a mechanistic explanation, and important parameters are often undefined (e.g., soil characteristics and nutrient availability). We recommend an increased study of the underlying mechanisms of these products to enable the selection of proper biostimulants, application methods, and dosage in viticulture. A detailed understanding of processes dictating beneficial effects in vineyards following application may allow for biostimulants with increased efficacy, uptake, and sustainability., Competing Interests: Author YA was employed by Chitose Laboratory Corp. The remaining authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Jindo, Goron, Pizarro-Tobías, Sánchez-Monedero, Audette, Deolu-Ajayi, van der Werf, Goitom Teklu, Shenker, Pombo Sudré, Busato, Ochoa-Hueso, Nocentini, Rippen, Aroca, Mesa, Delgado and Tortosa.)

Grasslands are now facing a continuously increasing supply of nitrogen (N) fertilizers, resulting in alterations in ecosystem functioning, including changes in carbon (C) and water cycling. Mowing, one of the most widely used grassland management techniques, has been shown to mitigate the negative impacts of increased N availability on species richness. However, knowledge of how N addition and mowing, alone and/or in combination, affect ecosystem-level C fluxes and water use efficiency (W N ) is still limited. We experimentally manipulated N fertilization (0 and 10 g N m -2 yr -1 ) and mowing (once per year at the end of the growing season) following a randomized block design in a meadow steppe characterized by salinization and alkalinization in northeastern China. We found that, compared to the control plots, N addition, mowing, and their interaction increased net ecosystem CO 2 exchange by 65.1%, 14.7%, and 133%, and W N by 40.7%, 18.5%, and 96.1%, respectively. Nitrogen enrichment also decreased soil pH, which resulted in greater aboveground biomass (AGB). Moreover, N addition indirectly increased AGB by inducing changes in species richness. Our results indicate that mowing enhances the positive effects of N addition on ecosystem C fluxes and W N . Therefore, appropriate grassland management practices are essential to improve ecosystem C sequestration, W N , and mitigate future species diversity declines due to ecosystem eutrophication., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Soil carbon (C) stabilization partially depends on its distribution within soil structural aggregates, and on the physicochemical processes of C within these aggregates. Changes in precipitation can alter the size distribution of aggregate classes within soils, and C input and output processes within these aggregates, which have potential consequences for soil C storage. However, the mechanisms underlying C accumulation within different aggregates under various precipitation regimes remain unclear. In this study, we conducted a 3-year field manipulation experiment to test the effects of a gradient of altered precipitation (-70%, -50%, -30%, 0%, +30%, and +50% amounts compared with ambient rainfall) on soil aggregate distribution and C accumulation in aggregates (53-250 μm, microaggregates; < 53 μm, silt and clay fractions) in a meadow steppe of northeastern China. Our results revealed that the distribution of soil microaggregates decreased along the precipitation gradient, with no detectable discrepant responses with respect to soil C accumulation within the microaggregates to precipitation treatments. In contrast, higher precipitation amounts coupled with a greater proportion of silt and clay fractions enhanced the accumulation of soil C. Importantly, structural equation models revealed that the pathways by which changes in precipitation control the accumulation of soil C varied across aggregate size fractions. Plant biomass was the main direct factor controlling the accumulation of C within soil microaggregates, whereas soil aggregate distribution and enzyme activities strongly interacted with soil C accumulation in the silt and clay fractions. Our findings imply that identifying how plant and soil aggregate properties respond to precipitation changes and drive C accumulation among soil particles will enhance the ability to predict responses of ecosystem processes to future global change., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier B.V. All rights reserved.)

