Genotype identity determines productivity and CO2 efflux across a genotype-species gradient of ectomycorrhizal fungi

Ectomycorrhizal (EM) fungal communities are taxonomically diverse, and independent manipulation of both intra- and interspecific diversity has previously been shown to positively influence the productivity and activity of EM fungi. Here, we combine manipulations of intra- and interspecific richness and test the effects of a genotype-species gradient on the biomass production and respiration of EM fungi in vitro. Genotype identity had the most pronounced effect on fungal productivity, and in some cases variation within species was greater than between species. We found small negative effects of both species and genotype richness on biomass production, CO₂ efflux and the final nitrogen (N) content of the fungal communities corresponding to mixed negative selection and complementarity effects. Our study highlights the degree of variability between individual EM fungi at the genotype level, and consequently emphasises the importance of individual genotypes for playing key roles in shaping belowground community functioning.     

Plant species richness, elevated CO2, and atmospheric nitrogen deposition alter soil microbial community composition and function

We determined soil microbial community composition and function in a field experiment in which plant communities of increasing species richness were exposed to factorial elevated CO₂ and nitrogen (N) deposition treatments. Because elevated CO₂ and N deposition increased plant productivity to a greater extent in more diverse plant assemblages, it is plausible that heterotrophic microbial communities would experience greater substrate availability, potentially increasing microbial activity, and accelerating soil carbon (C) and N cycling. We, therefore, hypothesized that the response of microbial communities to elevated CO₂ and N deposition is contingent on the species richness of plant communities. Microbial community composition was determined by phospholipid fatty acid analysis, and function was measured using the activity of key extracellular enzymes involved in litter decomposition. Higher plant species richness, as a main effect, fostered greater microbial biomass, cellulolytic and chitinolytic capacity, as well as the abundance of saprophytic and arbuscular mycorrhizal (AM) fungi. Moreover, the effect of plant species richness on microbial communities was significantly modified by elevated CO₂ and N deposition. For instance, microbial biomass and fungal abundance increased with greater species richness, but only under combinations of elevated CO₂ and ambient N, or ambient CO₂ and N deposition. Cellobiohydrolase activity increased with higher plant species richness, and this trend was amplified by elevated CO₂. In most cases, the effect of plant species richness remained significant even after accounting for the influence of plant biomass. Taken together, our results demonstrate that plant species richness can directly regulate microbial activity and community composition, and that plant species richness is a significant determinant of microbial response to elevated CO₂ and N deposition. The strong positive effect of plant species richness on cellulolytic capacity and microbial biomass indicate that the rates of soil C cycling may decline with decreasing plant species richness.     

Intraspecific Diversity Regulates Fungal Productivity and Respiration

Individuals and not just species are key components of biodiversity, yet the relationship between intraspecific diversity and ecosystem functioning in microbial systems remains largely untested. This limits our ability to understand and predict the effects of altered genetic diversity in regulating key ecosystem processes and functions. Here, we use a model fungal system to test the hypothesis that intraspecific genotypic richness of Paxillus obscurosporus stimulates biomass and CO₂ efflux, but that this is dependent on nitrogen supply. Using controlled experimental microcosms, we show that populations containing several genotypes (maximum 8) of the fungus had greater productivity and produced significantly more CO₂ than those with fewer genotypes. Moreover, intraspecific diversity had a much stronger effect than a four-fold manipulation of the carbon:nitrogen ratio of the growth medium. The effects of intraspecific diversity were underpinned by strong roles of individuals, but overall intraspecific diversity increased the propensity of populations to over-yield, indicating that both complementarity and selection effects can operate within species. Our data demonstrate the importance of intraspecific diversity over a range of nitrogen concentrations, and the need to consider fine scale phylogenetic information of microbial communities in understanding their contribution to ecosystem processes.     

