Disturbance regimes and nutrient inputs are changing worldwide, with consequences for the structure and functioning of plant communities. Classical life-history theory predicts that disturbance should shift communities from long-lived perennials toward short-lived annuals, and that nutrient enrichment may amplify this shift. However, these predictions have not been tested experimentally across broad environmental gradients. Here, using a global coordinated grassland experiment spanning 37 sites, we tested how physical disturbance, vegetation removal and shallow soil tillage, and fertilisation reshape annual-perennial balance, and whether disturbance relaxes the climatic limits of annual dominance. Disturbance nearly doubled the proportion of annual species and more than doubled the relative cover of annuals, whereas fertilisation had little influence and did not interact with disturbance. The disturbance-driven shift arose through contrasting pathways: in graminoids and legumes, it reflected the loss of perennial cover, while in forbs, the expansion of annual cover. In the absence of disturbance, annual dominance was restricted to systems with extremely hot and dry summers, but disturbance nearly tripled the extent of climate space in which annuals dominated. By rapidly reassembling after disturbance, annuals may help maintain vegetation cover, but their expansion also signals loss of perennial cover and the long-term ecosystem functions associated with it.

Plants are consumed by a variety of organisms, including herbivores and pathogens, which significantly impact plant biomass, diversity, community composition, and ecosystem functioning. While the impacts of vertebrate herbivores are well established, the effects of consumer groups such as insect herbivores, mollusks, and fungal pathogens on plant communities are less clear and remain understudied in many systems. Existing evidence of how they affect plant biomass, diversity, and community composition is mixed, and most studies have focused on individual consumer groups in isolation. However, different consumer groups interact with each other, directly or indirectly, in ways that alter their impacts on plants, and the consequences of these interactions for plant community structure and ecosystem function remain understudied. Further, consumer impacts vary across environmental gradients and likely depend on abiotic conditions such as climate, soil type, or elevation, and biotic conditions such as plant productivity, diversity, or community composition. Existing studies testing the impacts of invertebrate herbivores and fungal pathogens on plant communities differ substantially in methodology, making generalities across large scales difficult. This calls for experimental approaches that implement standardized protocols across many sites. Here, we introduce and report on the methodology of a novel global research network, The Bug-Network (BugNet), that implements standardized consumer-reduction experiments across 5 continents and 18 countries in diverse, herbaceous- or shrub-dominated ecosystems to investigate: (1) the influence of fungal pathogens, insect herbivores, and mollusks on plant diversity and ecosystem functioning, (2) interactions among these consumer groups, and (3) the abiotic and biotic drivers of context-dependent consumer impacts. BugNet aims to advance a predictive understanding of plant-consumer interactions in order to test fundamental ecological hypotheses and improve predictions of global change impacts on biodiversity and ecosystem functioning.

Ectomycorrhizal fungi (ECM) play a fundamental role in plant-soil carbon and nitrogen cycling in forest ecosystems, yet their influence on soil organic carbon (SOC) sequestration remains underexplored, particularly in process-based models. Here, we develop a Mycorrhizal fungi-mediated Microbial-ENzyme Decomposition (Myc-MEND) model to explore ECM effects on plant carbon fixation and nitrogen uptake. The model was calibrated with biomass of foliage, wood, and roots, and annual net primary productivity from forests in eastern NC, USA. Our findings show that ECM enhance plant nitrogen availability, increasing plant productivity but not directly promoting SOC sequestration. We find that the increased nitrogen provided by ECM to plants decreased plant C: N ratio and led to a 17 % increase in plant photosynthate. This increase in plant quality and quantity increased SOC storage up to 20 %. However, these increases also stimulated saprotrophic microbial activity and extra-cellular enzyme production, resulting in a 14 % decline in SOC, particularly a 19 % reduction in the particulate organic carbon pool. The most influential pathway for SOC stability was the stabilization of recalcitrant mycorrhizal mycelium necromass, which accounts for 36 % of mineral-associated organic carbon (MOC) storage and 31 % of overall SOC accumulation. Although mycorrhizal colonization led to a net 14 % decrease in total SOC storage, it contributed to a 10 % increase in MOC, highlighting its role in enhancing MOC formation. Our simulations demonstrated that ECM influence the microbial carbon pump by lowering plant C:N ratios, reducing microbial carbon use efficiency, and altering plant-soil carbon fluxes. Overall, our results underscore the critical role of ECM in regulating microbial carbon pump mechanisms and their indirect contributions to SOC persistence via MOC formation. By bridging empirical observations and theoretical modeling, this study lays the groundwork for integrating mycorrhizae processes into future research aimed at predicting ecosystem carbon fluxes and assessing their climate change mitigation potential.

