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Rising atmospheric carbon dioxide concentration ([CO2]) should stimulate biomass production directly via biochemical stimulation of carbon assimilation and indirectly via water savings caused by increased plant water use efficiency. Because of these water savings, the CO2 fertilisation effect should be stronger in drier sites, yet large differences among experiments in grassland biomass response to elevated CO2 appear unrelated to annual precipitation, preventing useful generalisations. Here we show that, as predicted, the impact of elevated CO2 on biomass production in 19 globally-distributed temperate grassland experiments reduces as mean precipitation in seasons other than spring increases but, unexpectedly, rises as mean spring precipitation increases. Moreover, because sites with high spring precipitation also tend to have high precipitation at other times, these effects of spring and non-spring precipitation on the CO2 response offset each other, constraining the response of ecosystem productivity to rising CO2. This explains why previous analyses were unable to discern a reliable trend between site dryness and the CO2 fertilisation effect. Thus, the CO2 fertilisation effect in temperate grasslands worldwide will be constrained by their natural rainfall seasonality such that the stimulation of biomass by rising CO2 could be substantially less than anticipated.

The extent to which ectomycorrhizal (ECM) fungi enable plants to access organic nitrogen (N) bound in soil organic matter (SOM) and transfer this growth-limiting nutrient to their plant host, has important implications for our understanding of plant-fungal interactions, and the cycling and storage of carbon (C) and N in terrestrial ecosystems. Empirical evidence currently supports a range of perspectives, suggesting that ECM vary in their ability to provide their host with N bound in SOM, and that this capacity can both positively and negatively influence soil C storage. To help resolve the multiplicity of observations, we gathered a group of researchers to explore the role of ECM fungi in soil C dynamics, and propose new directions that hold promise to resolve competing hypotheses and contrasting observations. In this Viewpoint, we summarize these deliberations and identify areas of inquiry that hold promise for increasing our understanding of these fundamental and widespread plant symbionts and their role in ecosystem-level biogeochemistry.

Rising atmospheric carbon dioxide concentration should stimulate biomass production directly via biochemical stimulation of carbon assimilation, and indirectly via water savings caused by increased plant water-use efficiency. Because of these water savings, the CO

Shrubs commonly form islands of fertility and are expanding their distribution and dominance in the arctic due to climate change, yet how soil properties may be influenced when different species of shrubs expand under warmer climates remains less explored. Important plant traits, such as their associated root community, are linked to functionally different and dominant shrub species in the arctic and these traits likely shape biogeochemical cycling in areas of shrub expansion. Using an elevational gradient as a proxy for warming, we explored how biochemical processes beneath two important arctic shrubs varied under warmer (low elevation) and cooler (high elevation) climates. Interestingly, the influence of elevation on biogeochemistry varied between the two shrubs. At the low elevation, Betula nana L., an ectomycorrhizal shrub, had high carbon (C) degrading enzyme activities, and relatively low potential net nitrogen (N) mineralization rates. Conversely, Empetrum nigrum ssp. hermaphroditum Hagerup, an cricoid mycorrhizal dwarf-shrub, had higher enzyme activities and net N immobilization rates at the higher elevation. Further, E. nigrum ssp. hermpahroditum appeared to have a more closed C and nutrient cycle than B. nana-enzymes degrading C, N, and phosphorus were tightly correlated with each other and with total C and ammonium concentrations in the humus beneath E. nigrum ssp. hermaphroditum, but not beneath B. nana. Our results suggest differences in the warming responses of C and N cycling beneath shrub species across an arctic tundra landscape.

Extreme drought is increasing in frequency and intensity in many regions globally, with uncertain consequences for the resistance and resilience of ecosystem functions, including primary production. Primary production resistance, the capacity to withstand change during extreme drought, and resilience, the degree to which production recovers, vary among and within ecosystem types, obscuring generalized patterns of ecological stability. Theory and many observations suggest forest production is more resistant but less resilient than grassland production to extreme drought; however, studies of production sensitivity to precipitation variability indicate that the processes controlling resistance and resilience may be influenced more by mean annual precipitation (MAP) than ecosystem type. Here, we conducted a global meta-analysis to investigate primary production resistance and resilience to extreme drought in 64 forests and grasslands across a broad MAP gradient. We found resistance to extreme drought was predicted by MAP; however, grasslands (positive) and forests (negative) exhibited opposing resilience relationships with MAP. Our findings indicate that common plant physiological mechanisms may determine grassland and forest resistance to extreme drought, whereas differences among plant residents in turnover time, plant architecture, and drought adaptive strategies likely underlie divergent resilience patterns. The low resistance and resilience of dry grasslands suggests that these ecosystems are the most vulnerable to extreme drought - a vulnerability that is expected to compound as extreme drought frequency increases in the future. (C) 2018 Elsevier B.V. All rights reserved.

Ecosystems containing multiple nonnative plant species are common, but mechanisms promoting their co-occurrence are understudied. Plant-soil interactions contribute to the dominance of singleton species in nonnative ranges because many nonnatives experience stronger positive feedbacks relative to co-occurring natives. Plant-soil interactions could impede other nonnatives if an individual nonnative benefits from its soil community to a greater extent than its neighboring nonnatives, as is seen with natives. However, plant-soil interactions could promote nonnative co-occurrence if a nonnative accumulates beneficial soil mutualists that also assist other nonnatives. Here, we use greenhouse and field experiments to ask whether plant-soil interactions (1) promote the codominance of two common nonnative shrubs (Ligustrum sinense and Lonicera maackii) and (2) facilitate the invasion of a less-common nonnative shrub (Rhamnus davurica) in deciduous forests of the southeastern United States. In the greenhouse, we found that two of the nonnatives, L. maackii and R. davurica, performed better in soils conditioned by nonnative shrubs compared to uninvaded forest soils, which. suggests that positive feedbacks among co-occurring nonnative shrubs can promote continued invasion of a site. In both greenhouse and field experiments, we found consistent signals that the codominance of the nonnatives L. sinense and L. maackii may be at least partially explained by the increased growth of L. sinense in L. maackii soils. Overall, significant effects of plant-soil interactions on shrub performance indicate that plant-soil interactions can potentially structure the co-occurrence patterns of these nonnatives.

Effective societal responses to rapid climate change in the Arctic rely on an accurate representation of region-specific ecosystem properties and processes. However, this is limited by the scarcity and patchy distribution of field measurements. Here, we use a comprehensive, geo-referenced database of primary field measurements in 1,840 published studies across the Arctic to identify statistically significant spatial biases in field sampling and study citation across this globally important region. We find that 31% of all study citations are derived from sites located within 50 km of just two research sites: Toolik Lake in the USA and Abisko in Sweden. Furthermore, relatively colder, more rapidly warming and sparsely vegetated sites are under-sampled and under-recognized in terms of citations, particularly among microbiology-related studies. The poorly sampled and cited areas, mainly in the Canadian high-Arctic archipelago and the Arctic coastline of Russia, constitute a large fraction of the Arctic ice-free land area. Our results suggest that the current pattern of sampling and citation may bias the scientific consensuses that underpin attempts to accurately predict and effectively mitigate climate change in the region. Further work is required to increase both the quality and quantity of sampling, and incorporate existing literature from poorly cited areas to generate a more representative picture of Arctic climate change and its environmental impacts.

