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Abstract Grasslands cover approximately a third of the Earth’s land surface and account for about a third of terrestrial carbon storage. Yet, we lack strong predictive models of grassland plant biomass, the primary source of carbon in grasslands. This lack of predictive ability may arise from the assumption of linear relationships between plant biomass and the environment and an underestimation of interactions of environmental variables. Using data from 116 grasslands on six continents, we show unimodal relationships between plant biomass and ecosystem characteristics, such as mean annual precipitation and soil nitrogen. Further, we found that soil nitrogen and plant diversity interacted in their relationships with plant biomass, such that plant diversity and biomass were positively related at low levels of nitrogen and negatively at elevated levels of nitrogen. Our results show that it is critical to account for the interactive and unimodal relationships between plant biomass and several environmental variables to accurately include plant biomass in global vegetation and carbon models.

Plants in grasslands navigate a complex landscape of interactions including competition for resources and defense against pathogens. Foliar fungi can suppress plant growth directly through pathogenic interactions, or indirectly via host growth-defense tradeoffs. The exclusion of foliar fungi allows the reallocation of resources from defense to growth and reproduction. In addition, plants also invest photosynthates in rhizodeposition, or root exudates, which play a significant role in shaping the rhizosphere microbial community. However, it remains unclear what impact the exclusion of foliar fungi has on the allocation of resources to rhizodeposition and the composition of the rhizosphere microbial community. Using a 6-year foliar fungicide study in plots planted with 16 species of native prairie plants, we asked whether foliar fungi influence the rhizosphere microbial composition of a common prairie grass (Andropogon gerardii) and a common legume (Lespedeza capatita). We found that foliar fungicide increased aboveground biomass and season-long plant production, but did not alter root biomass, seed production, or rhizosphere microbial diversity. The magnitude of change in aboveground season-long plant production was significantly associated with the magnitude of change in the rhizosphere microbial community in paired foliar fungicide-treated vs. control plots. These results suggest important coupling between foliar fungal infection and plant investment in rhizodeposition to modify the local soil microbial community.

Experiments comparing diploids with polyploids and in single grassland sites show that nitrogen and/or phosphorus availability influences plant growth and community composition dependent on genome size; specifically, plants with larger genomes grow faster under nutrient enrichments relative to those with smaller genomes. However, it is unknown if these effects are specific to particular site localities with speciifc plant assemblages, climates, and historical contingencies. To determine the generality of genome size-dependent growth responses to nitrogen and phosphorus fertilization, we combined genome size and species abundance data from 27 coordinated grassland nutrient addition experiments in the Nutrient Network that occur in the Northern Hemisphere across a range of climates and grassland communities. We found that after nitrogen treatment, species with larger genomes generally increased more in cover compared to those with smaller genomes, potentially due to a release from nutrient limitation. Responses were strongest for C3 grasses and in less seasonal, low precipitation environments, indicating that genome size effects on water-use-efficiency modulates genome size-nutrient interactions. Cumulatively, the data suggest that genome size is informative and improves predictions of species' success in grassland communities.

Plants serve as critical links between above- and below-ground microbial communitites, both influencing and being influenced by microbes in these two realms. Below-ground microbial communities are expected to respond to soil resource environments, which are mediated by the roots of plants that can, in turn, be influenced by the above-ground community of foliar endophytes. For instance, diverse plant communities deposit more, and more diverse, nutrients into the soil, and this deposition is often increased when foliar pathogens are removed. Differences in soil resources can alter soil microbial composition and phenotypes, including inhibitory capacity, resource use, and antibiotic resistance. In this work, we consider plots differing in plant richness and application of foliar fungicide, evaluating consequences on soil resource levels and root-associated Streptomyces phenotypes. Soil carbon, nitrogen, phosphorus, potassium, and organic matter were greater in samples from polyculture than monoculture, yet this increase was surprisingly offset when foliar fungal communities were disrupted. We find that Streptomyces phenotypes varied more between richness plots—with the Streptomyces from polyculture showing lower inhibitory capacity, altered resource-use profiles, and greater antibiotic resistance—than between subplots with/without foliar fungicide. Where foliar fungicide affected phenotypes, it did so differently in polyculture than in monoculture, for instance decreasing niche width and overlap in monoculture while increasing them in polyculture. No differences in phenotype were correlated with soil nutrient levels, suggesting the need for further research looking more closely at soil resource diversity and particular compounds that were found to differ between treatments.