The growing demand for timber and the boom in massive tree-planting programs could mean the spreading of mismanaged tree plantations worldwide. Here, we apply the concept of ecological intensification to forestry systems as a viable biodiversity-focused strategy that could be critical to develop productive, yet sustainable, tree plantations. Tree plantations can be highly productive if tree species are properly combined to complement their ecological functions. Simultaneously considering soil biodiversity and animal-mediated biocontrol will be critical to minimize the reliance on external inputs. Integrating genetic, functional, and demographic diversity across heterogeneous landscapes should improve resilience under climate change. Designing ecologically intensified plantations will mean breaking the timber productivity versus conservation dichotomy and assuring the maintenance of key ecosystem services at safe levels., Competing Interests: Declaration of interests None declared., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Spatial rarity is often used to predict extinction risk, but rarity can also occur temporally. Perhaps more relevant in the context of global change is whether a species is core to a community (persistent) or transient (intermittently present), with transient species often susceptible to human activities that reduce niche space. Using 5-12 yr of data on 1,447 plant species from 49 grasslands on five continents, we show that local abundance and species persistence under ambient conditions are both effective predictors of local extinction risk following experimental exclusion of grazers or addition of nutrients; persistence was a more powerful predictor than local abundance. While perturbations increased the risk of exclusion for low persistence and abundance species, transient but abundant species were also highly likely to be excluded from a perturbed plot relative to ambient conditions. Moreover, low persistence and low abundance species that were not excluded from perturbed plots tended to have a modest increase in abundance following perturbance. Last, even core species with high abundances had large decreases in persistence and increased losses in perturbed plots, threatening the long-term stability of these grasslands. Our results demonstrate that expanding the concept of rarity to include temporal dynamics, in addition to local abundance, more effectively predicts extinction risk in response to environmental change than either rarity axis predicts alone., (© 2021 by the Ecological Society of America.)

Background: Climate change models predict changes in the amount, frequency and seasonality of precipitation events, all of which have the potential to affect the structure and function of grassland ecosystems. While previous studies have examined plant or herbivore responses to these perturbations, few have examined their interactions; even fewer have included belowground herbivores. Given the ecological, economic and biodiversity value of grasslands, and their importance globally for carbon storage and agriculture, this is an important knowledge gap. To address this, we conducted a precipitation manipulation experiment in a former mesic pasture grassland comprising a mixture of C 4 grasses and C 3 grasses and forbs, in southeast Australia. Rainfall treatments included a control [ambient], reduced amount [50% ambient] and reduced frequency [ambient rainfall withheld for three weeks, then applied as a single deluge event] manipulations, to simulate predicted changes in both the size and frequency of future rainfall events. In addition, half of all experimental plots were inoculated with adult root herbivores (Scarabaeidae beetles)., Results: We found strong seasonal dependence in plant community responses to both rainfall and root herbivore treatments. The largest effects were seen in the cool season with lower productivity, cover and diversity in rainfall-manipulated plots, while root herbivore inoculation increased the relative abundance of C 3 , compared to C 4 , plants., Conclusions: This study highlights the importance of considering not only the seasonality of plant responses to altered rainfall, but also the important role of interactions between abiotic and biotic drivers of vegetation change when evaluating ecosystem-level responses to future shifts in climatic conditions., (© 2021. The Author(s).)

Despite their extent and socio-ecological importance, a comprehensive biogeographical synthesis of drylands is lacking. Here we synthesize the biogeography of key organisms (vascular and nonvascular vegetation and soil microorganisms), attributes (functional traits, spatial patterns, plant-plant and plant-soil interactions) and processes (productivity and land cover) across global drylands. These areas have a long evolutionary history, are centers of diversification for many plant lineages and include important plant diversity hotspots. This diversity captures a strikingly high portion of the variation in leaf functional diversity observed globally. Part of this functional diversity is associated with the large variation in response and effect traits in the shrubs encroaching dryland grasslands. Aridity and its interplay with the traits of interacting plant species largely shape biogeographical patterns in plant-plant and plant-soil interactions, and in plant spatial patterns. Aridity also drives the composition of biocrust communities and vegetation productivity, which shows large geographical variation. We finish our review by discussing major research gaps, which include: studying regular vegetation spatial patterns; establishing large-scale plant and biocrust field surveys assessing individual-level trait measurements; knowing whether the impacts of plant-plant and plant-soil interactions on biodiversity are predictable; and assessing how elevated CO 2 modulates future aridity conditions and plant productivity., (© 2021 The Authors New Phytologist © 2021 New Phytologist Foundation.)

