Bottom-up effects of plant genotype on aphids, ants, and predators

Theory predicts that bottom-up ecological forces can affect community dynamics, but whether this extends to the effects of heritable plant variation on tritrophic communities is poorly understood. In a field experiment, I contrasted the effects of plant genotype (28 genotypes; 1064 plants), aphid density, and the presence/absence of mutualistic ants in affecting the per capita population growth of a specialist aphid herbivore, as well as the effects of plant genotype on the third trophic level. Plant genotype strongly affected aphid population growth rate, explaining 29% of the total variation in growth rate, whereas aphid density and ant-aphid interactions explained substantially less variation (< 2%) in aphid population growth rate. Plant genotype also had direct and indirect effects on the third trophic level, affecting the abundance of aphid-tending ants and the richness of predators. Multiple regression identified several heritable plant traits that explained 49% of the variation in aphid growth rate and 30% of the variation in ant abundance among plant genotypes. These bottom-up effects of plant genotype on tritrophic interactions were independent of the effects of either initial aphid density or the presence/absence of mutualistic ants. This study shows that plant genotype can be one of the most important ecological factors shaping tritrophic communities.     

Functional and phylogenetic diversity as predictors of biodiversity--ecosystem-function relationships

How closely does variability in ecologically important traits reflect evolutionary divergence? The use of phylogenetic diversity (PD) to predict biodiversity effects on ecosystem functioning, and more generally the use of phylogenetic information in community ecology, depends in part on the answer to this question. However, comparisons of the predictive power of phylogenetic diversity and functional diversity (FD) have not been conducted across a range of experiments. To address how phylogenetic diversity and functional trait variation control biodiversity effects on biomass production, we summarized the results of 29 grassland plant experiments where both the phylogeny of plant species used in the experiments is well described and where extensive trait data are available. Functional trait variation was only partially related to phylogenetic distances between species, and the resulting FD values therefore correlate only partially with PD. Despite these differences, FD and PD predicted biodiversity effects across all experiments with similar strength, including in subsets that excluded plots with legumes and that focused on fertilization experiments. Two- and three-trait combinations of the five traits used here (leaf nitrogen percentage, height, specific root length, leaf mass per unit area, and nitrogen fixation) resulted in the FD values with the greatest predictive power. Both PD and FD can be valuable predictors of the effect of biodiversity on ecosystem functioning, which suggests that a focus on both community trait diversity and evolutionary history can improve understanding of the consequences of biodiversity loss.     

A direct comparison of the consequences of plant genotypic and species diversity on communities and ecosystem function

Biodiversity loss is proceeding at an unprecedented rate, yet we lack a thorough understanding of the consequences of losing diversity at different scales. While species diversity is known to impact community and ecosystem processes, genotypic diversity is assumed to have relatively smaller effects. Nonetheless, a few recent studies suggest that genotypic diversity may have quantitatively similar ecological consequences compared to species diversity. Here we show that increasing either genotypic diversity of common evening primrose (Oenothera biennis) or species diversity of old-field plant species resulted in nearly equivalent increases (approximately 17%) in aboveground primary production. The predominant mechanism explaining this effect, niche complementarity, was similar for each type of diversity. Arthropod species richness also increased with both types of plant diversity, but the mechanisms leading to this effect differed: abundance-driven accumulation of arthropod species was important in plant genotypic polycultures, whereas resource specialization was important in plant species polycultures. Thus, similar increases in primary productivity differentially impacted higher trophic levels in response to each type of plant diversity. These results highlight important ecological similarities and differences between genotypic and species diversity and suggest that genotypic diversity may play a larger role in community and ecosystem processes than previously realized.     