Abstract Plant disease often increases with N, decreases with CO2, and increases as biodiversity is lost (i.e., the dilution effect). Additionally, all these factors can indirectly alter disease by changing host biomass and hence density-dependent disease transmission. Yet over long periods of time as communities undergo compositional changes, these biomass-mediated pathways might fade, intensify, or even reverse in direction. Using a field experiment that has manipulated N, CO2, and species richness for over 20 years, we compared severity of a specialist rust fungus (Puccinia andropogonis) on its grass host (Andropogon gerardii) shortly after the experiment began (1999) and twenty years later (2019). Between these two sampling periods, two decades apart, we found that disease severity consistently increased with N and decreased with CO2. However, the relationship between diversity and disease reversed from a dilution effect in 1999 (more severe disease in monocultures) to an amplification effect in 2019 (more severe disease in mixtures). The best explanation for this reversal centered on host density (i.e., aboveground biomass), which was initially highest in monoculture, but became highest in mixtures two decades later. Thus, the diversity-disease pattern reversed, but disease consistently increased with host biomass. These results highlight the consistency of N and CO2 as drivers of plant disease in the Anthropocene and emphasize the critical role of host biomass—despite potentially variable effects of diversity—for relationships between biodiversity and disease.

Abstract Most species can be coinfected by multiple pathogens that may interact through shared resources (i.e., resource competition) or the host immune system (i.e., apparent competition). Community theory developed for free‐living organisms suggests that coinfecting pathogens can persist if they satisfy the mutual invasion criterion of coexistence, establishing infections in hosts that are already infected. Furthermore, the likelihood of coexistence may depend on host nutrition which can affect shared resources and host immunity. Here, we apply the novel approach of combining a dynamical model and experimental mutual invasibility trials to explore the effects of host nutrient supply on the coexistence of two viral plant pathogens. We focus on among‐pathogen interactions mediated by shared resources. First, we used a model to generate hypotheses about how nitrogen (N) and phosphorus (P) supply rates affect the ability of two plant viruses to invade established infections of the other virus. Then, we experimentally manipulated the N and P supplied to oats (Avena sativa) in a growth chamber experiment and tested mutual invasion of two RNA viral pathogens, BYDV‐PAV and CYDV‐RPV. Nutrient supplies ranged from rates that barely kept hosts alive up to high, but sub‐toxic, rates. Model simulations suggested that the viruses were more likely to invade established infections either when they could replicate at lower N and P concentrations or when plant N and P concentrations increased due to a combination of nutrient supply rates and resident virus nutrient use. In the experiment, each virus successfully invaded hosts infected by the other and had consistent growth rates across N and P supply rates. Our results suggest that BYDV‐PAV and CYDV‐RPV can coexist across a wide range environmental nutrient supply, which is consistent with the high levels of co‐occurrence of these two viruses in field populations.

Abstract Covering approximately 40% of land surfaces, grasslands provide critical ecosystem services that rely on soil organisms. However, the global determinants of soil biodiversity and functioning remain underexplored. In this study, we investigate the drivers of soil microbial and detritivore activity in grasslands across a wide range of climatic conditions on five continents. We apply standardized treatments of nutrient addition and herbivore reduction, allowing us to disentangle the regional and local drivers of soil organism activity. We use structural equation modeling to assess the direct and indirect effects of local and regional drivers on soil biological activities. Microbial and detritivore activities are positively correlated across global grasslands. These correlations are shaped more by global climatic factors than by local treatments, with annual precipitation and soil water content explaining the majority of the variation. Nutrient addition tends to reduce microbial activity by enhancing plant growth, while herbivore reduction typically increases microbial and detritivore activity through increased soil moisture. Our findings emphasize soil moisture as a key driver of soil biological activity, highlighting the potential impacts of climate change, altered grazing pressure, and eutrophication on nutrient cycling and decomposition within grassland ecosystems.