Interactive effects of diversity, nutrients and elevated CO2 on experimental plant communities

The effects of species richness and elevated CO₂ on community productivity under altered nutrient levels were studied in experimental herbaceous communities composed of species from the Midwestern United States annual community, which consists of three functional groups C₃, C₄ and N‐fixers. Aboveground and belowground biomass were measured at flowering stage and at the end of the experiment when fruits of most plants were ripe. At the low nutrient level, species richness did not have a significant effect on community productivity. However, at the high nutrient level, the community biomass decreased with decreasing species richness at both ambient and elevated CO₂ in the first harvest, and at elevated CO₂ in the second harvest. At low nutrient level, CO₂ slightly increased community biomass at medium and high species richness. At high nutrient level, CO₂ significantly increased community biomass in all species‐richness treatments in the first harvest, but a significant response was observed only in the high richness treatment in the second harvest. At the functional group level, biomass of C₃ responded positively to CO₂, and C₄ responded very negatively to CO₂. The N‐fixers responded positively to CO₂ at low and medium species richness, but negatively at high species richness, showing a CO₂×richness interaction. CO₂ increased species evenness in the communities, depending on nutrient level. Species varied in the responses of light‐saturated net photosynthesis (P_max) to elevated CO₂, even within functional groups. Our findings suggest that (1) the relationship between productivity and species diversity was dependent on nutrient levels. (2) Species diversity enhances responses of communities to elevated CO₂. (3) Harvest time can affect the results of diversity‐productivity experiments. (4) Responses of C₃, C₄, and N‐fixers to elevated CO₂ in communities did not follow the prediction based on functional groups or plants grown individually, rather it depended on species richness.     

Effects of macroalgal species identity and richness on primary production in benthic marine communities

Plant biodiversity can enhance primary production in terrestrial ecosystems, but biodiversity effects are largely unstudied in the ocean. We conducted a series of field and mesocosm experiments to measure the relative effects of macroalgal identity and richness on primary productivity (net photosynthetic rate) and biomass accumulation in hard substratum subtidal communities in North Carolina, USA. Algal identity consistently and strongly affected production; species richness effects, although often significent, were subtle. Partitioning of the net biodiversity effect indicated that complementarity effects were always positive and species were usually more productive in mixtures than in monoculture. Surprisingly, slow growing species performed relatively better in the most diverse treatments than the most productive species, thus selection effects were consistently negative. Our results suggest that several basic mechanisms underlying terrestrial plant biodiversity effects also operate in algal-based marine ecosystems, and thus may be general.     

Structure and phylogenetic diversity of post-fire ectomycorrhizal communities of maritime pine

Environmental disturbances define the diversity and assemblage of species, affecting the functioning of ecosystems. Fire is a major disturbance of Mediterranean pine forests. Pines are highly dependent on the ectomycorrhizal (EM) fungal symbiosis, which is critical for tree recruitment under primary succession. To determine the effects of time since fire on the structure and recovery of EM fungal communities, we surveyed the young Pinus pinaster regenerate in three sites differing in the elapsed time after the last fire event. Pine roots were collected, and EM fungi characterized by sequencing the internal transcribed spacer (ITS) and the large subunit (LSU) regions of the nuclear ribosomal (nr)-DNA. The effects of the elapsed time after fire on the EM community structure (richness, presence/absence of fungi, phylogenetic diversity) and on soil properties were analysed. Fungal richness decreased with the elapsed time since the fire; although, the phylogenetic diversity of the EM community increased. Soil properties were different depending on the elapsed time after fire and particularly, the organic matter, carbon-to-nitrogen (C/N) ratio, nitrogen and iron significantly correlated with the assemblage of fungal species. Ascomycetes, particularly Tuberaceae and Pezizales, were significantly over-represented on saplings in the burned site. On seedlings, a significant over-representation of Rhizopogonaceae and Atheliaceae was observed in the most recently burned site, while other fungi (i.e. Cortinariaceae) were significantly under-represented. Our results are consistent with the hypothesis that fire can act as a selective agent by printing a phylogenetic signal on the EM fungal communities associated with naturally regenerated pines, pointing out to some groups as potential fire-adapted fungi.     

Litter species traits, but not richness, contribute to carbon and nitrogen dynamics in an alpine meadow on the Tibetan Plateau

Aims

Litter, as afterlife of plants, plays an important role in driving belowground decomposition processes. Here we tested effects of litter species identity and diversity on carbon (C) and nitrogen (N) dynamics during litter decomposition in N-limited alpine meadow soil from the Qinghai–Tibet Plateau.

Methods

We incubated litters of four meadow species, a sedge (“S”, Kobresia humilis), a grass (“G”, Elymus nutans), a herb (“H”, Saussurea superba), and a legume (“L”, Oxytropis falcata), in monoculture and in mixture with meadow soil. CO₂ release was measured 21 times during the incubation, and soil available N and microbial biomass C and N were measured before and after the experiment.