BackgroundIn light of the biodiversity crisis and our limited ability to explain variation in biodiversity, tools to quantify spatial and temporal variation in biodiversity and its underlying drivers are critically needed. Inspired by the recently published ecospace framework, we developed and tested a sampling design for environmental and biotic mapping. We selected 130 study sites (40 × 40 m) across Denmark using stratified random sampling along the major environmental gradients underlying biotic variation. Using standardized methods, we collected site species data on vascular plants, bryophytes, macrofungi, lichens, gastropods and arthropods. To evaluate sampling efficiency, we calculated regional coverage (relative to the known species number per taxonomic group), and site scale coverage (i.e., sample completeness per taxonomic group at each site). To extend taxonomic coverage to organisms that are difficult to sample by classical inventories (e.g., nematodes and non-fruiting fungi), we collected soil for metabarcoding. Finally, to assess site conditions, we mapped abiotic conditions, biotic resources and habitat continuity.ResultsDespite the 130 study sites only covering a minute fraction (0.0005 %) of the total Danish terrestrial area, we found 1774 species of macrofungi (54 % of the Danish fungal species pool), 663 vascular plant species (42 %), 254 bryophyte species (41 %) and 200 lichen species (19 %). For arthropods, we observed 330 spider species (58 %), 123 carabid beetle species (37 %) and 99 hoverfly species (33 %). Correlations among species richness for taxonomic groups were predominantly positive. Overall, sample coverage was remarkably high across taxonomic groups and sufficient to capture substantial spatial variation in biodiversity across Denmark. This inventory is nationally unprecedented in detail and resulted in the discovery of 143 species with no previous record for Denmark. Comparison between plant OTUs detected in soil DNA and observed plant species confirmed the usefulness of carefully curated environmental DNA-data. Species richness did not correlate well among taxa suggesting differential and complex biotic responses to environmental variation.ConclusionsWe successfully and adequately sampled a wide range of diverse taxa along key environmental gradients across Denmark using an approach that includes multi-taxon biodiversity assessment and ecospace mapping. Our approach is applicable to assessments of biodiversity in other regions and biomes where species are structured along environmental gradient.

Herbivores modify their environment by consuming plant biomass and redistributing materials across the landscape. While small mammalian herbivores, such as rodents, are typically inconspicuous, their impacts on plant community structure and chemistry can be large. We used a small mammal exclosure experiment to explore whether rodents in a southeastern old field directly altered the aboveground plant species composition and chemistry, and indirectly altered the belowground soil community composition and activity. In general, when rodents were excluded, C3 graminoids increased in cover and biomass, contributing toward a shift in plant species composition relative to plots where rodents were present. The plant community chemistry also shifted; plant fiber concentration and carbon : nitrogen were higher, whereas plant nitrogen concentration was lower in exclosure plots relative to access plots. While microbial community enzyme activity increased when rodents were excluded, no significant changes in the fungal : bacterial or potential nitrogen mineralization occurred between treatments. Our results show that rodents can rapidly influence aboveground plant community composition and chemistry, but their influence on belowground processes may require plant inputs to the soil to accumulate over longer periods of time.

Summary 1. Climatic change is altering ecosystem structure and function, especially in the southwestern United States where trees are near their physiological water stress threshold. In pi~ (Pinus edulis-Juniperus monosperma; PJ) woodlands, increased drought is causing differential mortality of pi~ resulting in an ecosystem that is becoming juniper dominated. 2. Using a precipitation manipulation, we assessed how both increased and decreased precipitation altered ecosystem function beneath pi~ and juniper. We predicted that changes in precipitation would alter nitrogen (N) availability and mineralization at the site. Further, we predicted that these responses would differ beneath pi~ and juniper crowns due to plantlevel differences in transpiration and N uptake in response to drought. 3. We found minimal interactions between tree species and the precipitation treatments on N cycling. However, across all years measured, soil nitrate decreased with increasing soil volumetric water content; a pattern that is likely due to reduced turnover in dry plots. In contrast, potential soil net-nitrogen mineralization was reduced in water removal plots relative to water addition plots indicating that nitrogen cycling rates were slower under drought. Tree type also influenced nitrogen dynamics in this woodland. Across all 4 years, soil N availability and potential soil net-mineralization rates were higher in soils beneath pi~ relative to juniper across all treatments. Interestingly, the observed shifts in N cycling were not reflected in the abundance of N in microbial biomass or in ammonia-oxidizing bacteria, which are responsible for nitrification. The observed patterns may be due to increased N leaching from the soil during periods of increased rainfall or due to decreased microbial activity or plant N uptake when conditions are dry. 4. The effect of precipitation change on N cycling may have long-term consequences on the plant community in this semi-arid ecosystem. Nitrogen concentrations are highest in the soil when water availability is low, thus when N concentrations are high, plants and microbes are relatively inactive and unable to use this resource.

In the face of climate change, shifts in genetic structure and composition of terrestrial plant species are occurring worldwide. Because different genotypes of these plant species support different soil biota and soil processes, shifts in genetics are likely to have cascading effects on ecosystems. We explore plant genetic effects on soil function in the context of climate change, and selection by soils, soil biota and plant-soil feedbacks. We propose categories of genetically-based plant traits that should be prioritized in research on genetic-based effects on soil processes including plant productivity and C allocation, tissue quality, plant water-use, and rhizosphere mutualisms. Additionally, we posit that soil community responses to climate change should be considered in concert with plant genotype because of sensitivity of soil communities to climate. We use two case studies to highlight these points. We argue that the effects of climate change as an agent of selection on plants may cascade to affect soils, and ultimately the structure, composition and function of ecosystems. Understanding the ecological and evolutionary potential of plant-soil linkages may help us understand and mitigate the extended consequences of global change for ecosystems worldwide. Accordingly, we conclude with experimental approaches for examining genetically-based plant-soil interactions across climate change gradients.

Summary 1. Though co-occurrence of invasive plant species is common, few studies have compared the community and ecosystem impacts of invaders when they occur alone and when they co-occur. Prioritization of invasive species management efforts requires sufficient knowledge of impacts – both among individual invasive species and among different sets of co-occurring invaders – to target resources towards management of sites expected to undergo the largest change. 2. Here, we observed differences in above- and below-ground impacts of two invasive woody shrubs, Lonicera maackii and Ligustrum sinense, among plots containing both shrubs (mixed), each species singly or lacking both species (control). 3. We found additive and non-additive effects of these co-occurring invasives on plant communities and soil processes. Mixed plots contained two times more subdominant invasive plant species than L. maackii or L. sinense plots. Compared to control plots, mixed plots had three times the potential activity of b-glucosidase, a carbon-degrading extracellular soil enzyme. L. maackii plots and mixed plots had less acidic soils, while L. sinense plots had higher soil moisture than control plot soils. Differences in soil properties among plots explained plant- and ground-dwelling arthropod community composition as well as the potential microbial function in soils. 4. Synthesis and applications. Our study highlights the importance of explicitly studying the impacts of co-occurring invasive plant species singly and together. Though Lonicera maackii and Ligustrum sinense have similar effects on ecosystem structure and function when growing alone, our data show that two functionally similar invaders can have non-additive impacts on ecosystems. These results suggest that sites with both species should be prioritized for invasive plant management over sites containing only one of these species. Furthermore, this study provides a valuable template for future studies exploring how and when invasion by co-occurring species alters above- and below-ground function in ecosystems with different traits.