Soil nitrogen (N) availability is critical for grassland functioning. However, human activities have increased the supply of biologically limiting nutrients, and changed the density and identity of mammalian herbivores. These anthropogenic changes may alter net soil N mineralization (soil net N min ), that is, the net balance between N mineralization and immobilization, which could severely impact grassland structure and functioning. Yet, to date, little is known about how fertilization and herbivore removal individually, or jointly, affect soil net N min across a wide range of grasslands that vary in soil and climatic properties. Here we collected data from 22 grasslands on five continents, all part of a globally replicated experiment, to assess how fertilization and herbivore removal affected potential (laboratory-based) and realized (field-based) soil net N min . Herbivore removal in the absence of fertilization did not alter potential and realized soil net N min . However, fertilization alone and in combination with herbivore removal consistently increased potential soil net N min. Realized soil net N min , in contrast, significantly decreased in fertilized plots where herbivores were removed. Treatment effects on potential and realized soil net N min were contingent on site-specific soil and climatic properties. Fertilization effects on potential soil net N min were larger at sites with higher mean annual precipitation (MAP) and temperature of the wettest quarter (T.q.wet). Reciprocally, realized soil net N min declined most strongly with fertilization and herbivore removal at sites with lower MAP and higher T.q.wet. In summary, our findings show that anthropogenic nutrient enrichment, herbivore exclusion and alterations in future climatic conditions can negatively impact soil net N min across global grasslands under realistic field conditions. This is an important context-dependent knowledge for grassland management worldwide., (© 2020 John Wiley & Sons Ltd.)

Soil nitrogen (N) availability is a key driver of soil-atmosphere greenhouse gas (GHG) exchange, yet we are far from understanding how increases in N deposition due to human activities will influence the net soil-atmosphere fluxes of the three most important GHGs: nitrous oxide (N 2 O), methane (CH 4 ) and carbon dioxide (CO 2 ). We simulated four levels of N deposition (10, 20 and 50 kg N ha -1  yr -1 , plus unfertilised control) to evaluate their effects on N 2 O, CH 4 and CO 2 soil fluxes in a semiarid shrubland in central Spain. After 8 years of experimental fertilisation, increasing N availability led to a consistent increase in N 2 O emissions, likely due to simultaneous increases in soil microbial nitrification and/or denitrification processes. However, only intermediate levels of N fertilisation reduced CH 4 uptake, while increasing N fertilisation had no effects on CO 2 fluxes, suggesting complex interactions between N deposition loads and GHG fluxes. Our study provides novel insight into the responses of GHGs to N deposition in drylands, forecasting increases in N 2 O emissions, and decreases in CH 4 uptake rates, with likely consequences to the on-going climate change., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier B.V. All rights reserved.)

Microbial processing of aggregate-unprotected organic matter inputs is key for soil fertility, long-term ecosystem carbon and nutrient sequestration and sustainable agriculture. We investigated the effects of adding multiple nutrients (nitrogen, phosphorus and potassium plus nine essential macro- and micro-nutrients) on decomposition and biochemical transformation of standard plant materials buried in 21 grasslands from four continents. Addition of multiple nutrients weakly but consistently increased decomposition and biochemical transformation of plant remains during the peak-season, concurrent with changes in microbial exoenzymatic activity. Higher mean annual precipitation and lower mean annual temperature were the main climatic drivers of higher decomposition rates, while biochemical transformation of plant remains was negatively related to temperature of the wettest quarter. Nutrients enhanced decomposition most at cool, high rainfall sites, indicating that in a warmer and drier future fertilized grassland soils will have an even more limited potential for microbial processing of plant remains., (© 2020 John Wiley & Sons Ltd.)