Results

The organic C decay rate did not differ much among soils amended with monocultures or mixtures of litter, except in the H, S, L, and S+H treatments, which had much higher decay rates. Potential decomposable C pools were lowest in the control, highest in the L treatment, and intermediate in the S treatment. Mineralized N was completely immobilized by soil microbes in all treatments except the control, S+L, and S+G+L treatments. Litter mixtures had both additive and non-additive effects on CO₂-C emission (mainly antagonistic effects), net N mineralization (mainly synergistic), and microbial biomass C and N (both). Overall, these parameters were not significantly correlated with litter species richness. Similarly, microbial C or N was not significantly correlated with litter N content or C/N. However, cumulative CO₂-C emission and net N mineralization were positively correlated with litter N content and negatively correlated with litter C/N.

Conclusions

Litter N content and C/N rather than litter species richness drove the release of CO₂-C and net available N in this ecosystem. The antagonistic effects of litter mixtures contributed to a modest release of CO₂-C, but their synergistic effects enhanced net available N. We suggest that in alpine meadow communities, balancing species with high and low N contents will benefit soil carbon sequestration and plant competition for available N with soil microbes.     

Linking plant diversity–productivity relationships to plant functional traits of dominant species and changes in soil properties in 15‐year‐old experimental grasslands

Positive plant diversity–productivity relationships are known to be driven by complementary resource use via differences in plant functional traits. Moreover, soil properties related to nutrient availability were shown to change with plant diversity over time; however, it is not well‐understood whether and how such plant diversity‐dependent soil changes and associated changes in functional traits contribute to positive diversity–productivity relationships in the long run. To test this, we investigated plant communities of different species richness (1, 2, 6, and 9 species) in a 15‐year‐old grassland biodiversity experiment. We determined community biomass production and biodiversity effects (net biodiversity [NEs], complementarity [CEs], and selection effects [SEs]), as well as community means of plant functional traits and soil properties. First, we tested how these variables changed along the plant diversity gradient and were related to each other. Then, we tested for direct and indirect effects of plant and soil variables influencing community biomass production and biodiversity effects. Community biomass production, NEs, CEs, SEs, plant height, root length density (RLD), and all soil property variables changed with plant diversity and the presence of the dominant grass species Arrhenatherum elatius (increase except for soil pH, which decreased). Plant height and RLD for plant functional traits, and soil pH and organic carbon concentration for soil properties, were the variables with the strongest influence on biomass production and biodiversity effects. Our results suggest that plant species richness and the presence of the dominant species, A. elatius, cause soil organic carbon to increase and soil pH to decrease over time, which increases nutrient availability favoring species with tall growth and dense root systems, resulting in higher biomass production in species‐rich communities. Here, we present an additional process that contributes to the strengthening positive diversity–productivity relationship, which may play a role alongside the widespread plant functional trait‐based explanation.     

Biodiversity effects on productivity and stability of marine macroalgal communities: the role of environmental context

The influence of biodiversity on ecosystem functioning has been the focus of much recent research, but the role of environmental context and the mechanisms by which it may influence diversity effects on production and stability remain poorly understood. We assembled marine macroalgal communities in two mesocosm experiments that varied nutrient supply, and at four field sites that differed naturally in environmental conditions. Concordant with theory, nutrient addition promoted positive species richness effects on algal growth in the first mesocosm experiment; however, it tended to weaken the positive diversity relationship found under ambient conditions in a second experiment the next year. In the field experiments, species richness increased algal biomass production at two of four sites. Together, these experiments indicate that diversity effects on algal biomass production are strongly influenced by environmental conditions that vary over space and time. In decomposing the net biodiversity effect into its component mechanisms, seven of the eight experimental settings showed positive complementarity effects (suggesting facilitation or complementary resource use) countered by negative selection effects (i.e. enhanced growth in mixture of otherwise slow growing species) to varying degrees. Under no conditions, including nutrient enrichment, did we find evidence of positive selection effects commonly thought to drive positive diversity effects. Species richness enhanced stability of algal community biomass across a range of environmental settings in our field experiments. Hence, while species richness can increase production, enhanced stability is also an important functional outcome of maintaining diverse marine macroalgal communities.     