Abstract 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.

Abstract Eutrophication usually impacts grassland biodiversity, community composition, and biomass production, but its impact on the stability of these community aspects is unclear. One challenge is that stability has many facets that can be tightly correlated (low dimensionality) or highly disparate (high dimensionality). Using standardized experiments in 55 grassland sites from a globally distributed experiment (NutNet), we quantify the effects of nutrient addition on five facets of stability (temporal invariability, resistance during dry and wet growing seasons, recovery after dry and wet growing seasons), measured on three community aspects (aboveground biomass, community composition, and species richness). Nutrient addition reduces the temporal invariability and resistance of species richness and community composition during dry and wet growing seasons, but does not affect those of biomass. Different stability measures are largely uncorrelated under both ambient and eutrophic conditions, indicating consistently high dimensionality. Harnessing the dimensionality of ecological stability provides insights for predicting grassland responses to global environmental change.

Abstract Anthropogenic nutrient enrichment and shifts in herbivory can lead to dramatic changes in the composition and diversity of aboveground plant communities. In turn, this can alter seed banks in the soil, which are cryptic reservoirs of plant diversity. Here, we use data from seven Nutrient Network grassland sites on four continents, encompassing a range of climatic and environmental conditions, to test the joint effects of fertilization and aboveground mammalian herbivory on seed banks and on the similarity between aboveground plant communities and seed banks. We find that fertilization decreases plant species richness and diversity in seed banks, and homogenizes composition between aboveground and seed bank communities. Fertilization increases seed bank abundance especially in the presence of herbivores, while this effect is smaller in the absence of herbivores. Our findings highlight that nutrient enrichment can weaken a diversity maintaining mechanism in grasslands, and that herbivory needs to be considered when assessing nutrient enrichment effects on seed bank abundance.

Abstract All multicellular organisms host a diverse microbiome composed of microbial pathogens, mutualists, and commensals, and changes in microbiome diversity or composition can alter host fitness and function. Nonetheless, we lack a general understanding of the drivers of microbiome diversity, in part because it is regulated by concurrent processes spanning scales from global to local. Global-scale environmental gradients can determine variation in microbiome diversity among sites, however an individual host’s microbiome also may reflect its local micro-environment. We fill this knowledge gap by experimentally manipulating two potential mediators of plant microbiome diversity (soil nutrient supply and herbivore density) at 23 grassland sites spanning global-scale gradients in soil nutrients, climate, and plant biomass. Here we show that leaf-scale microbiome diversity in unmanipulated plots depended on the total microbiome diversity at each site, which was highest at sites with high soil nutrients and plant biomass. We also found that experimentally adding soil nutrients and excluding herbivores produced concordant results across sites, increasing microbiome diversity by increasing plant biomass, which created a shaded microclimate. This demonstration of consistent responses of microbiome diversity across a wide range of host species and environmental conditions suggests the possibility of a general, predictive understanding of microbiome diversity.

Abstract The value of large‐scale collaborations for solving complex problems is widely recognized, but many barriers hinder meaningful authorship for all on the resulting multi‐author publications. Because many professional benefits arise from authorship, much of the literature on this topic has focused on cheating, conflict and effort documentation. However, approaches specifically recognizing and creatively overcoming barriers to meaningful authorship have received little attention. We have developed an inclusive authorship approach arising from 15 years of experience coordinating the publication of over 100 papers arising from a long‐term, international collaboration of hundreds of scientists. This method of sharing a paper initially as a storyboard with clear expectations, assignments and deadlines fosters communication and creates unambiguous opportunities for all authors to contribute intellectually. By documenting contributions through this multi‐step process, this approach ensures meaningful engagement by each author listed on a publication. The perception that co‐authors on large authorship publications have not meaningfully contributed underlies widespread institutional bias against multi‐authored papers, disincentivizing large collaborations despite their widely recognized value for advancing knowledge. Our approach identifies and overcomes key barriers to meaningful contributions, protecting the value of authorship even on massively multi‐authored publications.