Although herbivores are well known to alter litter inputs and soil nutrient fluxes, their long-term influences on soil development are largely unknown because of the difficulty of detecting and attributing changes in carbon and nutrient pools against large background levels. The early phase of primary succession reduces this signal-to-noise problem, particularly in arid systems where individual plants can form islands of fertility. We used natural variation in tree-resistance to herbivory, and a 15 year herbivore-removal experiment in an Arizona pinon-juniper woodland that was established on cinder soils following a volcanic eruption, to quantify how herbivory shapes the development of soil carbon (C) and nitrogen (N) over 36-54 years (i.e., the ages of the trees used in our study). In this semi-arid ecosystem, trees are widely spaced on the landscape, which allows direct examination of herbivore impacts on the nutrient-poor cinder soils. Although chronic insect herbivory increased annual litterfall N per unit area by 50% in this woodland, it slowed annual tree-level soil C and N accumulation by 111% and 96%, respectively. Despite the reduction in soil C accumulation, short-term litterfall-C inputs and soil C-efflux rates per unit soil surface were not impacted by herbivory. Our results demonstrate that the effects of herbivores on soil C and N fluxes and soil C and N accumulation are not necessarily congruent: herbivores can increase N in litterfall, but over time their impact on plant growth and development can slow soil development. In sum, because herbivores slow tree growth, they slow soil development on the landscape.

International audience; Our basic understanding of plant litter decomposition informs the assumptions underlying widely applied soil biogeochemical models, including those embedded in Earth system models. Confidence in projected carbon cycle-climate feedbacks therefore depends on accurate knowledge about the controls regulating the rate at which plant biomass is decomposed into products such as CO2. Here we test underlying assumptions of the dominant conceptual model of litter decomposition. The model posits that a primary control on the rate of decomposition at regional to global scales is climate (temperature and moisture), with the controlling effects of decomposers negligible at such broad spatial scales. Using a regional-scale litter decomposition experiment at six sites spanning from northern Sweden to southern France-and capturing both within and among site variation in putative controls-we find that contrary to predictions from the hierarchical model, decomposer (microbial) biomass strongly regulates decomposition at regional scales. Furthermore, the size of the microbial biomass dictates the absolute change in decomposition rates with changing climate variables. Our findings suggest the need for revision of the hierarchical model, with decomposers acting as both local- and broad-scale controls on litter decomposition rates, necessitating their explicit consideration in global biogeochemical models.

International audience; Temperature is a primary driver of the distribution of biodiversity as well as of ecosystem boundaries. Declining temperature with increasing elevation in montane systems has long been recognized as a major factor shaping plant community biodiversity, metabolic processes, and ecosystem dynamics. Elevational gradients, as thermoclines, also enable prediction of long-term ecological responses to climate warming. One of the most striking manifestations of increasing elevation is the abrupt transitions from forest to treeless alpine tundra However, whether there are globally consistent above-and belowground responses to these transitions remains an open question. To disentangle the direct and indirect effects of temperature on ecosystem properties, here we evaluate replicate treeline ecotones in seven temperate regions of the world. We find that declining temperatures with increasing elevation did not affect tree leaf nutrient concentrations, but did reduce ground-layer community-weighted plant nitrogen, leading to the strong stoichiometric convergence of ground-layer plant community nitrogen to phosphorus ratios across all regions. Further, elevation-driven changes in plant nutrients were associated with changes in soil organic matter content and quality (carbon to nitrogen ratios) and microbial properties. Combined, our identification of direct and indirect temperature controls over plant communities and soil properties in seven contrasting regions suggests that future warming may disrupt the functional properties of montane ecosystems, particularly where plant community reorganization outpaces treeline advance.

Studies addressing the benefits of “directed dispersal” in ant seed dispersal systems have highlighted the beneficial soil properties of the nests of ants that disperse their seeds. No studies, however, have explored the properties of soils nearby exemplary seed-dispersing ant nests, where recent work indicates that seeds are quickly “redispersed” in eastern North America. To address this, we focused on a forested ecosystem in eastern United States where a keystone seed-dispersing ant, Aphaenogaster rudis, commonly disperses the seeds of numerous understory herbs, including Jeffersonia diphylla. We collected soil cores beneath J. diphylla, around A. rudis nests where seeds are dispersed, and from other forest locations. We analyzed the collected soils for microbial activity using potential soil enzyme activity as a proxy, as well as a number of environmental parameters. We followed this with a glasshouse experiment testing whether the soils collected from near nests, beneath J. diphylla, and from other forested areas altered seedling emergence. We found that microbial activities were higher in near-nest microsites than elsewhere. Specifically, the potential enzyme activities of a carbon-degrading enzyme (β-glucosidase), a phosphorus-acquiring enzyme (phosphatase), and a sulfur-acquiring enzyme (sulfatase) were all significantly higher in areas near ant nests than elsewhere; this same pattern, although not significant, was found for the nitrogen-acquiring enzyme NAGase. No differences were found in other environmental variables we investigated (e.g., soil temperature, soil moisture, soil pH). Our field results indicate that soil biological processes are significantly different in near-nest soils, where the seeds are ultimately dispersed. However, our glasshouse germination trials revealed no enhanced germination in near-nest soils, thereby refuting any near-term advantages of directed dispersal to near-nest locations. Future work should be directed toward addressing whether areas near ant nests provide biologically meaningful escape from seed predation and enhanced establishment, and further characterization of soil microbial communities in such settings.

Molecular innovations in microbial ecology are allowing scientists to correlate microbial community characteristics to a variety of ecosystem functions. However, to date the majority of soil microbial ecology studies target phylogenetic rRNA markers, while a smaller number target functional markers linked to soil processes. We validated a new primer set targeting citrate synthase ( gtl A), a central enzyme in the citric acid cycle linked to aerobic respiration. Primers for a 225 bp fragment suitable for qPCR were tested for specificity and assay performance verified on multiple soils. Clone libraries of the PCR-amplified gtl A gene exhibited high diversity and recovered most major groups identified in a previous 16S rRNA gene study. Comparisons among bacterial communities based on gtlA sequencing using UniFrac revealed differences among the experimental soils studied. Conditions for gtl A qPCR were optimized and calibration curves were highly linear ( R 2 > 0.99) over six orders of magnitude (4.56 × 10 5 to 4.56 × 10 11 copies), with high amplification efficiencies (>1.7). We examined the performance of the gtl A qPCR across a variety of soils and ecosystems, spanning forests, old fields and agricultural areas. We were able to amplify gtl A genes in all tested soils, and detected differences in gtl A abundance within and among environments. These results indicate that a fully developed gtl A-targeted qPCR approach may have potential to link microbial community characteristics with changes in soil respiration.