Partitioning the effects of algal species identity and richness on benthic marine primary production

Influential research in terrestrial habitats indicates that several ecosystem processes are related to plant biodiversity, yet these links remain poorly studied in marine ecosystems. We conducted one field and one mesocosm experiment to quantify the relative effects of macroalgal species identity and richness on primary production in coral reef macroalgal communities off the north coast of Jamaica. We measured production as the net accumulation of algal biomass in the absence of consumers and as photosynthetic rate using oxygen probes in sealed aquaria. We used two recently developed techniques to attribute deviations in expected relative yield to components associated with species identity or diversity and then to further partition diversity effects into mechanistic components based on dominance, trait-dependent complementarity, and trait-independent complementarity. Our results indicate that algal identity had far greater effects on absolute net growth and photosynthesis than richness. The most diverse mixture of macroalgae did not outperform the most productive monoculture or the average monoculture in either measure of primary production (i.e. we did not find evidence of either transgressive or non-transgressive overyielding). Trait-independent complementarity effects were positive but dominance and trait-dependent complementarity were both negative and became stronger when richness was increased. Thus the potentially positive influence of species interactions and niche partitioning on production were negated by dominance and other negative selection effects. These results demonstrate that the counteracting influence of component effects can diminish the net richness effects on production. This could explain frequently observed weak net richness effects in other aquatic and terrestrial systems and suggests that life history tradeoffs greatly reduce the potential for ecologically relevant plant biodiversity effects on ecosystem properties.     

Determining the mechanism by which fish diversity influences production

Understanding the ability of biodiversity to govern ecosystem function is essential with current pressures on natural communities from species invasions and extirpations. Changes in fish communities can be a major determinant of food web dynamics, and even small shifts in species composition or richness can translate into large effects on ecosystems. In addition, there is a large information gap in extrapolating results of small-scale biodiversity–ecosystem function experiments to natural systems with realistic environmental complexity. Thus, we tested the key mechanisms (resource complementarity and selection effect) for biodiversity to influence fish production in mesocosms and ponds. Fish diversity treatments were created by replicating species richness and species composition within each richness level. In mesocosms, increasing richness had a positive effect on fish biomass with an overyielding pattern indicating species mixtures were more productive than any individual species. Additive partitioning confirmed a positive net effect of biodiversity driven by a complementarity effect. Productivity was less affected by species diversity when species were more similar. Thus, the primary mechanism driving fish production in the mesocosms was resource complementarity. In the ponds, the mechanism driving fish production changed through time. The key mechanism was initially resource complementarity until production was influenced by the selection effect. Varying strength of intraspecific interactions resulting from differences in resource levels and heterogeneity likely caused differences in mechanisms between the mesocosm and pond experiments, as well as changes through time in the ponds. Understanding the mechanisms by which fish diversity governs ecosystem function and how environmental complexity and resource levels alter these relationships can be used to improve predictions for natural systems.     

Global change belowground: impacts of elevated CO2, nitrogen,and summer drought on soil food webs and biodiversity

The world's ecosystems are subjected to various anthropogenic global change agents, such as enrichment of atmospheric CO₂ concentrations, nitrogen (N) deposition, and changes in precipitation regimes. Despite the increasing appreciation that the consequences of impending global change can be better understood if varying agents are studied in concert, there is a paucity of multi‐factor long‐term studies, particularly on belowground processes. Herein, we address this gap by examining the responses of soil food webs and biodiversity to enrichment of CO₂, elevated N, and summer drought in a long‐term grassland study at Cedar Creek, Minnesota, USA (BioCON experiment). We use structural equation modeling (SEM), various abiotic and biotic explanatory variables, and data on soil microorganisms, protozoa, nematodes, and soil microarthropods to identify the impacts of multiple global change effects on drivers belowground. We found that long‐term (13‐year) changes in CO₂ and N availability resulted in modest alterations of soil biotic food webs and biodiversity via several mechanisms, encompassing soil water availability, plant productivity, and – most importantly – changes in rhizodeposition. Four years of manipulation of summer drought exerted surprisingly minor effects, only detrimentally affecting belowground herbivores and ciliate protists at elevated N. Elevated CO₂ increased microbial biomass and the density of ciliates, microarthropod detritivores, and gamasid mites, most likely by fueling soil food webs with labile C. Moreover, beneficial bottom‐up effects of elevated CO₂ compensated for detrimental elevated N effects on soil microarthropod taxa richness. In contrast, nematode taxa richness was lowest at elevated CO₂ and elevated N. Thus, enrichment of atmospheric CO₂ concentrations and N deposition may result in taxonomically and functionally altered, potentially simplified, soil communities. Detrimental effects of N deposition on soil biodiversity underscore recent reports on plant community simplification. This is of particular concern, as soils house a considerable fraction of global biodiversity and ecosystem functions.     