The insurance hypothesis posits that more diverse communities are more stable through time. Here, the authors show that plant biodiversity reduces the spatial variability of productivity in grassland communities, demonstrating that the insurance hypothesis applies also across space.

Abstract Human activities are altering ecological communities around the globe. Understanding the implications of these changes requires that we consider the composition of those communities. However, composition can be summarized by many metrics which in turn are influenced by different ecological processes. For example, incidence‐based metrics strongly reflect species gains or losses, while abundance‐based metrics are minimally affected by changes in the abundance of small or uncommon species. Furthermore, metrics might be correlated with different predictors. We used a globally distributed experiment to examine variation in species composition within 60 grasslands on six continents. Each site had an identical experimental and sampling design: 24 plots × 4 years. We expressed compositional variation within each site—not across sites—using abundance‐ and incidence‐based metrics of the magnitude of dissimilarity (Bray–Curtis and Sorensen, respectively), abundance‐ and incidence‐based measures of the relative importance of replacement (balanced variation and species turnover, respectively), and species richness at two scales (per plot‐year [alpha] and per site [gamma]). Average compositional variation among all plot‐years at a site was high and similar to spatial variation among plots in the pretreatment year, but lower among years in untreated plots. For both types of metrics, most variation was due to replacement rather than nestedness. Differences among sites in overall within‐site compositional variation were related to several predictors. Environmental heterogeneity (expressed as the CV of total aboveground plant biomass in unfertilized plots of the site) was an important predictor for most metrics. Biomass production was a predictor of species turnover and of alpha diversity but not of other metrics. Continentality (measured as annual temperature range) was a strong predictor of Sorensen dissimilarity. Metrics of compositional variation are moderately correlated: knowing the magnitude of dissimilarity at a site provides little insight into whether the variation is driven by replacement processes. Overall, our understanding of compositional variation at a site is enhanced by considering multiple metrics simultaneously. Monitoring programs that explicitly incorporate these implications, both when designing sampling strategies and analyzing data, will have a stronger ability to understand the compositional variation of systems and to quantify the impacts of human activities.

Abstract Biotic and abiotic factors interact with dominant plants—the locally most frequent or with the largest coverage—and nondominant plants differently, partially because dominant plants modify the environment where nondominant plants grow. For instance, if dominant plants compete strongly, they will deplete most resources, forcing nondominant plants into a narrower niche space. Conversely, if dominant plants are constrained by the environment, they might not exhaust available resources but instead may ameliorate environmental stressors that usually limit nondominants. Hence, the nature of interactions among nondominant species could be modified by dominant species. Furthermore, these differences could translate into a disparity in the phylogenetic relatedness among dominants compared to the relatedness among nondominants. By estimating phylogenetic dispersion in 78 grasslands across five continents, we found that dominant species were clustered (e.g., co‐dominant grasses), suggesting dominant species are likely organized by environmental filtering, and that nondominant species were either randomly assembled or overdispersed. Traits showed similar trends for those sites (

Abstract A growing body of literature links resources of hosts to their risk of infectious disease. Yet most hosts encounter multiple pathogens, and projections of disease risk based on resource availability could be fundamentally wrong if they do not account for interactions among pathogens within hosts. Here, we measured infection risk of grass hosts (Avena sativa) exposed to three naturally co‐occurring viruses either singly or jointly (barley and cereal yellow dwarf viruses [B/CYDVs]: CYDV‐RPV, BYDV‐PAV, and BYDV‐SGV) along experimental gradients of nitrogen and phosphorus supply. We asked whether disease risk (i.e., infection prevalence) differed in single versus co‐inoculations, and whether these differences varied with rates and ratios of nitrogen and phosphorus supply. In single inoculations, the viruses did not respond strongly to nitrogen or phosphorus. However, in co‐inoculations, we detected illustrative cases of 1) resource‐dependent antagonism (lower prevalence of RPV with increasing N; possibly due to competition), 2) resource‐dependent facilitation (higher prevalence of SGV with decreasing N:P; possibly due to immunosuppression), and 3) weak or no interactions within hosts (for PAV). Together, these within‐host interactions created emergent patterns for co‐inoculated hosts, with both infection prevalence and viral richness increasing with the combination of low nitrogen and high phosphorus supply. We demonstrate that knowledge of multiple pathogens is essential for predicting disease risk from host resources and that projections of risk that fail to acknowledge resource‐dependent interactions within hosts could be qualitatively wrong. Expansions of theory from community ecology theory may help anticipate such relationships by linking host resources to diverse pathogen communities.