The availability of metallic nutrients in dryland soils, many of which are essential for the metabolism of soil organisms and vascular plants, may be altered due to climate change-driven increases in aridity. Biocrusts, soil surface communities dominated by lichens, bryophytes and cyanobacteria, are ecosystem engineers known to exert critical functions in dryland ecosystems. However, their role in regulating metallic nutrient availability under climate change is uncertain. Here, we evaluated whether well-developed biocrusts modulate metallic nutrient availability in response to 7 years of experimental warming and rainfall reduction in a Mediterranean dryland located in southeastern Spain. We found increases in the availability of K, Mg, Zn and Na under warming and rainfall exclusion. However, the presence of a well-developed biocrust cover buffered these effects, most likely because its constituents can uptake significant quantities of available metallic nutrients. Our findings suggest that biocrusts, a biotic community prevalent in drylands, exert an important role in preserving and protecting metallic nutrients in dryland soils from leaching and erosion. Therefore, we highlight the need to protect them to mitigate undesired effects of soil degradation driven by climate change in this globally expanding biome.

Atmospheric carbon dioxide enrichment (eCO 2 ) can enhance plant carbon uptake and growth 1-5 , thereby providing an important negative feedback to climate change by slowing the rate of increase of the atmospheric CO 2 concentration 6 . Although evidence gathered from young aggrading forests has generally indicated a strong CO 2 fertilization effect on biomass growth 3-5 , it is unclear whether mature forests respond to eCO 2 in a similar way. In mature trees and forest stands 7-10 , photosynthetic uptake has been found to increase under eCO 2 without any apparent accompanying growth response, leaving the fate of additional carbon fixed under eCO 2 unclear 4,5,7-11 . Here using data from the first ecosystem-scale Free-Air CO 2 Enrichment (FACE) experiment in a mature forest, we constructed a comprehensive ecosystem carbon budget to track the fate of carbon as the forest responded to four years of eCO 2 exposure. We show that, although the eCO 2 treatment of +150 parts per million (+38 per cent) above ambient levels induced a 12 per cent (+247 grams of carbon per square metre per year) increase in carbon uptake through gross primary production, this additional carbon uptake did not lead to increased carbon sequestration at the ecosystem level. Instead, the majority of the extra carbon was emitted back into the atmosphere via several respiratory fluxes, with increased soil respiration alone accounting for half of the total uptake surplus. Our results call into question the predominant thinking that the capacity of forests to act as carbon sinks will be generally enhanced under eCO 2 , and challenge the efficacy of climate mitigation strategies that rely on ubiquitous CO 2 fertilization as a driver of increased carbon sinks in global forests.

The effects of short-term drought on soil microbial communities remain largely unexplored, particularly at large scales and under field conditions. We used seven experimental sites from two continents (North America and Australia) to evaluate the impacts of imposed extreme drought on the abundance, community composition, richness, and function of soil bacterial and fungal communities. The sites encompassed different grassland ecosystems spanning a wide range of climatic and soil properties. Drought significantly altered the community composition of soil bacteria and, to a lesser extent, fungi in grasslands from two continents. The magnitude of the fungal community change was directly proportional to the precipitation gradient. This greater fungal sensitivity to drought at more mesic sites contrasts with the generally observed pattern of greater drought sensitivity of plant communities in more arid grasslands, suggesting that plant and microbial communities may respond differently along precipitation gradients. Actinobateria, and Chloroflexi, bacterial phyla typically dominant in dry environments, increased their relative abundance in response to drought, whereas Glomeromycetes, a fungal class regarded as widely symbiotic, decreased in relative abundance. The response of Chlamydiae and Tenericutes, two phyla of mostly pathogenic species, decreased and increased along the precipitation gradient, respectively. Soil enzyme activity consistently increased under drought, a response that was attributed to drought-induced changes in microbial community structure rather than to changes in abundance and diversity. Our results provide evidence that drought has a widespread effect on the assembly of microbial communities, one of the major drivers of soil function in terrestrial ecosystems. Such responses may have important implications for the provision of key ecosystem services, including nutrient cycling, and may result in the weakening of plant-microbial interactions and a greater incidence of certain soil-borne diseases., (© 2018 John Wiley & Sons Ltd.)