Linking intraspecific variation in plant traits to ecosystem carbon uptake may allow us to better predict how shift in populations shape ecosystem function. We investigated whether plant populations of a dominant old-field plant species (Solidago altissima) differed in carbon dynamics and if variation in plant traits among genotypes and between populations predicted carbon dynamics. We established a common garden experiment with 35 genotypes from three populations of S. altissima from either Tennessee (southern populations) or Connecticut (northern populations) to ask whether: (1) southern and northern Solidago populations will differ in aboveground productivity, leaf area, flowering time and duration, and whole ecosystem carbon uptake, (2) intraspecific trait variation (growth and reproduction) will be related to intraspecific variation in gross ecosystem CO2 exchange (GEE) and net ecosystem CO2 exchange (NEE) within and between northern and southern populations. GEE and NEE were 4.8× and 2× greater in southern relative to northern populations. Moreover, southern populations produced 13× more aboveground biomass and 1.4× more inflorescence mass than did northern populations. Flowering dynamics (first- and last-day flowering and flowering duration) varied significantly among genotypes in both the southern and northern populations, but plant performance and ecosystem function did not. Both productivity and inflorescence mass predicted NEE and GEE between S. altissima southern and northern populations. Taken together, our data demonstrate that variation between S. altissima populations in performance and flowering traits are strong predictors of ecosystem function in a dominant old-field species and suggest that populations of the same species might differ substantially in their response to environmental perturbations.

Assembly history, including the order in which species arrive into a community, can infl uence long-term community structure; however we know less about how timing of species arrival may alter assembly especially under varying resource conditions. To explore how the timing of species arrival interacts with resource availability to alter community assembly, we constructed experimental plant communities and manipulated the interval between plantings of groups of seedlings (0, 5, 10, 15 or 20 days) at low and high levels of soil nutrient supply. To see if community changes infl uenced ecosystemscale processes, we measured parameters across the plant – soil continuum (e.g. plant biomass and net ecosystem carbon dioxide exchange). We found that the timing of species arrival had a large impact on community assembly, but the size of the eff ect depended on soil fertility. As planting interval increased, plant communities diverged further from the control, but the divergence was stronger at high than at low nutrient supply. Our data suggest that at high nutrient supply, early-planted species preempted light resources more quickly, thus preventing the successful establishment of later arriving species even at short planting intervals. Finally, we found that assembly related divergence in plant communities scaled to impact ecosystem-level characteristics such as green leaf chemistry, but had little eff ect on total community biomass and net ecosystem exchange of CO 2 and water vapor. Our data indicate that the eff ect of a stochastic factor, here the timing of species arrival on community composition, depends on the resource level under which the community assembles.

Background Ectomycorrhizal (ECM) fungi provide one of the main pathways for carbon (C) to move from trees into soils, where these fungi make significant contributions to microbial biomass and soil respiration. Scope ECM fungal species vary significantly in traits that likely influence C sequestration, such that forest C sequestration potential may be driven in part by the existing community composition of ECM fungi. Moreover, accumulating experimental data show that tree genotypes differ in their compatibility with particular ECM fungal species, i.e. mycorrhizal traits of forest trees are heritable. Those traits are genetically correlated with other traits for which tree breeders commonly select, suggesting that selection for traits of interest, such as disease resistance or growth rate, could lead to indirect selection for or against particular mycorrhizal traits of trees in forest plantations. Conclusions Altogether, these observations suggest that selection of particular tree genotypes could alter the community composition of symbiotic ECM fungi in managed forests, with cascading effects on soil functioning and soil C sequestration.

Climate change has the potential to alter both the composition and function of a soil’s microbial community, and interactions among climate change factors may alter soil communities in ways that are not possible to predict from experiments based on a single factor. This study evaluated the direct and interactive effects of three climate change factors—elevated CO 2 , altered amounts of precipitation, and elevated air temperature—on soil microbial communities from an old-field climate change experiment being conducted at Oak Ridge, TN. Soil microbial community composition and biomass were determined by phospholipid fatty acid (PLFA) and neutral lipid fatty acid composition. We found that the interactive effects of precipitation and temperature treatments, as well as the interactive effects of precipitation and CO 2 treatments, had significant impacts on microbial community composition. We found that total soil PLFA concentration, a measure of microbial biomass, was greater in the low-precipitation treatments, especially when low precipitation was combined with ambient CO 2 concentrations or ambient temperature. Ordination analysis indicated that temperature was the most significant predictor of shifts in the soil microbial community composition, explaining approximately 12% of the variance in relative abundance of PLFA biomarkers. The elevated-temperature treatment increased the abundance of Firmicutes (low-guanine–cytosine Gram positive) and decreased the abundance of Gram-negative bacteria. Elevated temperature also reduced the abundance of the arbuscular mycorrhizal fungi PLFA biomarker 16:1π5c and saprophytic fungal PLFA biomarker 18:2π6,9. Overall, our data indicate that the interactions among climate change factors alter the composition of soil microbial communities in old-field ecosystems, suggesting potential for changes in microbial community function under predicted future climate conditions.

Changes in plant species diversity can result in synergistic increases in decomposition rates, while elevated atmospheric CO2 can slow the decomposition rates; yet it remains unclear how diversity and changes in atmospheric CO2 may interact to alter root decomposition. To investigate how elevated CO2 interacts with changes in root-litter diversity to alter decomposition rates, we conducted a 120-day laboratory incubation. Roots from three species (Trifolium repens, Lespedeza cuneata, and Festuca pratense) grown under ambient or elevated CO2 were incubated individually or in combination in soils that were exposed to ambient or elevated CO2 for five years. Our experiment resulted in two main findings: (1) Roots from T. repens and L. cuneata, both nitrogen (N) fixers, grown under elevated CO2 treatments had significantly slower decomposition rates than similar roots grown under ambient CO2 treatments; but the decomposition rate of F. pratense roots (a non-N-fixing species) was similar regardless of CO2 treatment. (2) Roots of the three species grown under ambient CO2 and decomposed in combination with each other had faster decomposition rates than when they were decomposed as single species. However, roots of the three species grown under elevated CO2 had similar decomposition rates when they were incubated alone or in combination with other species. These data suggest that if elevated CO2 reduces the root decomposition rate of even a few species in the community, it may slow root decomposition of the entire plant community.

Nutrient availability and herbivory can regulate primary production in ecosystems, but little is known about how, or whether, they may interact with one another. Here, we investigate how nitrogen availability and insect herbivory interact to alter aboveground and belowground plant community biomass in an old-field ecosystem. In 2004, we established 36 experimental plots in which we manipulated soil nitrogen (N) availability and insect abundance in a completely randomized plot design. In 2009, after 6 years of treatments, we measured aboveground biomass and assessed root production at peak growth. Overall, we found a significant effect of reduced soil N availability on aboveground biomass and belowground plant biomass production. Specifically, responses of aboveground and belowground community biomass to nutrients were driven by reductions in soil N, but not additions, indicating that soil N may not be limiting primary production in this ecosystem. Insects reduced the aboveground biomass of subdominant plant species and decreased coarse root production. We found no statistical interactions between N availability and insect herbivory for any response variable. Overall, the results of 6 years of nutrient manipulations and insect removals suggest strong bottom-up influences on total plant community productivity but more subtle effects of insect herbivores on aspects of aboveground and belowground production.