Anthropogenic nutrient enrichment may drive shifts in soil microbial communities. Here, the authors analyse nitrogen and phosphorus addition effects on soil fungi in a distributed grassland experiment across four continents, finding promotion of pathogens, suppression of mutualists, and no shifts in saprotrophs.

Abstract Pathogen spread rates are determined, in part, by the performance of pathogens under altered environmental conditions and their ability to persist while switching among hosts and vectors. To determine the effects of new conditions (host, vector, and nutrient) on pathogen spread rate, we introduced a vector‐borne viral plant pathogen, Barley Yellow Dwarf Virus PAV (BYDV‐PAV) into hosts, vectors, and host nutrient supplies that it had not encountered for thousands of viral generations. We quantified pathogen prevalence over the course of two serial inoculations under the new conditions. Using individual‐level transmission rates from this experiment, we parameterized a dynamical model of disease spread and projected spread across host populations through a growing season. A change in nutrient conditions (increased supply of phosphorus) reduced viral transmission whereas shifting to a new vector or host species had no effect on infection prevalence. However, the reduction in the new nutrient environment was only temporary; infection prevalence recovered after the second inoculation. Synthesis. These results highlight how robust the pathogen, BYDV‐PAV, is to changes in its biotic and abiotic environment. Our study also highlights the need to quantify longitudinal infection information beyond snapshot assessments to project disease risk for pathogens in new environments.

Eutrophication has been shown to weaken diversity-stability relationships in grasslands, but it is unclear whether the effect depends on scale. Analysing a globally distributed network of grassland sites, the authors show a positive role of beta diversity and spatial asynchrony as drivers of stability but find that nitrogen enrichment weakens the diversity-stability relationships at different spatial scales.

Grassland ecosystems cover around 37% of the ice-free land surface on Earth and have critical socioeconomic importance globally. As in many terrestrial ecosystems, biological dinitrogen (N2) fixation represents an essential natural source of nitrogen (N). The ability to fix atmospheric N2 is limited to diazotrophs, a diverse guild of bacteria and archaea. To elucidate the abiotic (climatic, edaphic), biotic (vegetation), and spatial factors that govern diazotrophic community composition in global grassland soils, amplicon sequencing of the dinitrogenase reductase gene—nifH—was performed on samples from a replicated standardized nutrient [N, phosphorus (P)] addition experiment in 23 grassland sites spanning four continents. Sites harbored distinct and diverse diazotrophic communities, with most of reads assigned to diazotrophic taxa within the Alphaproteobacteria (e.g., Rhizobiales), Cyanobacteria (e.g., Nostocales), and Deltaproteobacteria (e.g., Desulforomonadales) groups. Likely because of the wide range of climatic and edaphic conditions and spatial distance among sampling sites, only a few of the taxa were present at all sites. The best model describing the variation among soil diazotrophic communities at the OTU level combined climate seasonality (temperature in the wettest quarter and precipitation in the warmest quarter) with edaphic (C:N ratio, soil texture) and vegetation factors (various perennial plant covers). Additionally, spatial variables (geographic distance) correlated with diazotrophic community variation, suggesting an interplay of environmental variables and spatial distance. The diazotrophic communities appeared to be resilient to elevated nutrient levels, as 2–4 years of chronic N and P additions had little effect on the community composition. However, it remains to be seen, whether changes in the community composition occur after exposure to long-term, chronic fertilization regimes.