Does richness of emergent plants affect CO2 and CH4 emissions in experimental wetlands?

相似文献   

Fertilization and plant diversity accelerate primary succession and restoration of dune communities

Plant species richness can increase primary production because plants occupy different niches or facilitate each other (“complementarity effects”) or because diverse mixtures have a greater chance of having more productive species (“selection effects”). To determine how complementarity and selection influence dune restoration, we established four types of plant communities [monocultures of sea oats (Uniola paniculata), bitter panicgrass (Panicum amarum) and saltmeadow cordgrass (Spartina patens) and the three-species mixture] under different soil treatments typical of dune restorations (addition of soil organic material, nutrients, both, or neither). This fully factorial design allowed us to determine if plant identity, diversity and soil treatments influenced the yield of both the planted species and species that recruited naturally (volunteers). Planted species responses in monocultures and mixtures varied among soil treatments. The composition of the plantings and soils also influenced the abundance of volunteers. The mixture of the three species had the lowest cover of volunteers. We also found that the effect of diversity on production increased with fertilizer. We partitioned the biodiversity effect into complementarity and selection effects and found that the increase in the diversity effect occurred because increased nutrients decreased dominance by the largest species and increased complementarity among species. Our findings suggest that different planting schemes can be used to meet specific goals of restoration (e.g., accelerate plant recovery while suppressing colonization of non-planted species).     

Inorganic soil nitrogen under grassland plant communities of different species composition and diversity

We measured aboveground plant biomass and soil inorganic nitrogen pools in a biodiversity experiment in northern Sweden, with plant species richness ranging from 1 to 12 species. In general, biomass increased and nitrate pools decreased with increasing species richness. Transgressive overyielding of mixed plant communities compared to the most productive of the corresponding monocultures occurred in communities with and without legumes. N₂-fixing legumes had a fertilizing function, while non-legumes had a N retaining function. Plant communities with only legumes had a positive correlation between biomass and soil nitrate content, whereas in plant communities without legumes they were negatively correlated. Both nitrate and ammonium soil pools in mixed non-legume communities were approximately equal to the lowest observed in the corresponding monocultures. In mixed legume/non-legume communities, no correlation was found for soil nitrate with either biomass or legume biomass as percentage of total biomass. The idea of complementarity among species in nitrogen acquisition was supported in both pure non-legume and mixed non-legume/legume communities. In the latter, however, facilitation through increased nitrogen availability and retention, was probably dominating. Our results suggest that diversity effects on biomass and soil N pools through resource use complementarity depend on the functional traits of species, especially N₂ fixation or high productivity.     

Consistent Effects of Biodiversity on Ecosystem Functioning Under Varying Density and Evenness

Biodiversity experiments typically vary only species richness and composition, yet the generality of their results relies on consistent effects of these factors even under varying starting conditions of density and evenness. We tested this assumption in a factorial species richness x density x evenness experiment using a pool of 60 common grassland species divided into four functional groups (grasses, legumes, tall herbs and short herbs). Richness varied from 1, 2, 4, 8 to 16 species, total planting density was 1,000 or 2,000 seeds/m², and species were sown in even or uneven proportions, where one functional group was made dominant. Aboveground plant biomass increased linearly with the logarithm of species richness in all density and evenness treatments during all three years of the experiment. This was due to a convergence of realized density and evenness within species richness levels, although functional groups which were initially made dominant retained their dominance. Between species richness levels, realized density increased, and realized evenness decreased with species richness. Thus, more individuals could coexist if they belonged to different species. Within species richness levels, higher biomass values were correlated with lower density, suggesting an underlying thinning process. However, communities with low realized evenness also had low biomass values; thus high biomass did not result from species dominance. So-called complementarity and selection effects were similar across density and evenness treatments, indicating that the mechanisms underpinning the biodiversity effects were not altered. Species richness was the dominant driver of aboveground biomass, irrespective of variations in total densities and species abundance distributions at the start of the experiment; rejecting the hypothesis that initial differences in species abundance distributions might lead to different “stable states” in community structure or biomass. Thus, results from previous biodiversity experiments that only manipulated species richness and composition should be quite robust and broadly generalizable.     