Biocrusts are key drivers of the structure and functioning of drylands and are very sensitive to disturbance, including atmospheric nitrogen (N) deposition. We studied the impacts of simulated N deposition on biocrust community composition and soil photosynthetic and photoprotective pigment content after five years of N application in a European semiarid Mediterranean shrubland. The experiment consisted in six experimental blocks with four plots, each receiving 0, 10, 20, or 50 kg NH 4 NO 3 -N ha -1  year -1  + 6-7 kg N ha -1  year -1 background. After 5 years of N application, total lichen cover decreased up to 50% compared to control conditions and these changes were only clearly evident when evaluated from a temporal perspective (i.e. as the percentage of change from the first survey in 2008 to the last survey in 2012). In contrast, moss cover did not change in response to N, suggesting that biocrust community alterations operate via species- and functional group-specific effects. Interestingly, between-year variations in biocrust cover tracked variations in autumnal precipitation, showing that these communities are more dynamic than previously thought. Biocrust species alterations in response to N were, however, often secondary when compared to the role of ecologically relevant drivers such as soil pH and shrub cover, which greatly determined the composition and inter-annual dynamics of the biocrust community. Similarly, cyanobacterial abundance and soil pigment concentration were greatly determined by biotic and abiotic interactions, soil pH for pigments, and organic matter content and shrub cover for cyanobacteria. Biocrusts, and particularly the lichen component, are highly sensitive to N deposition and their responses to pollutant N can be best understood when evaluated from a temporal and multivariate perspective, including impacts mediated by interactions with biotic and abiotic drivers.

Increased atmospheric nitrogen (N) deposition is known to alter ecosystem carbon source-sink dynamics through changes in soil CO 2 fluxes. However, a limited number of experiments have been conducted to assess the effects of realistic N deposition in the Mediterranean Basin, and none of them have explored the effects of N addition on soil respiration (R s ). To fill this gap, we assessed the effects of N supply on R s dynamics in the following two Mediterranean sites: Capo Caccia (Italy), where 30 kg ha -1  year -1 was supplied for 3 years, and El Regajal (Spain), where plots were treated with 10, 20, or 50 kg N ha -1  year -1 for 8 years. Results show a complex, non-linear response of soil respiration (R s ) to N additions with R s overall increasing at Capo Caccia and decreasing at El Regajal. This suggests that the response of R s to N addition depends on dose and duration of N supply, and the existence of a threshold above which the N introduced in the ecosystem can affect the ecosystem's functioning. Soil cover and seasonality of precipitations also play a key role in determining the effects of N on R s as shown by the different responses observed across seasons and in bare soil vs. the soil under canopy of the dominant species. These results show how increasing rates of N addition may influence soil C dynamics in semiarid ecosystems in the Mediterranean Basin and represent a valuable contribution for the understanding and the protection of Mediterranean ecosystems.

Nitrogen (N) deposition due to anthropogenic pollution is a major driver of the global biodiversity loss. We studied the effect of experimental N and phosphorus (P) fertilization (0, 10, 20, and 50 kg N ha -1 year -1 and 14 kg P ha -1  year -1 over the background deposition levels) on plant cover dynamics of a rosemary (Rosmarinus officinalis L.) shrubland after 8 years of nutrient addition in a semiarid Mediterranean ecosystem from Central Spain. We specifically aimed at testing whether N deposition has the potential to influence the observed expanding trend of woody vegetation into areas dominated by grassland, biological soil crusts, and bare soil. Our results show that N addition loads above 10 kg N ha -1 year -1 reverted the cover dynamics of shrubs. Under N addition conditions, N was no longer a limiting nutrient and other elements, especially P and calcium, determined the seasonal growth of young twigs. Interestingly, N fertilization did not inhibit the growth of young shoots; our estimates point to a reduced rosemary leaf lifespan that is driving individuals to death. This may be triggered by long-term accumulation of N compounds in leaves, suggesting the need to consider the old organs and tissues in long-lived perennial plants, where N toxicity effects could be more mediated by accumulation processes. Shrublands are a widely distributed ecosystem type in biodiverse Mediterranean landscapes, where shrubs play a key role as nurse plants. Therefore, the disappearance of shrublands may accelerate the biodiversity loss associated with other global change drivers, hamper the recruitment of seedlings of woody species, and, as a consequence, accelerate desertification.