Soil microbes drive ecosystem function and play a critical role in how ecosystems respond to global change. Research surrounding soil microbial communities has rapidly increased in recent decades, and substantial data relating to phospholipid fatty acids (PLFAs) and potential enzyme activity have been collected and analysed. However, studies have mostly been restricted to local and regional scales, and their accuracy and usefulness are limited by the extent of accessible data. Here we aim to improve data availability by collating a global database of soil PLFA and potential enzyme activity measurements from 12,258 georeferenced samples located across all continents, 5.1% of which have not previously been published. The database contains data relating to 113 PLFAs and 26 enzyme activities, and includes metadata such as sampling date, sample depth, and soil pH, total carbon, and total nitrogen. This database will help researchers in conducting both global- and local-scale studies to better understand soil microbial biomass and function.

Responses of ecological communities to perturbations are inherently variable because responses of their constituent populations also vary. Species within a single community may show combinations of no response, positive responses, and negative responses to any given perturbation often canceling each other out resulting in small or no signal that the community level. Here we explore the impacts of warming and loss of the dominant species on alpine ecosystems in a global study. We investigate warming and species-loss treatments on population- and community-level dynamics across alpine-plant communities at two elevations in five globally-distributed mountain locations. Communities showed varied responses to treatments; no community showed strong responses to a single treatment. Rather, most sites were influenced by both perturbations. Populations within these communities responded idiosyncratically, suggesting that constituent species are not all equally robust to perturbations even when community-level effects appear weak. Our results highlight the challenge of making general predictions about population- and community-level responses of alpine ecosystems in the face of present and future perturbations. ### Competing Interest Statement The authors have declared no competing interest.

Final report for the FEMC Ecosystem Monitoring Funded project.

High elevation and latitude ecosystems are experiencing high levels of anthropogenic atmospheric warming. Climate warming may directly change soil microbial activity and alter ecosystem carbon dynamics and productivity, but increasing evidence suggests these responses may depend on other biotic factors such as plant community composition and abiotic factors such as moisture. We examined how abiotic (warming) and biotic (presence of dominant plant species) factors interact to affect soil microbial processes. Our experiment deployed the independent and combined treatments of experimental warming and dominant plant species removal in a high and low elevation montane meadows. We analysed multiple soil microbial responses to warming and the presence of a dominant plant at three times throughout the growing season, including soil respiration, microbial metabolic functional diversity, microbial biomass carbon and nitrogen, and extracellular enzyme potential activity. Overall, there were few independent microbial responses to either the warming or the removal treatments. There was a significant interaction between warming and the removal of a dominant plant species, where microbial biomass and the activity of some microbial enzymes were lower in warmed plots where the dominant species was removed relative to control plots. The effect of warming on extracellular enzyme activity was typically observed only at the high-elevation site. In contrast, we found that effects of warming were consistent across the growing season, despite strong temporal variation in microbial properties. Our results emphasize the need to further consider soil microbial responses to warming under multiple environments, including shifts in both biotic and abiotic factors, to aid in predictions of carbon dynamics under future global change.Read the free Plain Language Summary for this article on the Journal blog. Read the free Plain Language Summary for this article on the Journal blog.image

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.

A growing body of work examines the direct and indirect effects of climate change on ecosystems, typically by using manipulative experiments at a single site or performing meta-analyses across many independent experiments. However, results from single-site studies tend to have limited generality. Although meta-analytic approaches can help overcome this by exploring trends across sites, the inherent limitations in combining disparate datasets from independent approaches remain a major challenge. In this paper, we present a globally distributed experimental network that can be used to disentangle the direct and indirect effects of climate change. We discuss how natural gradients, experimental approaches, and statistical techniques can be combined to best inform predictions about responses to climate change, and we present a globally distributed experiment that utilizes natural environmental gradients to better understand long-term community and ecosystem responses to environmental change. The warming and (species) removal in mountains (WaRM) network employs experimental warming and plant species removals at high- and low-elevation sites in a factorial design to examine the combined and relative effects of climatic warming and the loss of dominant species on community structure and ecosystem function, both above- and belowground. The experimental design of the network allows for increasingly common statistical approaches to further elucidate the direct and indirect effects of warming. We argue that combining ecological observations and experiments along gradients is a powerful approach to make stronger predictions of how ecosystems will function in a warming world as species are lost, or gained, in local communities.