Abstract Host nutrient supply can mediate host–pathogen and pathogen–pathogen interactions. In terrestrial systems, plant nutrient supply is mediated by soil microbes, suggesting a potential role of soil microbes in plant diseases beyond soil‐borne pathogens and induced plant defenses. Long‐term nitrogen (N) enrichment can shift pathogenic and nonpathogenic soil microbial community composition and function, but it is unclear if these shifts affect plant–pathogen and pathogen–pathogen interactions. In a growth chamber experiment, we tested the effect of long‐term N enrichment on infection by Barley Yellow Dwarf Virus (BYDV‐PAV) and Cereal Yellow Dwarf Virus (CYDV‐RPV), aphid‐vectored RNA viruses, in a grass host. We inoculated sterilized growing medium with soil collected from a long‐term N enrichment experiment (ambient, low, and high N soil treatments) to isolate effects mediated by the soil microbial community. We crossed soil treatments with a N supply treatment (low, high) and virus inoculation treatment (mock‐, singly‐, and co‐inoculated) to evaluate the effects of long‐term N enrichment on plant–pathogen and pathogen–pathogen interactions, as mediated by N availability. We measured the proportion of plants infected (i.e., incidence), plant biomass, and leaf chlorophyll content. BYDV‐PAV incidence (0.96) declined with low N soil (to 0.46), high N supply (to 0.61), and co‐inoculation (to 0.32). Low N soil mediated the effect of N supply on BYDV‐PAV: instead of N supply reducing BYDV‐PAV incidence, the incidence increased. Additionally, ambient and low N soil ameliorated the negative effect of co‐inoculation on BYDV‐PAV incidence. BYDV‐PAV infection only reduced chlorophyll when plants were grown with low N supply and ambient N soil. There were no significant effects of long‐term N soil on CYDV‐RPV incidence. Soil inoculant with different levels of long‐term N enrichment had different effects on host–pathogen and pathogen–pathogen interactions, suggesting that shifts in soil microbial communities with long‐term N enrichment may mediate disease dynamics.

Abstract The plant microbiome can affect host function in many ways and characterizing the ecological factors that shape endophytic (microbes living inside host plant tissues) community diversity is a key step in understanding the impacts of environmental change on these communities. Phylogenetic relatedness among members of a community offers a way of quantifying phylogenetic diversity of a community and can provide insight into the ecological factors that shape endophyte microbiomes. We examined the effects of experimental nutrient addition and herbivory exclusion on the phylogenetic diversity of foliar fungal endophyte communities of the grass species Andropogon gerardii at four sites in the Great Plains of the central USA. Using amplicon sequencing, we characterized the effects of fertilization and herbivory on fungal community phylogenetic diversity at spatial scales that spanned within‐host to between sites across the Great Plains. Despite increasing fungal diversity and richness, at larger spatial scales, fungal microbiomes were composed of taxa showing random phylogenetic associations. Phylogenetic diversity did not differ systematically when summed across increasing spatial scales from a few meters within plots to hundreds of kilometers among sites. We observed substantial shifts in composition across sites, demonstrating distinct but similarly diverse fungal communities were maintained within sites across the region. In contrast, at the scale of within leaves, fungal communities tended to be comprised of closely related taxa regardless of the environment, but there were no shifts in phylogenetic composition among communities. We also found that nutrient addition (fertilization) and herbivory have varying effects at different sites. These results suggest that the direction and magnitude of the outcomes of environmental modifications likely depend on the spatial scale considered, and can also be constrained by regional site differences in microbial diversity and composition.

Abstract Biodiversity—both above‐ and belowground—influences multiple functions in terrestrial ecosystems. Yet, it is unclear whether differences in above‐ and belowground species composition (β‐diversity) are associated with differences in multiple ecosystem functions (e.g., spatial turnover in ecosystem function). Here, we partitioned the contributions of above‐ and belowground β‐diversity and abiotic factors (geographic distance, differences in environments) on the spatial turnover of multiple grassland ecosystem functions. We compiled a dataset of plant and soil microbial communities and six indicators of grassland ecosystem functions (i.e., plant aboveground live biomass, plant nitrogen [N], plant phosphorus [P], root biomass, soil total N, and soil extractable P) from 18 grassland sites on four continents contributing to the Nutrient Network experiment. We used Mantel tests and structural equation models to disentangle the relationship between above‐ and belowground β‐diversity and spatial turnover in grassland ecosystem functions. We found that the effects of abiotic factors on the spatial turnover of ecosystem functions were largely indirect through their influences on above‐ and belowground β‐diversity, and that spatial turnover of ecosystem function was more strongly associated with plant β‐diversity than with soil microbial β‐diversity. These results indicate that changes in above‐ and belowground species composition are one mechanism that interacts with environmental change to determine variability in multiple ecosystem functions across spatial scales. As grasslands face global threats from shrub encroachment, conversion to agriculture, or are lost to development, the functions and services they provide will more strongly converge with increased aboveground community homogenization than with soil microbial community homogenization.

