Plant Functional Group Identity Research Articles

How arbuscular mycorrhizal fungi drives herbaceous plants' C: N: P stoichiometry? A meta-analysis

Plant element stoichiometry is fundamental for preserving growth-related terrestrial ecosystem structures and functions. However, effects of arbuscular mycorrhizal fungi (AMF) on herbaceous plant element stoichiometry (carbon (C), nitrogen (N), and phosphorus (P)) remain unclear. In this study, we aimed at evaluating the potential effects of AMF on herbaceous plant C, N and P concentration and their C:N:P stoichiometry worldwide through a quantitative meta-analysis. We observed that AMF reduced C:P and N:P ratios in the shoot of plants by 35.83 % and 54.23 %, respectively, and in plant root organs by 36.24 % and 46.35 %, respectively. Conversely, C:N ratios increased in roots by 6.61 %. The negative effect of AMF on N:P and C:P ratios in plant shoots and root organs is mainly attributed to the plant benefits in P and N concentrations. AMF impact on plant C:N:P stoichiometry depends on fungal and plant functional group identities and soil nutrient availability. Our results suggest that plant functional group identity affects plant nutrient concentration, which, in turn, controls herbaceous plant C:N:P stoichiometry. Overall, we emphasize the importance of abiotic and biotic environmental factors in changing AMF effects on plant element stoichiometry. Therefore, clarifying the relationship between AMF and herbaceous plant C:N:P stoichiometry will improve our understanding of herbaceous plant stoichiometric variations in terrestrial ecosystems.

Contrasting responses of nitrogen: Phosphorus stoichiometry in plants and soils under grazing: A global meta‐analysis

Abstract Grazing by ungulate herbivores can greatly alter nitrogen ([N]) and phosphorus ([P]) concentrations in plants and soils. It is not clear, however, how grazing might affect N:P co‐limitation in grasslands depending on soil N and P availability. Here we selected 173 peer‐reviewed studies, which measured 12 key variables associated with changes in [N], [P] and N:P ratios (i.e. N:P stoichiometry) in soils and plants in the presence or absence of herbivore grazing. Subsequently, we addressed the magnitude and direction of grazing effects on these variables using a meta‐analysis approach. Grazing increased leaf [N] and [P] but decreased total and available soil [N] and [P]. Grazing also increased leaf N:P ratios while decreasing root and total soil N:P ratios. The response ratio (RR) of leaf N:P was negatively correlated with RR of plant‐available soil [P] and positively correlated with RR of available soil N:P ratio (rather than with RR of total soil N:P). Intensive grazing (e.g. heavy grazing or long‐term grazing) had in general more positive effects on plant N:P stoichiometry and negative effects on soil N:P stoichiometry than light grazing. Responses of plant‐soil N:P stoichiometry to grazing greatly varied depending on plant functional group identity, plant organizational level (i.e. species and community) and grassland type. Synthesis and applications. Our study suggests that understanding changes in available soil N:P stoichiometry (rather than total soil N:P) in response to grazing is crucial to predict nutrient co‐limitation in grassland biomes. Our findings show that P is more important for plant growth than generally thought due to greater reduction of plant‐available soil [P] under grazing. A better mechanistic understanding of the relationships between plant and available soil N:P stoichiometry under grazing will greatly help improve the sustainability of natural and semi‐natural grassland ecosystems.

Ecosystem functioning in urban grasslands: The role of biodiversity, plant invasions and urbanization.