Interactions among mycorrhizae, atmospheric CO2 and soil N impact plant community composition

We examined plant community responses to interactions between arbuscular mycorrhizal (AM) fungi and availability of atmospheric CO₂ and soil N. Communities of 14 plant species were grown in mesocosms containing living or killed AM fungal inoculum, ambient or elevated atmospheric CO₂ and low or enriched soil N. After one growing season, significantly different plant communities existed in the different treatments. Plant species richness was lowest in +N mesocosms and highest in +AM + CO₂ mesocosms. At ambient CO₂, AM fungi reduced richness but at elevated CO₂ they increased it. This was caused by changes in mortality rates of several C₃ forbs and may suggest that CO₂ enrichment ameliorates the carbon cost of some AM symbioses. Soil moisture was higher in +CO₂ mesocosms but +AM counteracted this effect. These results suggest that AM symbioses may be important mediators of plant community responses to anthropogenic CO₂ and N enrichment.     

The relationship between species richness and biomass changes from boreal to subtropical forests in China

Diversity‐manipulation experiments suggest a positive effect of biodiversity on ecosystem properties (EPs), but variable relationships between species richness and EPs have been reported in natural communities. An explanation for this discrepancy is that observed richness–EPs relationships in natural communities change with environment and species functional identities. But how the relationships change along broad‐scale climatic gradients has rarely been examined. In this paper, we sampled 848 plots of 20 × 30 m² from boreal to tropical forests across China. We examined plot biomass with respect to environmental factors, tree species richness and functional group identity (FGI, i.e. evergreen vs deciduous, and coniferous vs broadleaf). Variation partitioning was used to evaluate the relative effects of the three classes of factors. We found that, most of the ‘effects’ (percentage of variation explained) of richness and FGI on forest biomass were shared with environmental factors, but species richness and FGI still revealed significant effects in addition to environment for plots across China. Richness and FGI explained biomass mainly through their shared effects instead of independent effects, suggesting that the positive biodiversity effect is closely associated with a sampling effect. The relative effects of richness, FGI and environment varied latitudinally: the independent effects of environment and richness decreased from boreal to subtropical forests, whereas the total effect of FGI increased. We also found that the slope of richness–biomass relationship decreased monotonically from boreal to subtropical forests, possibly because of decreasing complementarity and increasing competition with increasing productivity. Our results suggest that while species richness does have significant effects on forest biomass it is less important than environmental gradients and other biotic factors in shaping large‐scale biomass patterns. We suggest that understanding how and why the diversity–EPs relationships change along climatic gradient would be helpful for a better understanding of real biodiversity effects in natural communities.     

Disentangling species and functional group richness effects on soil N cycling in a grassland ecosystem

Species richness (SR) and functional group richness (FGR) are often confounded in both observational and experimental field studies of biodiversity and ecosystem function. This precludes discernment of their separate influences on ecosystem processes, including nitrogen (N) cycling, and how those influences might be moderated by global change factors. In a 17‐year field study of grassland species, we used two full factorial experiments to independently vary SR (one or four species, with FGR = 1) and FGR (1–4 groups, with SR = 4) to assess SR and FGR effects on ecosystem N cycling and its response to elevated carbon dioxide (CO₂) and N addition. We hypothesized that increased plant diversity (either SR or FGR) and elevated CO₂ would enhance plant N pools because of greater plant N uptake, but decrease soil N cycling rates because of greater soil carbon inputs and microbial N immobilization. In partial support of these hypotheses, increasing SR or FGR (holding the other constant) enhanced total plant N pools and decreased soil nitrate pools, largely through higher root biomass, and increasing FGR strongly reduced mineralization rates, because of lower root N concentrations. In contrast, increasing SR (holding FGR constant and despite increasing total plant C and N pools) did not alter root N concentrations or net N mineralization rates. Elevated CO₂ had minimal effects on plant and soil N metrics and their responses to plant diversity, whereas enriched N increased plant and soil N pools, but not soil N fluxes. These results show that functional diversity had additional effects on both plant N pools and rates of soil N cycling that were independent of those of species richness.