Genotypic richness and phenotypic dissimilarity enhance population performance

Increases in biodiversity can result from an increase in species richness, as well as from a higher genetic diversity within species. Intraspecific genetic diversity, measured as the number of genotypes, can enhance plant primary productivity and have cascading effects at higher trophic levels, such as an increase in herbivore and predator richness. The positive effects of genotypic mixtures are not only determined by additive effects, but also by interactions among genotypes, such as facilitation or inhibition. However, so far there has been no effort to predict the extent of such effects. In this study, we address the question of whether the magnitude of the effect of genotype number on population performance can be explained by the extent of dissimilarity in key traits among genotypes in a mixture. We examine the relative contribution of genotype number and phenotypic dissimilarity among genotypes to population performance of the soil arthropod, Orchesella cincta. Nearly homogeneous genotypes were created from inbred isofemale lines. Phenotypic dissimilarity among genotypes was assessed in terms of three life-history traits that are associated with population growth rate, i.e., egg size, egg development time, and juvenile growth rate. A microcosm experiment with genotype mixtures consisting of one, two, four, and eight genotypes, showed that genotypic richness strongly increased population size and biomass production and was associated with greater net diversity effects. Most importantly, there was a positive log-linear relationship between phenotypic dissimilarity in a mixture and the net diversity effects for juvenile population size and total biomass. In other words, the degree of phenotypic dissimilarity among genotypes determined the magnitude of the genotypic richness effect, although this relationship leveled off at higher values of phenotypic dissimilarity. Although the exact mechanisms responsible for these effects are currently unknown, similar advantages of trait dissimilarity have been found among species. Hence, to better understand population performance, genotype number and phenotypic dissimilarity should be considered collectively.     

Plant-soil microorganism interactions: heritable relationship between plant genotype and associated soil microorganisms

Although soil microbial communities are known to play crucial roles in the cycling of nutrients in forest ecosystems and can vary by plant species, how microorganisms respond to the subtle gradients of plant genetic variation is just beginning to be appreciated. Using a model Populus system in a common garden with replicated clones of known genotypes, we evaluated microbial biomass and community composition as quantitative traits. Two main patterns emerged. (1) Plant genotype influenced microbial biomass nitrogen in soils under replicated genotypes of Populus angustifolia, F1, and backcross hybrids, but not P. fremontii. Genotype explained up to 78% of the variation in microbial biomass as indicated by broad-sense heritability estimates (i.e., clonal repeatability). A second estimate of microbial biomass (total phospholipid fatty acid) was more conservative and showed significant genotype effects in P. angustifolia and backcross hybrids. (2) Plant genotype significantly influenced microbial community composition, explaining up to 70% of the variation in community composition within P. angustifolia genotypes alone. These findings suggest that variation in above- and belowground traits of individual plant genotypes can alter soil microbial dynamics, and suggests that further investigations of the evolutionary implications of genetic feedbacks are warranted.     

Beyond the ecological: biological invasions alter natural selection on a native plant species

Biological invasions can have strong ecological effects on native communities by altering ecosystem functions, species interactions, and community composition. Even though these ecological effects frequently impact the population dynamics and fitness of native species, the evolutionary consequences of biological invasions have received relatively little attention. Here, I show that invasions impose novel selective pressures on a native plant species. By experimentally manipulating community composition, I found that the exotic plant Medicago polymorpha and the exotic herbivore Hypera brunneipennis alter the strength and, in some instances, the direction of natural selection on the competitive ability and anti-herbivore defenses of the native plant Lotus wrangelianus. Furthermore, the community composition of exotics influenced which traits were favored. For example, high densities of the exotic herbivore Hypera selected for increased resistance to herbivores in the native Lotus; however, when Medicago also was present, selection on this defense was eliminated. In contrast, selection on tolerance, another plant defense trait, was highest when both Hypera and Medicago were present at high densities. Thus, multiple exotic species may interact to influence the evolutionary trajectories of native plant populations, and patterns of selection may change as additional exotic species invade the community.     

Genetics, environment, and their interaction determine efficacy of chemical defense in trembling aspen

Optimal defense theories suggest that a trade-off between defense costs and benefits maintains genetic variation within plant populations. This study assessed the independent and interactive effects of genetic- and environment-based variation in aspen leaf chemistry on insect performance, preference, and defoliation. Gypsy moth larvae were released into screenhouses containing eight aspen genotypes growing with high and low levels of nutrient availability. Plant chemistry, defoliation, and larval growth rates varied in response to genotype, nutrient availability, and their interaction. Total phenolic glycoside concentrations were inversely correlated with patterns of larval preference and were the best predictor of larval performance and defoliation among genotypes. Low-nutrient trees were less heavily defoliated and afforded decreased larval growth rates compared with high-nutrient trees. Nutrient availability mediated the defense benefits of phenolic glycosides, as plant chemistry explained significantly less variation in defoliation in low- compared with high-nutrient trees (7% vs. 44% of variation explained). These results suggest that spatial and temporal variation in resource availability may influence the relative magnitude of defense benefits in plants. Environmental mediation of the defense costs and benefits likely leads to diversifying selection and may maintain genetic polymorphisms in chemical defense traits in plant populations.     