Elevated atmospheric carbon dioxide concentrations [CO2] is projected to increase forest production, which could increase ecosystem carbon (C) storage. This study contributes to our broad goal of understanding the causes and consequences of increased fine-root production and mortality under elevated [CO2] by examining potential gross nitrogen (N) cycling rates throughout the soil profile. Our study was conducted in a CO2-enriched sweetgum (Liquidambar styraciflua L.) plantation in Oak Ridge, TN, USA. We used 15 N isotope pool dilution methodology to measure potential gross N cycling rates in laboratory incubations of soil from four depth increments to 60cm. Our objectives were twofold: (1) to determine whether N is available for root acquisition in deeper soil and (2) to determine whether elevated [CO2], which has increased inputs of labile C resulting from greater fine-root mortality at depth, has altered N cycling rates. Although gross N fluxes declined with soil depth, we found that N is potentially available for roots to access, especially below 15cm depth where rates of microbial consumption of mineral N were reduced relative to production. Overall, up to 60% of potential gross N mineralization and 100% of potential net N mineralization occurred below 15cm depth at this site. This finding was supported by in situ measurements from ion-exchange resins, where total inorganic N availability at 55cm depth was equal to or greater than N availability at 15cm depth. While it is likely that trees grown under elevated [CO2] are accessing a larger pool of inorganic N by mining deeper soil, we found no effect of elevated [CO2] on potential gross or net N cycling rates. Thus, increased root exploration of the soil volume under elevated [CO2] may be more important than changes in potential gross N cycling rates in sustaining forest responses to rising atmospheric CO2.

Many serious ecosystem consequences of climate change will take decades or even centuries to emerge. Long-term ecological responses to global change are strongly regulated by slow processes, such as changes in species composition, carbon dynamics in soil and by long-lived plants, and accumulation of nutrient capitals. Understanding and predicting these processes require experiments on decadal time scales. But decadal experiments by themselves may not be adequate because many of the slow processes have characteristic time scales much longer than experiments can be maintained. This article promotes a coordinated approach that combines long-term, large-scale global change experiments with process studies and modeling. Long-term global change manipulative experiments, especially in high-priority ecosystems such as tropical forests and high-latitude regions, are essential to maximize information gain concerning future states of the earth system. The long-term experiments should be conducted in tandem with complementary process studies, such as those using model ecosystems, species replacements, laboratory incubations, isotope tracers, and greenhouse facilities. Models are essential to assimilate data from long-term experiments and process studies together with information from long-term observations, surveys, and space-for-time studies along environmental and biological gradients. Future research programs with coordinated long-term experiments, process studies, and modeling have the potential to be the most effective strategy to gain the best information on long-term ecosystem dynamics in response to global change.

The potential for soil carbon (C) sequestration under short-rotation woody crops, like hybrid poplar ( Populus spp.), is a significant uncertainty in our understanding of how managed tree plantations might be used to partially offset increasing atmospheric CO 2 concentrations. Through development of a multi-compartment model, we reviewed information from studies on hybrid poplar and analyzed the potential impact of changes in plant traits and nitrogen (N) fertilization on soil C storage. For a hypothetical setting in the southeastern U.S.A., and starting from soils that are relatively depleted in organic matter (2.5 kg C m −2 ), the model predicted an increase in mineral soil C stocks (1.7 kg C m −2 ) over four 7-year rotations of hybrid poplar. However, at the end of the fourth rotation, both cumulative soil C gains and annual rates of soil C accrual (23–93 g C m −2 yr −1 ) varied widely depending on fertilization rate, biomass yield, and rates of dead root decomposition (three factors that were examined in a factorial model-based experiment). Our analysis indicated that processes linked to genetically modifiable poplar traits (aboveground biomass production, belowground C allocation, root decomposition) are potential controls on soil C sequestration. Key measures of model performance were sensitive to how aboveground biomass production responded to N fertilization. Site specific properties that were independent of plant traits were also important to predicted soil C accumulation and point to possible genotype x site interactions that may explain contradictory data from both empirical and theoretical studies of C sequestration under hybrid poplar plantations.

Long-term ecosystem responses to climate change strongly depend on how the soil subsystem and its inhabitants respond to these perturbations. Using open-top chambers, we studied the response of soil microarthropods to single and combined effects of ambient and elevated atmospheric [CO2], ambient and elevated temperatures and changes in precipitation in constructed old-fields in Tennessee, USA. Microarthropods were assessed five years after treatments were initiated and samples were collected in both November and June. Across treatments, mites and collembola were the most dominant microarthropod groups collected. We did not detect any treatment effects on microarthropod abundance. In November, but not in June, microarthropod richness, however, was affected by the climate change treatments. In November, total microarthropod richness was lower in dry than in wet treatments, and in ambient temperature treatments, richness was higher under elevated [CO2] than under ambient [CO2]. Differential responses of individual taxa to the climate change treatments resulted in shifts in community composition. In general, the precipitation and warming treatments explained most of the variation in community composition. Across treatments, we found that collembola abundance and richness were positively related to soil moisture content, and that negative relationships between collembola abundance and richness and soil temperature could be explained by temperature-related shifts in soil moisture content. Our data demonstrate how simultaneously acting climate change factors can affect the structure of soil microarthropod communities in old-field ecosystems. Overall, changes in soil moisture content, either as direct effect of changes in precipitation or as indirect effect of warming or elevated [CO2], had a larger impact on microarthropod communities than did the direct effects of the warming and elevated [CO2] treatments. Moisture-induced shifts in soil microarthropod abundance and community composition may have important impacts on ecosystem functions, such as decomposition, under future climatic change. © 2010 Elsevier B.V. All rights reserved.

Soil respiration is an important component of the global carbon cycle and is highly responsive to changes in soil temperature and moisture. Accurate prediction of soil respiration and its changes under future climatic conditions requires a clear understanding of the processes involved. Most current empirical soil respiration models incorporate just few of the underlying mechanisms that may influence its response. In this study, a new partially process-based component model that separately treated several source components of soil respiration was tested with data from a climate change experiment that manipulated atmospheric [CO2], air temperature and soil moisture. Results from this model were compared to results from other widely used models with the parameters fitted using experimental data. Using the component model, we were able to estimate the relative proportions of heterotrophic and autotrophic respiration in total soil respiration for each of the different treatments. The value of the Q 10 parameters for temperature response component of all of the models showed sensitivity to soil moisture. Estimated Q 10 parameters were higher for wet treatments and lower for dry treatments compared to the values estimated using either the data from all treatments or from only the control treatments. Our results suggest that process-based models provide a better understanding of soil respiration dynamics under changing environmental conditions, but the extent and contribution of different source components need to be included in mechanistic and process-based soil respiration models at corresponding scales.