Urbanization is driving the transformation of natural and rural ecosystems worldwide by affecting both the abiotic environment and the biota. This raises the question whether urban ecosystems are able to provide services in a comparable way to their non-urban counterparts. In urban grasslands, the effects of urbanization-driven ecological novelty and the role of plant diversity in modulating ecosystem functioning have received little attention. In this study, we assessed the influence of biodiversity, abiotic and biotic novelty on ecosystem functioning based on in situ measurements in non-manipulated grasslands along an urbanization gradient in Berlin (Germany). We focused on plant aboveground biomass (AGB), intrinsic water-use efficiency (iWUE) and 15N enrichment factor (Δδ15N) as proxies for biomass production, water and N cycling, respectively, within grassland communities, and tested how they change with plant biogeographic status (native vs alien), functional group and species identity. Approximately one third of the forb species were alien to Berlin and they were responsible for 13.1% of community AGB. Community AGB was positively correlated with plant-species richness. In contrast, iWUE and Δδ15N were mostly determined by light availability (depicted by sky view factor) and urban parameters like the percentage of impervious surface or human population density. We found that abiotic novelty potentially favors aliens in Berlin, mainly by enhancing their dispersal and fitness under drought. Mainly urban parameters indicating abiotic novelty were significantly correlated to both alien and native Δδ15N, but to AGB and iWUE of alien plants only, pointing to a stronger impact of abiotic novelty on N cycling compared to C and water cycling. At the species level, sky view factor appeared to be the prevailing driver of photosynthetic performance and resource-use efficiency. Although we identified a significant impact of abiotic novelty on AGB, iWUE and Δδ15N at different levels, the relationship between species richness and community AGB found in the urban grasslands studied in Berlin was comparable to that described in non-urban experimental grasslands in Europe. Hence, our results indicate that conserving and enhancing biodiversity in urban ecosystems is essential to preserve ecosystem services related to AGB production. For ensuring the provision of ecosystem services associated to water and N use, however, changes in urban abiotic parameters seem necessary.

Precrop Functional Group Identity Affects Yield of Winter Barley but Less so High Carbon Amendments in a Mesocosm Experiment.

Nitrate leaching is a pressing environmental problem in intensive agriculture. Especially after the crop harvest, leaching risk is greatest due to decomposing plant residues, and low plant nutrient uptake and evapotranspiration. The specific crop also matters: grain legumes and canola commonly result in more leftover N than the following winter crop can take up before spring. Addition of a high carbon amendment (HCA) could potentially immobilize N after harvest. We set up a 2-year mesocosm experiment to test the effects of N fertilization (40 or 160 kg N/ha), HCA addition (no HCA, wheat straw, or sawdust), and precrop plant functional group identity on winter barley yield and soil C/N ratio. Four spring precrops were sown before winter barley (white lupine, faba bean, spring canola, spring barley), which were selected based on a functional group approach (colonization by arbuscular mycorrhizal fungi [AMF] and/or N2-fixing bacteria). We also measured a subset of faba bean and spring barley for leaching over winter after harvest. As expected, N fertilization had the largest effect on winter barley yield, but precrop functional identity also significantly affected the outcome. The non-AMF precrops white lupine and canola had on average a positive effect on yield compared to the AMF precrops spring barley and faba bean under high N (23% increase). Under low N, we found only a small precrop effect. Sawdust significantly reduced the yield compared to the control or wheat straw under either N level. HCAs reduced nitrate leaching over winter, but only when faba bean was sown as a precrop. In our setup, short-term immobilization of N by HCA addition after harvest seems difficult to achieve. However, other effects such as an increase in SOM or nutrient retention could play a positive role in the long term. Contrary to the commonly found positive effect of AMF colonization, winter barley showed a greater yield when it followed a non-AMF precrop under high fertilization. This could be due to shifts of the agricultural AMF community toward parasitism.

Differential responses of soil bacteria, fungi, archaea and protists to plant species richness and plant functional group identity.

Plants are known to influence belowground microbial community structure along their roots, but the impacts of plant species richness and plant functional group (FG) identity on microbial communities in the bulk soil are still not well understood. Here, we used 454-pyrosequencing to analyse the soil microbial community composition in a long-term biodiversity experiment at Jena, Germany. We examined responses of bacteria, fungi, archaea, and protists to plant species richness (communities varying from 1 to 60 sown species) and plant FG identity (grasses, legumes, small herbs, tall herbs) in bulk soil. We hypothesized that plant species richness and FG identity would alter microbial community composition and have a positive impact on microbial species richness. Plant species richness had a marginal positive effect on the richness of fungi, but we observed no such effect on bacteria, archaea and protists. Plant species richness also did not have a large impact on microbial community composition. Rather, abiotic soil properties partially explained the community composition of bacteria, fungi, arbuscular mycorrhizal fungi (AMF), archaea and protists. Plant FG richness did not impact microbial community composition; however, plant FG identity was more effective. Bacterial richness was highest in legume plots and lowest in small herb plots, and AMF and archaeal community composition in legume plant communities was distinct from that in communities composed of other plant FGs. We conclude that soil microbial community composition in bulk soil is influenced more by changes in plant FG composition and abiotic soil properties, than by changes in plant species richness per se.