Abstract Ecosystems are changing in complex and unpredictable ways, and analysis of these changes is facilitated by coordinated, long‐term research. Meeting diverse societal needs requires an understanding of what populations and communities will be dominant in 20, 50, and 100 yr. This paper is a product of a synthesis effort of the U.S. National Science Foundation funded Long‐Term Ecological Research (LTER) network addressing the LTER core research area of populations and communities. This analysis revealed that each LTER site had at least one compelling story about what their site would look like in 50 or 100 yr. As the stories were prepared, themes emerged, and the stories were grouped into papers along five themes for this special issue: state change, connectivity, resilience, time lags, and cascading effects. This paper addresses the resilience theme and includes stories from the Baltimore (urban), Hubbard Brook (northern hardwood forest), Andrews (temperate rain forest), Moorea (coral reef), Cedar Creek (grassland), and North Temperate Lakes (lakes) sites. The concept of resilience (the capacity of a system to maintain structure and processes in the face of disturbance) is an old topic that has seen a resurgence of interest as the nature and extent of global environmental change have intensified. The stories we present here show the power of long‐term manipulation experiments (Cedar Creek), the value of long‐term monitoring of forests in both natural (Andrews, Hubbard Brook) and urban settings (Baltimore), and insights that can be gained from modeling and/or experimental approaches paired with long‐term observations (North Temperate Lakes, Moorea). Three main conclusions emerge from the analysis: (1) Resilience research has matured over the past 40 yr of the LTER program; (2) there are many examples of high resilience among the ecosystems in the LTER network; (3) there are also many warning signs of declining resilience of the ecosystems we study. These stories highlight the need for long‐term studies to address this complex topic and show how the diversity of sites within the LTER network facilitates the emergence of overarching concepts about this important driver of ecosystem structure, function, services, and futures.

Human behavior (movement, social contacts) plays a central role in the spread of pathogens like SARS-CoV-2. The rapid spread of SARS-CoV-2 was driven by global human movement, and initial lockdown measures aimed to localize movement and contact in order to slow spread. Thus, movement and contact patterns need to be explicitly considered when making reopening decisions, especially regarding return to work. Here, as a case study, we consider the initial stages of resuming research at a large research university, using approaches from movement ecology and contact network epidemiology. First, we develop a dynamical pathogen model describing movement between home and work; we show that limiting social contact, via reduced people or reduced time in the workplace are fairly equivalent strategies to slow pathogen spread. Second, we develop a model based on spatial contact patterns within a specific office and lab building on campus; we show that restricting on-campus activities to labs (rather than labs and offices) could dramatically alter (modularize) contact network structure and thus, potentially reduce pathogen spread by providing a workplace mechanism to reduce contact. Here we argue that explicitly accounting for human movement and contact behavior in the workplace can provide additional strategies to slow pathogen spread that can be used in conjunction with ongoing public health efforts.