Summary • Root carbon (C) inputs may regulate decomposition rates in soil, and in this study we ask: how do labile C inputs regulate decomposition of plant residues, and soil microbial communities? • In a 14 d laboratory incubation, we added C compounds often found in root exudates in seven different concentrations (0, 0.7, 1.4, 3.6, 7.2, 14.4 and 21.7 mg C g )1 soil) to soils amended with and without 13 C-labeled plant residue. We measured CO2 respiration and shifts in relative fungal and bacterial rRNA gene copy numbers using quantitative polymerase chain reaction (qPCR). • Increased labile C input enhanced total C respiration, but only addition of C at low concentrations (0.7 mg C g )1 ) stimulated plant residue decomposition (+2%). Intermediate concentrations (1.4, 3.6 mg C g )1 ) had no impact on plant residue decomposition, while greater concentrations of C (> 7.2 mg C g )1 ) reduced decomposition ()50%). Concurrently, high exudate concentrations (> 3.6 mg C g )1 ) increased fungal and bacterial gene copy numbers, whereas low exudate concentrations (< 3.6 mg C g )1 ) increased metabolic activity rather than gene copy numbers. • These results underscore that labile soil C inputs can regulate decomposition of more recalcitrant soil C by controlling the activity and relative abundance of fungi and bacteria.

Soil fauna can be an important regulator of community parameters and ecosystem processes, but there have been few quantitative syntheses of the role of soil fauna in terrestrial soil communities and ecosystems. Here, we conducted a meta-analysis to investigate the impacts of invertebrate soil micro- and mesofauna (grazers and predators) on plant productivity and microbial biomass. Overall our results indicate that an increase in the biomass of soil fauna increased aboveground plant productivity across ecosystems by 35% and decreased microbial biomass by 8%. In addition, we found no evidence for trophic cascades in terrestrial soil food webs, but the bacterivorous component of soil fauna influenced plant productivity and microbial biomass more than did the fungivorous component. Furthermore, changes in the biomass of soil fauna differentially affected plant productivity among plant functional groups: a higher biomass of soil fauna increased aboveground productivity by 70% in coniferous systems. However, in ecosystems dominated by legumes, a functional group with lower inorganic nitrogen requirements, there was no response of aboveground productivity to increases in the biomass of soil fauna. In sum, the results of this meta-analysis indicate that soil fauna help to regulate ecosystem production, especially in nutrient-limited ecosystems.

Atmospheric and climatic change can alter plant biomass production and plant community composition. However, we know little about how climate change-induced alterations in biomass production affect plant species composition. To better understand how climate change will alter both individual plant species and community biomass, we manipulated atmospheric [CO2], air temperature, and precipitation in a constructed old-field ecosystem. Specifically, we compared the responses of dominant and subdominant species to our climatic treatments, and explored how changes in plant dominance patterns alter community evenness over 2 years. Our study resulted in four major findings: (1) all treatments, elevated [CO2], warming, and increased precipitation increased plant community biomass and the effects were additive rather than interactive, (2) plant species differed in their response to the treatments, resulting in shifts in the proportional biomass of individual species, which altered the plant community composition; however, the plant community response was largely driven by the positive precipitation response of Lespedeza, the most dominant species in the community, (3) precipitation explained most of the variation in plant community composition among treatments, and (4) changes in precipitation caused a shift in the dominant species proportional biomass that resulted in lower community evenness in the wet relative to dry treatments. Interestingly, compositional and evenness responses of the subdominant community to the treatments did not always follow the responses of the whole plant community. Our data suggest that changes in plant dominance patterns and community evenness are an important part of community responses to climatic change, and generally, that such compositional shifts can alter ecosystem biomass production and nutrient inputs.

Plant species can differ in the quantity and quality of leaf litter they produce, and many studies have examined whether plant species diversity affects leaf-litter decomposition and nutrient release. A growing number of studies have indicated that intra-specific variation within plant species can also affect key ecosystem processes. However, the relative importance of intra- versus inter-specific variation for the functioning of ecosystems remains poorly known. Here, we investigate the effects of intra-specific variation in a dominant old-field plant species, tall goldenrod (Solidago altissima), and inter-specific variation among goldenrod species on litter quality, decomposition, and nitrogen (N) release. We found that the nutrient concentration of leaf litter varied among genotypes, which translated into ∼50% difference in decomposition rates. Variation among other goldenrod species in decomposition rate was more than twice that of genetic variation within S. altissima. Furthermore, by manipulating litterbags to contain 1, 3, 6, or 9 genotypes, we found that S. altissima genotype identity had much stronger effects than did genotypic diversity on leaf-litter quality, decomposition, and N release. Taken together, these results suggest that the order of ecological importance for controlling leaf-litter decomposition and N release dynamics is plant species identity⪢genotype identity>genotypic diversity.

Cryptic belowground organisms are difficult to observe and their responses to global changes are not well understood. Nevertheless, there is reason to believe that interactions among above- and belowground communities may mediate ecosystem responses to global change. We used grassland mesocosms to manipulate the abundance of one important group of soil organisms, arbuscular mycorrhizal (AM) fungi, and to study community and ecosystem responses to CO2 and N enrichment. After two growing seasons, biomass responses of plant communities were recorded, and soil community responses were measured using microscopy, phospholipid fatty acids (PLFA) and community-level physiological profiles (CLPP). Ecosystem responses were examined by measuring net primary production (NPP), evapotranspiration, total soil organic matter (SOM), and extractable mineral N. Structural equation modeling was used to examine the causal relationships among treatments and response variables. We found that while CO2 and N tended to directly impact ecosystem functions (evapotranspiration and NPP, respectively), AM fungi indirectly impacted ecosystem functions by strongly influencing the composition of plant and soil communities. For example, the presence of AM fungi had a strong influence on other root and soil fungi and soil bacteria. We found that the mycotrophic status of the dominant plant species in the mesocosms determined whethermore » the presence of AM fungi increased or decreased NPP. Mycotrophic grasses dominated the mesocosm communities during the first growing season, and thus, the mycorrhizal treatments had the highest NPP. In contrast, non-mycotrophic forbs were dominant during the second growing season and thus, the mycorrhizal treatments had the lowest NPP. The composition of the plant community strongly influenced soil N; and the composition of the soil organisms strongly influenced SOM accumulation in the mesocosms. These results show how linkages between above- and belowground communities can determine ecosystem responses to global change.« less

Increased vegetative growth and soil carbon (C) storage under elevated carbon dioxide concentration ([CO 2 ]) has been demonstrated in a number of experiments. However, the ability of ecosystems, either above- or belowground, to maintain increased C storage relies on the response of soil processes, such as those that control nitrogen (N) mineralization, to climatic change. These soil processes are mediated by microbial communities whose activity and structure may also respond to increasing atmospheric [CO 2 ]. We took advantage of a long-term (ca 10 y) CO 2 enrichment experiment in a sweetgum plantation located in the southeastern United States to test the hypothesis that observed increases in root production in elevated relative to ambient CO 2 plots would alter microbial community structure, increase microbial activity, and increase soil nutrient cycling. We found that elevated [CO 2 ] had no detectable effect on microbial community structure using 16S rRNA gene clone libraries, on microbial activity measured with extracellular enzyme activity, or on potential soil N mineralization and nitrification rates. These results support findings at other forested Free Air [CO 2 ] Enrichment (FACE) sites.