Incubation time, functional litter diversity, and habitat characteristics predict litter-mixing effects on decomposition

Plant diversity influences many fundamental ecosystem functions, including carbon and nutrient dynamics, during litter breakdown. Mixing different litter species causes litter mixtures to lose mass at different rates than expected from component species incubated in isolation. Such nonadditive litter-mixing effects on breakdown processes often occur idiosyncratically because their direction and magnitude change with incubation time, litter species composition, and ecosystem characteristics. Taking advantage of results from 18 litter mixture experiments in streams, we examined whether the direction and magnitude of nonadditive mixing effects are randomly determined. Across 171 tested litter mixtures and 510 incubation time-by-mixture combinations, nonadditive effects on breakdown were common and on average resulted in slightly faster decomposition than expected. In addition, we found that the magnitude of nonadditive effects and the relative balance of positive and negative responses in mixtures change predictably over time, and both were related to an index of functional litter diversity and selected environmental characteristics. Based on these, it should be expected that nonadditive effects are stronger for litter mixtures made of functionally dissimilar species especially in smaller streams. Our findings demonstrate that effects of litter diversity on plant mixture breakdown are more predictable than generally thought. We further argue that the consequences of current worldwide homogenization in the composition of plant traits on carbon and nutrient dynamics could be better inferred from long-duration experiments that manipulate both functional litter diversity and ecosystem characteristics in "hotspots of biodiversity effects," such as small streams.     

Disturbance and resource availability act differently on the same suite of plant traits: revisiting assembly hypotheses

Understanding the mechanisms of trait selection at the scale of plant communities is a crucial step toward predicting community assembly. Although it is commonly assumed that disturbance and resource availability constrain separate suites of traits, representing the regenerative and established phases, respectively, a quantification and test of this accepted hypothesis is still lacking due to limitations of traditional statistical techniques. In this paper we quantify, using structural equation modeling (SEM), the relative contributions of disturbance and resource availability to the selection of suites of traits at the community scale. Our model specifies and reflects previously obtained ecological insights, taking disturbance and nutrient availability as central drivers affecting leaf, allometric, seed, and phenology traits in 156 (semi-) natural plant communities throughout The Netherlands. The common hypothesis positing that disturbance and resource availability each affect a set of mutually independent traits was not consistent with the data. Instead, our final model shows that most traits are strongly affected by both drivers. In addition, trait-trait constraints are more important in community assembly than environmental drivers in half of the cases. Both aspects of trait selection are crucial for correctly predicting ecosystem processes and community assembly, and they provide new insights into hitherto underappreciated ecological interactions.     

Ecological consequences of carbon substrate identity and diversity in a laboratory study

Plants return a wide range of carbon (C) substrates to the soil system. The decomposition rate of these substrates is determined by their chemical nature, yet few studies have examined the relative ecological role of specific substrates (i.e., substrate identity) or mixtures of substrates. Carbon substrate identity and diversity may alter soil chemistry and soil community composition, resulting in changes in belowground ecosystem functions such as decomposition and nutrient transfer, creating feedbacks that may affect plant growth and the aboveground community. A laboratory experiment was set up in which eight C substrates of varying chemical complexity were added to a base soil singly, in pairs, fours, or with all eight together every four days over a 92-day period. After 92 days these soils were analyzed for changes in chemistry, microbial community structure, and components of ecosystem functioning. The identity of the added C substrates significantly affected soil chemistry, microbial basal and substrate-induced respiration, and soil microbial community structure measured by either the catabolic response profile (CRP) technique or phospholipid fatty acid composition. These belowground changes strongly affected the ability of the soil microflora to decompose cellulose paper, probably because of differential effects of the C substrates on soil energy supplies and enzyme activities. The addition of C substrates to soils also reduced plant growth compared to the unamended control soil, but less so in soils amended with a tannin than those amended with other substrates. Carbon substrate diversity effects saturated at low diversity levels, tended to have neutral or negative effects on ecosystem functions, and depended strongly on which C substrates were added. It increased CRP compound use but had little effect on other measures of the soil microbial community. Overall, results showed that the chemical nature of C substrates added to soil, and sometimes their diversity, can affect the soil microbial community and soil chemistry, which subsequently affect other ecosystem processes such as decomposition and plant growth. The identity and diversity of substrates that plants add to soil may therefore have important consequences for both above- and belowground ecosystem functions.     