Plant roots play a crucial role in regulating key ecosystem processes such as carbon (C) sequestration and nutrient solubilisation. Elevated (e)CO 2 is expected to alter the biomass of fine, coarse and total roots to meet increased demand for other resources such as water and nitrogen (N), however, the magnitude and direction of observed changes vary considerably between ecosystems. Here, we assessed how climate and soil properties mediate root responses to eCO 2 by comparing 24 field-based CO 2 experiments across the globe including a wide range of ecosystem types. We calculated response ratios (i.e. effect size) and used structural equation modelling (SEM) to achieve a system-level understanding of how aridity, mean annual temperature and total soil nitrogen simultaneously drive the response of total, coarse and fine root biomass to eCO 2 . Models indicated that increasing aridity limits the positive response of fine and total root biomass to eCO 2 , and that fine (but not coarse or total) root responses to eCO 2 are positively related to soil total N. Our results provide evidence that consideration of factors such as aridity and soil N status is crucial for predicting plant and ecosystem-scale responses to future changes in atmospheric CO 2 concentrations, and thus feedbacks to climate change.

Mediterranean Basin ecosystems, their unique biodiversity, and the key services they provide are currently at risk due to air pollution and climate change, yet only a limited number of isolated and geographically-restricted studies have addressed this topic, often with contrasting results. Particularities of air pollution in this region include high O 3 levels due to high air temperatures and solar radiation, the stability of air masses, and dominance of dry over wet nitrogen deposition. Moreover, the unique abiotic and biotic factors (e.g., climate, vegetation type, relevance of Saharan dust inputs) modulating the response of Mediterranean ecosystems at various spatiotemporal scales make it difficult to understand, and thus predict, the consequences of human activities that cause air pollution in the Mediterranean Basin. Therefore, there is an urgent need to implement coordinated research and experimental platforms along with wider environmental monitoring networks in the region. In particular, a robust deposition monitoring network in conjunction with modelling estimates is crucial, possibly including a set of common biomonitors (ideally cryptogams, an important component of the Mediterranean vegetation), to help refine pollutant deposition maps. Additionally, increased attention must be paid to functional diversity measures in future air pollution and climate change studies to establish the necessary link between biodiversity and the provision of ecosystem services in Mediterranean ecosystems. Through a coordinated effort, the Mediterranean scientific community can fill the above-mentioned gaps and reach a greater understanding of the mechanisms underlying the combined effects of air pollution and climate change in the Mediterranean Basin., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Global environmental change (GEC) is affecting species interactions and causing a rapid decline in biodiversity. In this study, I present a new Ecosystem Disruption Index to quantify the impacts of simulated nitrogen (N) deposition (0, 10, 20, and 50 kg N·ha -1 ·yr -1  + 6-7 kg N·ha -1 ·yr -1 background) on abiotic and biotic ecological interactions. This comparative index is based on pairwise linear and quadratic regression matrices. These matrices, calculated at the N treatment level, were constructed using a range of abiotic and biotic ecosystem constituents: soil pH, shrub cover, and the first component of several separate principal component analyses using soil fertility data (total carbon and N) and community data (annual plants, microorganisms, biocrusts, edaphic fauna) for a total of seven ecosystem constituents. Four years of N fertilization in a semiarid shrubland completely disrupted the network of ecological interactions, with a greater proportional increase in ecosystem disruption at low N addition levels. Biotic interactions, particularly those involving microbes, shrubs, and edaphic fauna, were more prone to be lost in response to N, whereas interactions involving soil properties were more resilient. In contrast, edaphic fauna was the only group directly affected by N addition, with mites and collembolans increasing their abundance with up to 20 kg N·ha -1 ·yr -1 and then decreasing, which supports the idea of higher-trophic-level organisms being more sensitive to disturbance due to more complex links with other ecosystem constituents. Future experimental studies evaluating the impacts of N deposition, and possibly other GEC drivers, on biodiversity and biotic and abiotic interactions may be able to explain results more effectively in the context of ecological networks as a key feature of ecosystem sensitivity., (© 2016 by the Ecological Society of America.)