Mycorrhizal Symbiotic Efficiency on C3 and C4 Plants under Salinity Stress - A Meta-Analysis.

A wide range of C3 and C4 plant species could acclimatize and grow under the impact of salinity stress. Symbiotic relationship between plant roots and arbuscular mycorrhizal fungi (AMF) are widespread and are well known to ameliorate the influence of salinity stress on agro-ecosystem. In the present study, we sought to understand the phenomenon of variability on AMF symbiotic relationship on saline stress amelioration in C3 and C4 plants. Thus, the objective was to compare varied mycorrhizal symbiotic relationship between C3 and C4 plants in saline conditions. To accomplish the above mentioned objective, we conducted a random effects models meta-analysis across 60 published studies. An effect size was calculated as the difference in mycorrhizal responses between the AMF inoculated plants and its corresponding control under saline conditions. Responses were compared between (i) identity of AMF species and AMF inoculation, (ii) identity of host plants (C3 vs. C4) and plant functional groups, (iii) soil texture and level of salinity and (iv) experimental condition (greenhouse vs. field). Results indicate that both C3 and C4 plants under saline condition responded positively to AMF inoculation, thereby overcoming the predicted effects of symbiotic efficiency. Although C3 and C4 plants showed positive effects under low (EC < 4 ds/m) and high (>8 ds/m) saline conditions, C3 plants showed significant effects for mycorrhizal inoculation over C4 plants. Among the plant types, C4 annual and perennial plants, C4 herbs and C4 dicot had a significant effect over other counterparts. Between single and mixed AMF inoculants, single inoculants Rhizophagus irregularis had a positive effect on C3 plants whereas Funneliformis mosseae had a positive effect on C4 plants than other species. In all of the observed studies, mycorrhizal inoculation showed positive effects on shoot, root and total biomass, and in nitrogen, phosphorous and potassium (K) uptake. However, it showed negative effects in sodium (Na) uptake in both C3 and C4 plants. This influence, owing to mycorrhizal inoculation, was significantly higher in K uptake in C4 plants. For our analysis, we concluded that AMF-inoculated C4 plants showed more competitive K+ ions uptake than C3 plants. Therefore, maintenance of high cytosolic K+/Na+ ratio is a key feature of plant salt tolerance. Studies on the detailed mechanism for the selective transport of K in C3 and C4 mycorrhizal plants under salt stress is lacking, and this needs to be explored.

Plant diversity and functional groups affect Si and Ca pools in aboveground biomass of grassland systems

Plant diversity is an important driver of nitrogen and phosphorus stocks in aboveground plant biomass of grassland ecosystems, but plant diversity effects on other elements also important for plant growth are less understood. We tested whether plant species richness, functional group richness or the presence/absence of particular plant functional groups influences the Si and Ca concentrations (mmolg(-1)) and stocks (mmolm(-2)) in aboveground plant biomass in a large grassland biodiversity experiment (Jena Experiment). In the experiment including 60 temperate grassland species, plant diversity was manipulated as sown species richness (1, 2, 4, 8, 16) and richness and identity of plant functional groups (1-4; grasses, small herbs, tall herbs, legumes). We found positive species richness effects on Si as well as Ca stocks that were attributable to increased biomass production. The presence of particular functional groups was the most important factor explaining variation in aboveground Si and Ca stocks (mmol m(-2)). Grass presence increased the Si stocks by 140% and legume presence increased the Ca stock by 230%. Both the presence of specific plant functional groups and species diversity altered Si and Ca stocks, whereas Si and Ca concentration were affected mostly by the presence of specific plant functional groups. However, we found a negative effect of species diversity on Si and Ca accumulation, by calculating the deviation between mixtures and mixture biomass proportions, but in monoculture concentrations. These changes may in turn affect ecosystem processes such as plant litter decomposition and nutrient cycling in grasslands.