Aim An important research question in ecology is how climate and the biodiversity of aboveground plants and belowground microbiomes affect ecosystem functions such as nutrient pools. However, little is studied on the concurrent role of above- and belowground species composition in shaping the spatial distribution patterns of ecosystem functions across environmental gradients. Here, we investigated the relationships between the taxonomic composition of plants, soil bacteria and soil fungi and spatial turnover in nutrient pools, and assessed how species composition-nutrient pool relationships were mediated by contemporary climatic conditions. Location Qinghai-Tibetan Plateau. Time period Current. Major taxa studied Plants, soil bacteria and soil fungi. Methods We surveyed plant assemblages, sampled the taxonomic composition of soil bacteria and soil fungi, and measured plant- and soil-mediated nutrient pools at 60 alpine grasslands on the Qinghai-Tibetan Plateau. Using Mantel tests, structural equation models and general linear models, we investigated the relative importance of the taxonomic composition of plant, soil bacterial, and soil fungal communities on the spatial turnover of alpine grassland nutrient pools. Results We found that the taxonomic composition of plant, soil bacterial, and soil fungal communities was associated with local climate. However, the effects of local climate on the spatial turnover of plant- and soil-mediated nutrient pools were mainly indirect and mediated through plant and soil bacterial species composition, but not through soil fungal species composition. We further found that the replacement component of soil bacterial beta-diversity and the richness difference of plant beta-diversity were the direct predictors of nutrient pools in the alpine grasslands. Main conclusions These results highlight that belowground bacterial composition together with aboveground plant species composition are related to spatial turnover in nutrient pools, perhaps even driving it. Conserving above- and belowground biodiversity may therefore safeguard against the impacts of local climate on the functions of climate-sensitive alpine grasslands.

Aim A fundamental challenge in soil macroecology is to understand how microbial community structure shapes ecosystem functions along environmental gradients of land surface (i.e., horizontal dimension). However, little is known of microbial community structure-function relationships along environmental gradients of soil depth (i.e., vertical dimension) in extreme environments. A full understanding of the consequences of environmental change for microbial communities structure and subsequent changes in microbial functions could enable more accurate predictions of extreme environmental change effects. Here, we leveraged a 200-km desert soil salinity gradient that is created by a 12-year saline-water irrigation to evaluate how soil microbial community structure-function relationships change with soil salinity in the horizontal and vertical dimensions. Location The Tarim basin of Taklamakan desert. Taxa Soil bacteria and fungi. Methods We assessed the prime ecological processes controlling the assembly of microbial communities and the activity of enzymes relevant to carbon, nitrogen, and phosphorus cycling along soil salinity gradients across study sites (horizontal dimension) and soil depths (vertical dimension) by using the general linear model, hierarchical variance partitioning, and path model. Results Differences in soil depth (on the scale of meters) was as important as geographic distance (on the scale of kilometers) in shaping the structure of bacterial and fungal communities, while both the vertical and horizontal variability in enzymatic activity were largely attributed to the increase in the heterogeneity of soil properties, such as soil texture, water content, and pH. Main conclusions Our results suggest that dispersal limitation and environmental heterogeneity, not soil salinization, along soil depth governs microbial community assembly and enzymatic activity, respectively. This work highlights that conservation efforts of soil macroecology should consider soil depth as a key attribute in the face of ongoing salinization in arid ecosystems.