Climate models predict shifts in the amount, frequency and seasonality of rainfall. Given close links between grassland productivity and rainfall, such changes are likely to have profound effects on the functioning of grassland ecosystems and modify species interactions. Here, we introduce a unique, new experimental platform - DRI-Grass ( D rought and R oot Herbivore I nteractions in a Grass land) - that exposes a south-eastern Australian grassland to five rainfall regimes [Ambient (AMB), increased amount (IA, +50%), reduced amount (RA, -50%), reduced frequency (RF, single rainfall event every 21 days, with total amount unchanged) and summer drought (SD, 12-14 weeks without water, December-March)], and contrasting levels of root herbivory. Incorporation of a belowground herbivore (root-feeding scarabs) addition treatment allows novel investigation of ecological responses to the twin stresses of altered rainfall and root herbivory. We quantified effects of permanently installed rain shelters on microclimate by comparison with outside plots, identifying small shelter effects on air temperature (-0.19°C day, +0.26°C night), soil water content (SWC; -8%) and photosynthetically active radiation (PAR; -16%). Shelters were associated with modest increases in net primary productivity (NPP), particularly during the cool season. Rainfall treatments generated substantial differences in SWC, with the exception of IA; the latter is likely due to a combination of higher transpiration rates associated with greater plant biomass in IA and the low water-holding capacity of the well-drained, sandy soil. Growing season NPP was strongly reduced by SD, but did not respond to the other rainfall treatments. Addition of root herbivores did not affect plant biomass and there were no interactions between herbivory and rainfall treatments in the 1st year of study. Root herbivory did, however, induce foliar silicon-based defenses in Cynodon dactylon and Eragrostis curvula . Rapid recovery of NPP following resumption of watering in SD plots indicates high functional resilience at the site, and may reflect adaptation of the vegetation to historically high variability in rainfall, both within- and between years. DRI-Grass provides a unique platform for understanding how ecological interactions will be affected by changing rainfall regimes and, specifically, how belowground herbivory modifies grassland resistance and resilience to climate extremes.

Nitrogen (N) deposition has doubled the natural N inputs received by ecosystems through biological N fixation and is currently a global problem that is affecting the Mediterranean regions. We evaluated the existing relationships between increased atmospheric N deposition and biogeochemical indicators related to soil chemical factors and cryptogam species across semiarid central, southern, and eastern Spain. The cryptogam species studied were the biocrust-forming species Pleurochaete squarrosa (moss) and Cladonia foliacea (lichen). Sampling sites were chosen in Quercus coccifera (kermes oak) shrublands and Pinus halepensis (Aleppo pine) forests to cover a range of inorganic N deposition representative of the levels found in the Iberian Peninsula (between 4.4 and 8.1 kg N ha(-1) year(-1)). We extended the ambient N deposition gradient by including experimental plots to which N had been added for 3 years at rates of 10, 20, and 50 kg N ha(-1) year(-1). Overall, N deposition (extant plus simulated) increased soil inorganic N availability and caused soil acidification. Nitrogen deposition increased phosphomonoesterase (PME) enzyme activity and PME/nitrate reductase (NR) ratio in both species, whereas the NR activity was reduced only in the moss. Responses of PME and NR activities were attributed to an induced N to phosphorus imbalance and to N saturation, respectively. When only considering the ambient N deposition, soil organic C and N contents were positively related to N deposition, a response driven by pine forests. The PME/NR ratios of the moss were better predictors of N deposition rates than PME or NR activities alone in shrublands, whereas no correlation between N deposition and the lichen physiology was observed. We conclude that integrative physiological measurements, such as PME/NR ratios, measured on sensitive species such as P. squarrosa, can provide useful data for national-scale biomonitoring programs, whereas soil acidification and soil C and N storage could be useful as additional corroborating ecosystem indicators of chronic N pollution.