Aims The direct effects of atmospheric and climatic change factors— atmospheric [CO2], air temperature and changes in precipitation— can shape plant community composition and alter ecosystem function. It is essential to understand how these factors interact to make better predictions about how ecosystems may respond to change. We investigated the direct and interactive effects of [CO2], warming and altered soil moisture in open-top chambers (OTCs) enclosing a constructed old-field community to test how these factors shape plant communities. Materials and methods The experimental facility in Oak Ridge, TN, USA, made use of 4-m diameter OTCs and rain shelters to manipulate [CO2] (ambient, ambient + 300 ppm), air temperature (ambient, ambient + 3.5C) and soil moisture (wet, dry). The plant communities within the chambers comprised seven common old-field species, including grasses, forbs and legumes. We tracked foliar cover for each species and calculated community richness, evenness and diversity from 2003 to 2005. Important findings This work resulted in three main findings: (1) warming had speciesspecific effects on foliar cover that varied through time and were altered by soil moisture treatments; (2) [CO2] had little effect on individual species or the community; (3) diversity, evenness and richness were influenced most by soil moisture, primarily reflecting the response of one dominant species. We conclude that individualistic species responses to atmospheric and climatic change can alter community composition and that plant populations and communities should be considered as part of analyses of terrestrial ecosystem response to climate change. However, prediction of plant community responses may be difficult given interactions between factors and changes in response through time.

Some single-factor experiments suggest that elevated CO2 concentrations can increase soil carbon, but few experiments have examined the effects of interacting environmental factors on soil carbon dynamics. We undertook studies of soil carbon and nitrogen in a multi-factor (CO2 × temperature × soil moisture) climate change experiment on a constructed old-field ecosystem. After four growing seasons, elevated CO2 had no measurable effect on carbon and nitrogen concentrations in whole soil, particulate organic matter (POM), and mineral-associated organic matter (MOM). Analysis of stable carbon isotopes, under elevated CO2, indicated between 14 and 19% new soil carbon under two different watering treatments with as much as 48% new carbon in POM. Despite significant belowground inputs of new organic matter, soil carbon concentrations and stocks in POM declined over four years under soil moisture conditions that corresponded to prevailing precipitation inputs (1,300 mm yr−1). Changes over time in soil carbon and nitrogen under a drought treatment (approximately 20% lower soil water content) were not statistically significant. Reduced soil moisture lowered soil CO2 efflux and slowed soil carbon cycling in the POM pool. In this experiment, soil moisture (produced by different watering treatments) was more important than elevated CO2 and temperature as a control on soil carbon dynamics.

Amplification of the hydrological cycle as a consequence of global warming is forecast to lead to more extreme intra-annual precipitation regimes characterized by larger rainfall events and longer intervals between events. We present a conceptual framework, based on past investigations and ecological theory, for predicting the consequences of this underappreciated aspect of climate change. We consider a broad range of terrestrial ecosystems that vary in their overall water balance. More extreme rainfall regimes are expected to increase the duration and severity of soil water stress in mesic ecosystems as intervals between rainfall events increase. In contrast, xeric ecosystems may exhibit the opposite response to extreme events. Larger but less frequent rainfall events may result in proportional reductions in evaporative losses in xeric systems, and thus may lead to greater soil water availability. Hydric (wetland) ecosystems are predicted to experience reduced periods of anoxia in response to pr...

Plant–insect interactions can alter ecosystem processes, especially if the insects modify plant architecture, quality, or the quantity of leaf litter inputs. In this study, we investigated the interactions between the rosette gall midge Rhopalomyia solidaginis and tall goldenrod, Solidago altissima, to quantify the degree to which the midge alters plant architecture and how the galls affect rates of litter decomposition and nutrient release in an old-field ecosystem. R. solidaginis commonly leads to the formation of a distinct apical rosette gall on S. altissima and approximately 15% of the ramets in a S. altissima patch were galled (range: 3–34%). Aboveground biomass of galled ramets was 60% higher and the leaf area density was four times greater on galled leaf tissue relative to the portions of the plant that were not affected by the gall. Overall decomposition rate constants did not differ between galled and ungalled leaf litter. However, leaf-litter mass loss was lower in galled litter relative to ungalled litter, which was likely driven by modest differences in initial litter chemistry; this effect diminished after 12 weeks of decomposition in the field. The proportion of N remaining was always higher in galled litter than in ungalled litter at each collection date indicating differential release of nitrogen in galled leaf litter. Several studies have shown that plant–insect interactions on woody species can alter ecosystem processes by affecting the quality or quantity of litter inputs. Our results illustrate how plant–insect interactions in an herbaceous species can affect ecosystem processes by altering the quality and quantity of litter inputs. Given that S. altissima dominates fields and that R. solidaginis galls are highly abundant throughout eastern North America, these interactions are likely to be important for both the structure and function of old-field ecosystems.

Summary 1. It is generally assumed that the same factors drive the decomposition of both litter and roots and that nutrient release from litter and roots is synchronized. However, few studies have explicitly tested these assumptions, and no studies have examined whether plant genetics (i.e. plant susceptibility to herbivory) could affect these relationships. 2. Here we examine the effects of herbivore susceptibility and resistance on needle and fine root litter decomposition of pinon pine, Pinus edulis . The study population consists of individual trees that are either susceptible or resistant to herbivory by the pinon needle scale, Matsucoccus acalyptus , or the stem-boring moth, Dioryctria albovittella . Genetic analyses and long-term experimental removals and additions of these insects to individual trees have identified trees that are naturally resistant or susceptible to M. acalyptus and D. albovittella . In addition, these herbivores increase litter chemical quality and alter soil microclimate, both of which mediate decomposition in ecosystems. 3. The effects of herbivore susceptibility and resistance on needle litter mass and phosphorus (P) loss, when significant, are largely mediated by herbivore-induced changes to microclimate. But the effects of herbivore susceptibility and resistance on root litter nitrogen (N) and P retention, and needle litter N retention, are largely governed by herbivore-induced changes to litter chemical quality. Whether a particular tree was resistant or susceptible to herbivores exerted a large influence on net nutrient release, but the direction of herbivore influence varied temporally. 4. The controls on decomposition vary between herbivore-susceptible and herbivoreresistant phenotypes. This suggests that understanding decomposition and nutrient retention in some ecosystems may require considering the effects of herbivores on aboveand below-ground processes and how these effects may be governed by plant genetics. 5. Synthesis . Because so few studies have attempted to quantify genetic components of ecosystem processes, the integration of ecosystem ecology with population genetics has the potential to place ecosystem science within a genetic and evolutionary framework. Using field trials of known genetic composition, ecosystem scientists may use quantitative genetics techniques to explore ecosystem traits just as population geneticists have used these techniques to explore traditional traits such as resistance to insects.