While numerous studies focus on the ecosystem effects of invasive mammals, few explore the causal mechanisms of such effects. Wild boar is one of the most widely introduced invasive mammal species in the world. By overturning extensive areas of vegetation and soil to feed on belowground resources, wild boar alter the soil food web and thus many microbial-mediated soil processes. Here, we take advantage of a long-term, 8-year, wild boar exclosure experiment across three plant community types in Patagonia, Argentina to explore how wild boar impact soil communities and their potential function. Previous work in this experimental system found that wild boar significantly impacted litter decomposition in the field, but it remained unclear if this effect was mediated through changes in abiotic or biotic soil properties. To explore both the abiotic and biotic drivers of decomposition, we measured soil moisture, soil temperature, soil bulk density, and soil respiration as well as soil micro-arthropod richness and abundance, earthworm abundance, and microbial biomass inside and outside of 10 exclosures in each of three plant community types. To assess potential microbial activity, we measured potential decomposition rates, substrate-induced respiration, and soil microbial enzyme activity. Rooting decreased soil moisture by 18

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.

The loss of aboveground plant diversity alters belowground ecosystem function; yet, the mechanisms underpinning this relationship and the degree to which plant community structure and climate mediate the effects of plant species loss remain unclear. Here, we explored how plant species loss through experimental removal shaped belowground function in ecosystems characterized by different climatic regimes and edaphic properties. We measured plant community composition as well as potential carbon (C) and nitrogen (N) mineralization and microbial extracellular enzyme activity in soils collected from four unique plant removal experiments located along an elevational gradient in Colorado, USA. We found that, regardless of the identity of the removed species or the climate at each site, plant removal decreased the absolute variation in potential N mineralization rates and marginally reduced the magnitude of N mineralization rates. While plant species removal also marginally reduced C mineralization rates, C mineralization, unlike N mineralization, displayed sensitivity to the climatic and edaphic differences among sites, where C mineralization was greatest at the high elevation site that receives the most precipitation annually and contains the largest soil total C pool. Plant removal had little impact on soil enzyme activity. Removal effects were not contingent on the amount of biomass removed annually, and shifts in mineralization rates occurred despite only marginal shifts in plant community structure following plant species removal. Our results present a surprisingly simple and consistent pattern of belowground response to the loss of dominant plant species across an elevational gradient with different climatic and edaphic properties, suggesting a common response of belowground ecosystem function to plant species loss regardless of which plant species are lost or the broader climatic context.

Climate change alters frequencies and intensities of soil drying-rewetting and freezing-thawing cycles. These fluctuations affect soil water availability, a crucial driver of soil microbial activity. While these fluctuations are leaving imprints on soil microbiome structures, the question remains if the legacy of one type of weather fluctuation (e.g., drying-rewetting) affects the community response to the other (e.g., freezing-thawing). As both phenomenons give similar water availability fluctuations, we hypothesized that freezing-thawing and drying-rewetting cycles have similar effects on the soil microbiome. We tested this hypothesis by establishing targeted microcosm experiments. We created a legacy by exposing soil samples to a freezing-thawing or drying-rewetting cycle (phase 1), followed by an additional drying-rewetting or freezing-thawing cycle (phase 2). We measured soil respiration and analyzed soil microbiome structures. Across experiments, larger CO 2 pulses and changes in microbiome structures were observed after rewetting than thawing. Drying-rewetting legacy affected the microbiome and CO 2 emissions upon the following freezing-thawing cycle. Conversely, freezing-thawing legacy did not affect the microbial response to the drying-rewetting cycle. Our results suggest that drying-rewetting cycles have stronger effects on soil microbial communities and CO 2 production than freezing-thawing cycles and that this pattern is mediated by sustained changes in soil microbiome structures.

Abstract The impacts of warming on communities and ecosystems are predicted to be significant in mountain ecosystems because physiological processes, including rates of carbon (C) cycling, are often more temperature‐sensitive in colder environments. Plant biodiversity can also influence C exchange, yet few studies integrate how biotic and abiotic factors may directly or interactively impact ecosystem C flux. Here, we examine the link between simultaneous changes in multiple dimensions of plant diversity and peak growing season ecosystem C uptake across a climatic gradient in the Rocky Mountains, Colorado, USA. We found that taxonomic diversity (species richness), functional diversity (functional evenness), and phylogenetic diversity (mean pairwise distance) were significantly and positively related to peak growing season ecosystem C uptake (i.e., net ecosystem exchange) when considered independently. However, when abiotic and biotic factors were integrated in a structural equation model, only plant phylogenetic diversity was significantly related to C uptake. In addition, we found that actual evapotranspiration (AET—a measure that integrates precipitation and temperature) affected ecosystem C exchange indirectly via its impact on the three dimensions of plant diversity that we examined. These findings highlight complex relationships among key measures of biodiversity and ecosystem C uptake in a rapidly warming ecosystem, and the possible mechanisms that underlie relationships between biodiversity and ecosystem functioning. They also point to the need for integrating multiple dimensions of biodiversity into studies of community and ecosystem ecology.