Effects of plant species traits on ecosystem processes: experiments in the Patagonian steppe

Several experiments have shown that aboveground net primary productivity increases with plant species richness. The main mechanism proposed to explain this relationship is niche complementarity, which is determined by differences in plant traits that affect resource use. We combined field and laboratory experiments using the most abundant species of the Patagonian steppe to identify which are the traits that determine niche complementarity in this ecosystem. We estimated traits that affect carbon, water, microclimate, and nitrogen dynamics. The most important traits distinguishing among species, from the standpoint of their effects on ecosystem functioning, were potential soil nitrification, rooting depth, and soil thermal amplitude. Additionally, we explored the relationship between trait diversity and aboveground net primary production (ANPP) using a manipulative field experiment. ANPP and the fraction of ANPP accounted for by trait diversity increased with number of traits. The effect of trait diversity decreased as the number of traits increased. Here, the use of traits gave us a mechanistic understanding of niche complementarity in the Patagonian steppe.     

The relative importance of host-plant genetic diversity in structuring the associated herbivore community

Recent studies suggest that intraspecific genetic diversity in one species may leave a substantial imprint on the surrounding community and ecosystem. Here, we test the hypothesis that genetic diversity within host-plant patches translates into consistent and ecologically important changes in the associated herbivore community. More specifically, we use potted, grafted oak saplings to construct 41 patches of four saplings each, with one, two, or four tree genotypes represented among the host plants. These patches were divided among two common gardens. Focusing first at the level of individual trees, we assess how tree-specific genotypic identity, patch-level genetic diversity, garden-level environmental variation, and their interactions affect the structure of the herbivore community. At the level of host-plant patches, we analyze whether the joint responses of herbivore species to environmental variation and genetic diversity result in differences in species diversity among tree quartets. Strikingly, both species-specific abundances and species diversity varied substantially among host-tree genotypes, among common gardens, and among specific locations within individual gardens. In contrast, the genetic diversity of the patch left a detectable imprint on local abundances of only two herbivore taxa. In both cases, the effect of genetic diversity was inconsistent among gardens and among host-plant genotypes. While the insect community differed significantly among individual host-plant genotypes, there were no interactive effects of the number of different genotypes within the patch. Overall, additive effects of intraspecific genetic diversity of the host plant explained a similar or lower proportion (7-10%) of variation in herbivore species diversity than did variation among common gardens. Combined with the few previous studies published to date, our study suggests that the impact of host-plant genetic diversity on the herbivore community can range from none to nonadditive, is generally low, and reaches its most pronounced impact at small spatial scales. Overall, our findings strengthen the emerging view that the impacts of genetic diversity are system, scale, and context dependent. As the next step in community genetics, we should then start asking not only whether genetic diversity matters, but under what circumstances its imprint is accentuated.     

Plant-trait-based modeling assessment of ecosystem-service sensitivity to land-use change

Evidence is accumulating that the continued provision of essential ecosystem services is vulnerable to land-use change. Yet, we lack a strong scientific basis for this vulnerability as the processes that drive ecosystem-service delivery often remain unclear. In this paper, we use plant traits to assess ecosystem-service sensitivity to land-use change in subalpine grasslands. We use a trait-based plant classification (plant functional types, PFTs) in a landscape modeling platform to model community dynamics under contrasting but internally consistent land-use change scenarios. We then use predictive models of relevant ecosystem attributes, based on quantitative plant traits, to make projections of ecosystem-service delivery. We show that plant traits and PFTs are effective predictors of relevant ecosystem attributes for a range of ecosystem services including provisioning (fodder), cultural (land stewardship), regulating (landslide and avalanche risk), and supporting services (plant diversity). By analyzing the relative effects of the physical environment and land use on relevant ecosystem attributes, we also show that these ecosystem services are most sensitive to changes in grassland management, supporting current agri-environmental policies aimed at maintaining mowing of subalpine grasslands in Europe.     