Effects of plant functional group loss on soil biota and net ecosystem exchange: a plant removal experiment in the Mongolian grassland

Summary The rapid loss of global biodiversity can greatly affect the functioning of above‐ground components of ecosystems. However, how such biodiversity losses affect below‐ground communities and linkages to soil carbon (C) sequestration is unclear. Here, we describe how losses in plant functional groups (PFGs) affect soil microbial and nematode communities and net ecosystem exchange (NEE) in a 4‐year removal experiment conducted on the Mongolian plateau, the world's largest remaining natural grassland. Our results demonstrated that the biomasses or abundances of most components of the two below‐ground communities (microbes and nematodes) were negatively affected by PFG loss and were positively related to above‐ground plant biomass. The removal of dominant PFGs (perennial bunchgrasses and perennial rhizomatous grasses) reduced the biomass or abundance of below‐ground community components while removal of less dominant PFGs (perennial forbs and annuals/biennials) did not change or increased the biomass or abundance of below‐ground community components. The biomass‐based ratio of fungal to bacterial microbes and the number‐based ratio of fungal‐feeding to bacterial‐feeding nematodes decreased with increasing PFG losses. Variation partitioning analyses showed that the identity of PFGs together with above‐ground plant biomass explained most of the total variation in soil microbes and that the identity of PFGs and above‐ground plant biomass together with nematode food resources explained most of the total variation in soil nematodes. The increase in NEE with PFG loss was mainly explained by decreases in above‐ground plant biomass and the ratio of fungi to bacteria. Synthesis. The shift of below‐ground communities from a fungal‐based to a bacterial‐based energy channel as PFG richness decreases indicates that less diverse grassland ecosystems will have lower nutrient retention and hence be more sensitive to land‐use or climate change. The dominant effects of above‐ground plant biomass and below‐ground communities on NEE indicate that PFG loss resulting from land‐use or climate change has the potential to reduce C sequestration in semi‐arid grassland soils. These findings suggest that predictive models may need to consider the composition of above‐ground and below‐ground communities in order to accurately simulate the dynamics of CO2 fluxes in terrestrial ecosystems.

Functional trait responses to grazing are mediated by soil moisture and plant functional group identity.

Abundant evidence has shown that grazing alters plant functional traits, community structure and ecosystem functioning of grasslands. Few studies, however, have tested how plant responses to grazing are mediated by resource availability and plant functional group identity. We examined the effects of grazing on functional traits across a broad range of species along a soil moisture gradient in Inner Mongolia grassland. Our results showed that trait syndromes of plant size (individual biomass) and shoot growth (leaf N content and leaf density) distinguished plant species responses to grazing. The effects of grazing on functional traits were mediated by soil moisture and dependent on functional group identity. For most species, grazing decreased plant height but increased leaf N and specific leaf area (SLA) along the moisture gradient. Grazing enhanced the community-weighted attributes (leaf NCWM and SLACWM), which were triggered mainly by the positive trait responses of annuals and biennials and perennial grasses, and increased relative abundance of perennial forbs. Our results suggest that grazing-induced species turnover and increased intraspecific trait variability are two drivers for the observed changes in community weighted attributes. The dominant perennial bunchgrasses exhibited mixed tolerance–resistance strategies to grazing and mixed acquisitive–conservative strategies in resource utilization.