Ruminant livestock are a significant contributor to global methane emissions. Infectious diseases have the potential to exacerbate these contributions by elevating methane outputs associated with animal production. With the increasing spread of many infectious diseases, the emergence of a vicious climate-livestock-disease cycle is a looming threat.

Aims Biodiversity is often positively related to the capacity of an ecosystem to provide multiple functions simultaneously (i.e. multifunctionality). However, there is some controversy over whether biodiversity-multifunctionality relationships depend on the number of functions considered. Particularly, investigators have documented contrasting findings that the effects of biodiversity on ecosystem multifunctionality do not change or increase with the number of ecosystem functions. Here, we provide some clarity on this issue by examining the statistical underpinnings of different multifunctionality metrics. Methods We used simulations and data from a variety of empirical studies conducted across spatial scales (from local to global) and biomes (temperate and alpine grasslands, forests and drylands). We revisited three methods to quantify multifunctionality including the averaging approach, summing approach and threshold-based approach. Important Findings Biodiversity-multifunctionality relationships either did not change or increased as more functions were considered. These results were best explained by the statistical underpinnings of the averaging and summing multifunctionality metrics. Specifically, by averaging the individual ecosystem functions, the biodiversity-multifunctionality relationships equal the population mean of biodiversitysingle function relationships, and thus will not change with the number of functions. Likewise, by summing the individual ecosystem functions, the strength of biodiversity-multifunctionality relationships increases as the number of functions increased. We proposed a scaling standardization method by converting the averaging or summing metrics into a scaling metric, which would make comparisons among different biodiversity studies. In addition, we showed that the range-relevant standardization can be applied to the threshold-based approach by solving for the mathematical artefact of the approach (i.e. the effects of biodiversity may artificially increase with the number of functions considered). Our study highlights different approaches yield different results and that it is essential to develop an understanding of the statistical underpinnings of different approaches. The standardization methods provide a prospective way of comparing biodiversity-multifunctionality relationships across studies.

Roots release carbon into soil and can alleviate energy limitation of microbial organic matter decomposition. We know little about the effects of roots on microbial decomposition of different organic matter substrates, despite the importance for soil carbon stocks and turnover. Through implementing root-microbe interactions, the Carbon, Organisms, Rhizosphere and Protection in the Soil Environment (CORPSE) model was previously shown to represent dynamics of total soil carbon in temperate forest field experiments. However, the model permits alternative hypotheses concerning microbial-substrate affinity. We investigated how root inputs affect decomposition of soil organic carbon (SOC) with variable decomposability. We simulated SOC stocks in CORPSE and compared microbial degradation of two substrate types with varying root-microbe interactions under two alternative hypotheses that varied in microbial-substrate affinity. We compared our modelled hypotheses to a forest field experiment where we quantified decomposition of isotopically labelled starch and leaf tissues in soils with manipulated root access to microbes. We tested the hypothesis that decomposition of leaves would be more sensitive to root inputs than decomposition of starch, corresponding to the alternative model hypotheses. In the field study, leaf decomposition increased with root density, whereas starch decomposition was unchanged by root density. Microbial biomass and enzyme activity consistently increased with root inputs in CORPSE and the field study. Our field experiment supported the CORPSE simulations with high microbial-substrate affinity. Roots stimulated microbial growth and enzyme production, which increased the degradation of more complex substrates such as leaf tissues. Substrates that were easily decomposed, such as starch, may already be degrading at a maximum rate in the absence of rhizosphere influence because their decomposition rate was unchanged by root inputs. We found that the degree to which roots stimulate microbial decomposition depends on the substrate being decomposed, and that root-microbe interactions influenced SOC stocks in both our model and field experiment. Environmental changes that alter root-microbe interactions could, therefore, alter soil C stocks and biogeochemical cycling, and models of these interactions should incorporate differential influence of rhizosphere inputs on different substrates. A free Plain Language Summary can be found within the Supporting Information of this article.