Microorganisms mobilize phosphorus (P) in soil by solubilizing bound inorganic P from soil minerals and by mineralizing organic P via phosphatase enzymes. Nitrogen (N) inputs are predicted to increase through human activities and shift plants to be more P limited, increasing the importance of P mobilization processes for plant nutrition. We studied how the relative abundance of P-solubilizing bacteria (PSB), PSB community composition, and phosphatase activity respond to N and P addition (+N, +P, +NP) in grassland soils spanning large biogeographic gradients. The studied soils are located in South Africa, USA, and UK and part of a globally coordinated nutrient addition experiment. We show that the abundance of PSB in the topsoil was reduced by −18% in the N and by −41% in the NP treatment compared to the control. In contrast, phosphatase activity was significantly higher in the N treatment than in the control across all soils. Soil C:P ratio, sand content, pH, and water-extractable P together explained 71% of the variance of the abundance of PSB across all study sites and all treatments. Further, the community of PSB in the N and NP addition treatment differed significantly from the control. Taken together, this study shows that N addition reduced the relative abundance of PSB, altered the PSB community, and increased phosphatase activity, whereas P addition had no impact. Increasing atmospheric N deposition may therefore increase mineralization of organic P and decrease solubilization of bound inorganic P, possibly inducing a switch in the dominant P mobilization processes from P solubilization to P mineralization.

A Correction to this paper has been published: https://doi.org/10.1038/s41467-021-20997-9.

Ecological stoichiometry and resource competition theory both predict that nutrient rates and ratios can alter infectious disease dynamics. Pathogens such as viruses hijack nutrient rich host metabolites to complete multiple steps of their epidemiological cycle. As the synthesis of these molecules requires nitrogen (N) and phosphorus (P), environmental supply rates, and ratios of N and P to hosts can directly limit disease dynamics. Environmental nutrient supplies also may alter virus epidemiology indirectly by changing host phenotype or the dynamics of coinfecting pathogens. We tested whether host nutrient supplies and coinfection control pathogen growth within hosts and transmission to new hosts, either directly or through modifications of plant tissue chemistry (i.e., content and stoichiometric ratios of nutrients), host phenotypic traits, or among-pathogen interactions. We examined two widespread plant viruses (BYDV-PAV and CYDV-RPV) in cultivated oats (Avena sativa) grown along a range of N and of P supply rates. N and P supply rates altered plant tissue chemistry and phenotypic traits; however, environmental nutrient supplies and plant tissue content and ratios of nutrients did not directly alter virus titer. Infection with CYDV-RPV altered plant traits and resulted in thicker plant leaves (i.e., higher leaf mass per area) and there was a positive correlation between CYDV-RPV titer and leaf mass per area. CYDV-RPV titer was reduced by the presence of a competitor, BYDV-PAV, and higher CYDV-RPV titer led to more severe chlorotic symptoms. In our experimental conditions, virus transmission was unaffected by nutrient supply rates, co-infection, plant stoichiometry, or plant traits, although nutrient supply rates have been shown to increase infection and coinfection rates. This work provides a robust test of the role of plant nutrient content and ratios in the dynamics of globally important pathogens and reveals a more complex relationship between within-host virus growth and alterations of plant traits. A deeper understanding of the differential effects of environmental nutrient supplies on virus epidemiology and ecology is particularly relevant given the rapid increase of nutrients flowing into Earth's ecosystems as a result of human activities.

Abstract Managing multiple ecosystem services (ESs) across landscapes presents a central challenge for ecosystem‐based management, because services often exhibit spatiotemporal variation and weak associations with co‐occurring ESs. Further focus on the mechanistic relationships among ESs and their underlying biophysical processes provides greater insight into the causes of variation and covariation among ESs, thus serving as a guide to enhance their supply while preventing adverse outcomes. Here, we used the U.S. Pacific Northwest coastal dune ecosystem to examine how invasive beachgrass management affects three ESs: coastal protection, western snowy plover conservation, and endemic foredune plant conservation. At seven coastal dune habitat restoration areas, we observed spatial variation in the supply of each ES and further identified a tradeoff between western snowy plover conservation and coastal protection. While the ESs were collectively influenced by the invasive beachgrasses and the foredunes they create, the magnitude of the synergies and tradeoffs were influenced by numerous non‐shared drivers, including nearshore geomorphology, changes in foredune shape as a result of restoration, and other management actions irrespective of restoration. Incorporation of these shared and non‐shared drivers into future coastal management planning may reduce tradeoffs among Pacific Northwest dune ESs. With better understanding of ES relationships, it becomes possible to identify management actions that may enhance synergies and mitigate tradeoffs, leading to better decisions for nature and people.