Species phylogenetic relatedness, priority effects, and ecosystem functioning

Species immigration history can structure ecological communities through priority effects, which are often mediated by competition. As competition tends to be stronger between species with more similar niches, we hypothesize that species phylogenetic relatedness, under niche conservatism, may be a reasonable surrogate of niche similarity between species, and thus influence the strength of priority effects. We tested this hypothesis using a laboratory microcosm experiment in which we established bacterial species pools with different levels of phylogenetic relatedness and manipulated the immigration history of species from each pool into microcosms. Our results showed that strong priority effects, and hence multiple community states, only emerged for the species pool with the greatest phylogenetic relatedness. Community assembly also resulted in a significant positive relationship between bacterial phylogenetic diversity and ecosystem functions. Interestingly, these results emerged despite a lack of phylogenetic conservatism for most of the bacterial functional traits considered. Our results highlight the utility of phylogenetic information for understanding the structure and functioning of ecological communities, even when phylogenetically conserved functional traits are not identified or measured.     

Invasive ecosystem engineer selects for different phenotypes of an associated native species

Invasive habitat-forming ecosystem engineers modify the abiotic environment and thus represent a major perturbation to many ecosystems. Because native species often persist in these invaded habitats but have no shared history with the ecosystem engineer, the engineer may impose novel selective pressure on native species. In this study, we used a phenotypic selection framework to determine whether an invasive habitat-forming ecosystem engineer (the seaweed Caulerpa taxifolia) selects for different phenotypes of a common co-occurring native species (the bivalve Anadara trapezia). Compared to unvegetated habitat, Caulerpa habitat has lower water flow, lower dissolved oxygen, and sediments are more silty and anoxic. We determined the performance consequences of variation in key functional traits that may be affected by these abiotic changes (shell morphology, gill mass, and palp mass) for Anadara transplanted into Caulerpa and unvegetated habitat. Both linear and nonlinear performance gradients in Anadara differed between habitats, and these gradients were stronger in Caulerpa compared to unvegetated sediment. Moreover, in Caulerpa alternate phenotypes performed well, and these phenotypes were different from the dominant phenotype in unvegetated sediment. By demonstrating that phenotype-performance gradients differ between habitats, we have highlighted a role for Caulerpa as an agent of selection on native species.     

A genetic basis to community repeatability and stability

Recent studies have shown that genetically based traits of plants can structure associated arthropod and microbial communities, but whether the effects are consistent and repeatable across years is unknown. If communities are both heritable (i.e., related individuals tend to support similar communities) and repeatable (i.e., the same patterns observed over multiple years), then plant genetics may also affect community properties previously thought to be emergent, such as "stability." Using replicated clones of narrowleaf cottonwood (Populus angustifolia) and examining an arthropod community of 103 species, we found that (1) individual tree genotypes supported significantly different arthropod communities, which exhibited broad-sense heritability; (2) these findings were highly repeatable over three consecutive years (repeatability = 0.91) indicating that community responses to individual tree genotypes are consistent from year to year; (3) differences among tree genotypes in community stability (i.e., changes in community composition over multiple years) exhibited broad-sense heritability (H(C)2 = 0.32). In combination, these findings suggest that an emergent property such as stability can be genetically based and thus subject to natural selection.     

Predators have large effects on ecosystem properties by changing plant diversity, not plant biomass

Cascading effects of predators on total plant trophic-level biomass tend to be weaker in terrestrial than in aquatic systems. Accordingly, it is hypothesized that top predator effects on terrestrial plant diversity and on ecosystem function should likewise be weak or unimportant. This report presents a test of this hypothesis using data from a long-term field experiment. The five-year experiment manipulated the trophic structure of an old field ecosystem by excluding either predators or predators and herbivores relative to an unmanipulated, natural control. Long-term manipulations led to systematic treatment effects on community properties (plant trophic-level biomass, plant species biomass, plant species evenness) and on ecosystem properties (supply rate of solar radiation, N mineralization rate). The strengths of top predator effects on community properties were modest compared with nonterrestrial systems. But, predator-caused changes in plant community structure via alteration of plant dominance, and hence plant species evenness, strengthened effects on ecosystem properties. Counter to the hypothesis, weak trophic cascades do not necessarily lead to weak indirect effects of predators on ecosystem properties.     