Herbivores can directly impact ecosystem function by altering litter quality of an ecosystem or indirectly by shifting the composition of microbial communities that mediate nutrient processes. We examined the effects of tree susceptibility and resis- tance to herbivory on litter microarthropod and soil microbial communities to test the general hypothesis that herbivore driven changes in litter inputs and soil microclimate will feedback to the microbial commu- nity. Our study population consisted of individual pinon pine trees that were either susceptible or resistant to the stem-boring moth (Dioryctria albovit- tella) and susceptible pinon pine trees from which the moth herbivores have been manually removed since 1982. Moth herbivory increased pinon litter nitrogen concentrations (16%) and decreased canopy precipi- tation interception (28%), both potentially significant factors influencing litter and soil microbial commu- nities. Our research resulted in three major findings: (1) In spite of an apparent increase in litter quality, herbivory did not change litter microarthropod abun- dance or species richness. (2) However, susceptibility to herbivores strongly influenced bulk soil microbial communities (i.e., 52% greater abundance beneath herbivore-resistant and herbivore-removal trees than susceptible trees) and alkaline phosphatase activity (i.e., 412% increase beneath susceptible trees relative to other groups). (3) Season had a strong influence on microbial communities (i.e., microbial biomass and alkaline phosphatase activity increased after the summer rains), and their response to herbivore inputs, in this semi-arid ecosystem. Thus, during the dry season plant resistance and susceptibility to a common insect herbivore had little or no observable effects on the belowground organisms and processes we studied, but after the rains, some pronounced effects emerged.

Nitrogen (N) deposition is impacting the services that ecosystems provide to humanity. However, the mechanisms determining impacts on the N cycle are not fully understood. To explore the mechanistic underpinnings of N impacts on N cycle processes, we reviewed and synthesised recent progress in ecosystem N research through empirical studies, conceptual analysis and model simulations. Experimental and observational studies have revealed that the stimulation of plant N uptake and soil retention generally diminishes as N loading increases, while dissolved and gaseous losses of N occur at low N availability but increase exponentially and become the dominant fate of N at high loading rates. The original N saturation hypothesis emphasises sequential N saturation from plant uptake to soil retention before N losses occur. However, biogeochemical models that simulate simultaneous competition for soil N substrates by multiple processes match the observed patterns of N losses better than models based on sequential competition. To enable better prediction of terrestrial N cycle responses to N loading, we recommend that future research identifies the response functions of different N processes to substrate availability using manipulative experiments, and incorporates the measured N saturation response functions into conceptual, theoretical and quantitative analyses.

Plant biodiversity is often correlated with ecosystem functioning in terrestrial ecosystems. However, we know little about the relative and combined effects of above- and belowground biodiversity on multiple ecosystem functions (for example, ecosystem multifunctionality, EMF) or how climate might mediate those relationships. Here we tease apart the effects of biotic and abiotic factors, both above- and belowground, on EMF on the Tibetan Plateau, China. We found that a suite of biotic and abiotic variables account for up to 86% of the variation in EMF, with the combined effects of above- and belowground biodiversity accounting for 45% of the variation in EMF. Our results have two important implications: first, including belowground biodiversity in models can improve the ability to explain and predict EMF. Second, regional-scale variation in climate, and perhaps climate change, can determine, or at least modify, the effects of biodiversity on EMF in natural ecosystems., Plant biodiversity often has a positive influence on ecosystem functioning. Here Jing et al. show that belowground diversity can also significantly impact ecosystem multifunctionality, but its relative effect varies by regional-scale climate variation.

Summary � Although many plant communities are invaded by multiple nonnative species, we have limited information on how a species’ origin affects ecosystem function. We tested how differences in species richness and origin affect productivity and seedling establishment. � We created phylogenetically paired native and nonnative plant communities in a glasshouse experiment to test diversity–productivity relationships and responsible mechanisms (i.e. selection or complementarity effects). Additionally, we tested how productivity and associated mechanisms influenced seedling establishment. We used diversity-interaction models to describe how species’ interactions influenced diversity–productivity relationships. � Communities with more species had higher total biomass than did monoculture communities, but native and nonnative communities diverged in root : shoot ratios and the mechanism responsible for increased productivity: positive selection effect in nonnative communities and positive complementarity effect in native communities. Seedling establishment was 46% lower in nonnative than in native communities and was correlated with the average selection effect. Interspecific interactions contributed to productivity patterns, but the specific types of interactions differed between native and nonnative communities. � These results reinforce findings that the diversity–productivity mechanisms in native and nonnative communities differ and are the first to show that these mechanisms can influence seedling establishment and that different types of interactions influence diversity–productivity relationships.

Changes in nutrient inputs due to aboveground herbivory may influence the litter and soil microbial community responsible for processes such as decomposition. The mesophyll-feeding scale insect (Matsucoccus acalyptus) found near Sunset Crater National Monument in northern Arizona, USA significantly increases pinon (Pinus edulis) needle litter nitrogen (N) and phosphorus (P) concentrations by 50%, as well as litter inputs to soil by 21%. Because increases in needle litter quality and quantity of this magnitude should affect the microbial communities responsible for decomposition, we tested the hypothesis that insect herbivory causes a shift in soil microbial and litter microarthropod function. Four major findings result from this research: (1) Despite increases in needle inputs due to herbivory, soil carbon (C) was 56% lower beneath scale-susceptible trees than beneath resistant trees; however, soil moisture, N, and pH were similar among treatments. (2) Microbial biomass was 80% lower in soils beneath scale-susceptible trees when compared to resistant trees in the dry season, while microbial enzyme activities were lower beneath susceptible trees in the wet season. (3) Bacterial community-level physiological profiles differed significantly between susceptible and resistant trees during the dry season but not during the wet season. (4) There was a 40% increase in Oribatida and 23% increase in Prostigmata in susceptible needle litter relative to resistant litter. Despite these changes, the magnitude of microbial biomass, activity, and community structure response to herbivory was lower than expected and appears to take a long time to develop. These results suggest that herbivores impact soils in subtle, but important ways; we suggest that while litter chemistry may strongly mediate soil fertility and microbial communities in mesic ecosystems, the influence is lower than expected in this primary succession xeric ecosystem where season mediates differences in microbial populations. Understanding how insect herbivores alter the distribution of susceptible and resistant trees and their associated decomposer communities in arid environments may lead to better prediction of how these ecosystems respond to climatic change.

Changes in vegetation due to drought-influenced herbivory may influence microclimate in ecosystems. In combination with studies of insect resistant and susceptible trees, we used long-term herbivore removal experiments with two herbivores of pinon (Pinus edulis Endelm.) to test the general hypothesis that herbivore alteration of plant architecture affects soil microclimate, a major driver of ecosystem-level processes. The pinon needle scale (Matsucoccus acalyptus, Herbert) attacks needles of juvenile trees causing them to develop an open crown. In contrast, the stem-boring moth (Dioryctria albovittella Hulst.) kills the terminal shoots of mature trees, causing the crown to develop a dense form. Our studies focused on how the microclimate effects of these architectural changes are likely to accumulate over time. Three patterns emerged: (1) scale herbivory reduced leaf area index (LAI) of susceptible trees by 39%, whereas moths had no effect on LAI; (2) scale herbivory increased soil moisture and temperature beneath susceptible trees by 35 and 26%, respectively, whereas moths had no effect; and (3) scale and moth herbivory decreased crown interception of precipitation by 51 and 29%, respectively. From these results, we conclude: (1) the magnitude of scale effects on soil moisture and temperature is large, similar to global change scenarios, andmore » sufficient to drive changes in ecosystem processes. (2) The larger sizes of moth-susceptible trees apparently buffered them from most microclimate effects of herbivory, despite marked changes in crown architecture. (3) The phenotypic expression of susceptibility or resistance to scale insects extends beyond plant-herbivore interactions to the physical environment.« less