Leaf herbivory and decomposability in a Malaysian tropical rain forest

There is accumulating evidence that similar suites of plant traits may affect leaf palatability and leaf litter decomposability. However, the possible association between leaf herbivory and litter decomposition rates across species in species-diverse natural ecosystems such as tropical rain forests remains unexplored, despite its importance in estimating the herbivory effects on carbon and nutrient cycling of ecosystems. We found no strong association between leaf herbivory and litter decomposition rates across 40 tree species in a Malaysian tropical rain forest, even though the leaf and litter traits were tightly correlated. This is because the leaf and litter traits related to herbivory and decomposition rates in the field were inconsistent. Leaf toughness accounted for only a small part of the variation in the herbivory rate, whereas a number of litter traits (the leaf mass per area, lignin to nitrogen ratio, and condensed tannin concentration) accurately predicted the decomposition rate across species. These results suggest that herbivory rate across species may not be strongly related to single leaf traits, probably because plant-herbivore interactions in tropical rain forests are highly diverse; on the other hand, plant-decomposer interactions are less specific and can be governed by litter chemicals. We also investigated two factors, phylogeny and tree functional types, that could affect the relationship between herbivory and decomposition across species. Phylogenetic relatedness among the species did not affect the relationship between herbivory and decomposition. In contrast, when the plants were segregated according to their leaf emergence pattern, we found a significant positive relationship between herbivory and decomposition rates for continuous-leafing species. In these species, the condensed tannin to N ratios in leaves and litter were related to herbivory and decomposition rates, respectively. However, we did not observe a similar trend for synchronous-leafing species. These results suggest that the relationship between herbivory and decomposition may be more greatly affected by functional types than by phylogenetic relatedness among species. In conclusion, our results suggest that well-defended leaves are not necessarily less decomposable litter in a tropical rain forest community, implying that herbivory may not generate positive feedback for carbon and nutrient cycling in this type of ecosystem.     

Vole response to unintentional changes in the chemistry of GM poplars

Summary.  There is an increased interest for the use of GM trees in forestry and several commercially promising lines are now available. However, the ecological implications of the use of GM trees, e.g. effects on non-target natural enemies, have rarely been explored. The aim of this study was to determine if modification of non-defensive traits in GM poplars unintentionally can influence plant chemistry in a way that has consequences for palatability to voles. In a greenhouse experiment, we used two lines, SPS33A and SPS26, of GM hybrid poplars (Populus tremula x tremuloides) with 1.5 and 4 times, respectively, over-expression sucrose-phosphate synthase (SPS). This enzyme plays a central role in sucrose synthesis, affecting cold acclimation, mesophyll sucrose content and biomass production. As a control we used the isogenic unmodified wild type. Stems of these poplars were presented to bank voles and field voles in cafeteria experiments. The concentration of condensed tannins was higher in leaves of lines SPS33A and SPS26 than in the isogenic wild-type and the concentration of nitrogen was higher in line SPS33A than in both the wild-type and line SPS26. Although the bank voles consumed slightly less bark from SPS33A, there were no significant differences in the preference of bank vole or field vole for the different poplar lines. This indicates that the changes in plant chemistry were insufficient to produce any strong herbivore response or that alteration in tannins and nitrogen counteracted each other. Still, changes in the interactions between mammalian herbivores and GM trees are important to consider in future cost-benefits analyses of GM trees.     

Consistent effects of nitrogen fertilization on soil bacterial communities in contrasting systems

Ecosystems worldwide are receiving increasing amounts of reactive nitrogen (N) through anthropogenic activities. Although the effects of increased N inputs on plant communities have been reasonably well studied, few comparable studies have examined impacts on whole soil bacterial communities, though they play critical roles in ecosystem functioning. We sampled soils from two long-term ecological research (LTER) experimental N gradients, both of which have been amended with NH4NO3; a grassland at Cedar Creek (27 years of N additions) and an agricultural field at Kellogg Biological Station (8 years of N additions). By examining shifts in bacterial communities across these contrasting ecosystem types, we could test competing hypotheses about the direct and indirect factors that might drive bacterial responses to elevated N inputs. Bacterial community structure was highly responsive to N additions. We observed predictable and consistent changes in the structure of the bacterial communities across both ecosystem types. Our results suggest that bacterial communities across these gradients are more structured by N and/or soil carbon availability than by shifts in the plant community or soil pH associated with the elevated nitrogen inputs. In contrast to the pronounced shifts in bacterial community composition and in direct contrast to the patterns often observed in plant communities, increases in N availability did not have consistent effects on the richness and diversity of soil bacterial communities.