Soil fertility and the carbon:nutrient stoichiometry of herbaceous plant species

Biomass allocation to plant aboveground and belowground compartments may change in response to changes in soil nutrient fertility (e.g., nitrogen (N) and phosphorous (P) availability). It is not clear however whether and how changes in soil fertility could influence carbon(C):nutrient stoichiometry of wild plants distributed along environmental gradients. Here we use ecological stoichiometric theory to test two hypotheses: (1) C:N (or C:P) ratios of different plant compartments (i.e., roots, stems and leaves) increase when soil N (or P) availability decreases, (2) the relative availability of N compared to P in soils predictably influences C:nutrient ratios (e.g., high soil N:P ratios are associated with high plant C:P ratios). Data from 72 wild plant species were collected along a gradient of soil development determined by the temporal progression of four primary ecological successions across Europe. We found significant changes in soil N and P availability and content along the four vegetation successions, and our hypotheses were partially confirmed: (1) stem and root C:N ratios increased with decreases in soil N availability, and (2) stem C:P ratios increased with increases in soil N:P ratios. We also found that plant functional group identity strongly influenced plant C:nutrient stoichiometry whereby plant C:N ratios were significantly lower in legumes and C:P ratios were significantly higher in grasses. Our study suggests that changes in (1) soil N and P availability and content, and (2) soil N:P ratios can influence plant C:nutrient stoichiometry. However, we also bring evidence that plant functional group identity can be more important than soil fertility in influencing plant C:nutrient stoichiometry. Further studies could address how changes in soil fertility interact with multiple plant functional traits to better understand patterns of plant species distribution along environmental gradients.

Biotic and abiotic properties mediating plant diversity effects on soil microbial communities in an experimental grassland.

Plant diversity drives changes in the soil microbial community which may result in alterations in ecosystem functions. However, the governing factors between the composition of soil microbial communities and plant diversity are not well understood. We investigated the impact of plant diversity (plant species richness and functional group richness) and plant functional group identity on soil microbial biomass and soil microbial community structure in experimental grassland ecosystems. Total microbial biomass and community structure were determined by phospholipid fatty acid (PLFA) analysis. The diversity gradient covered 1, 2, 4, 8, 16 and 60 plant species and 1, 2, 3 and 4 plant functional groups (grasses, legumes, small herbs and tall herbs). In May 2007, soil samples were taken from experimental plots and from nearby fields and meadows. Beside soil texture, plant species richness was the main driver of soil microbial biomass. Structural equation modeling revealed that the positive plant diversity effect was mainly mediated by higher leaf area index resulting in higher soil moisture in the top soil layer. The fungal-to-bacterial biomass ratio was positively affected by plant functional group richness and negatively by the presence of legumes. Bacteria were more closely related to abiotic differences caused by plant diversity, while fungi were more affected by plant-derived organic matter inputs. We found diverse plant communities promoted faster transition of soil microbial communities typical for arable land towards grassland communities. Although some mechanisms underlying the plant diversity effect on soil microorganisms could be identified, future studies have to determine plant traits shaping soil microbial community structure. We suspect differences in root traits among different plant communities, such as root turnover rates and chemical composition of root exudates, to structure soil microbial communities.

Plant functional group identity and diversity determine biotic resistance to invasion by an exotic grass

Summary Biotic resistance, the ability of species in a community to limit invasion, is central to our understanding of how communities at risk of invasion assemble after disturbances, but it has yet to translate into guiding principles for the restoration of invasion‐resistant plant communities. We combined experimental, functional, and modelling approaches to investigate processes of community assembly contributing to biotic resistance to an introduced lineage of Phragmites australis, a model invasive species in North America. We hypothesized that (i) functional group identity would be a good predictor of biotic resistance to P. australis, while species identity effect would be redundant within functional group (ii) mixtures of species would be more invasion resistant than monocultures. We classified 36 resident wetland plants into four functional groups based on eight functional traits. We conducted two competition experiments based on the additive competition design with P. australis and monocultures or mixtures of wetland plants. As an indicator of biotic resistance, we calculated a relative competition index (RCIavg) based on the average performance of P. australis in competition treatment compared with control. To explain diversity effect further, we partitioned it into selection effect and complementarity effect and tested several diversity–interaction models. In monoculture treatments, RCIavg of wetland plants was significantly different among functional groups, but not within each functional group. We found the highest RCIavg for fast‐growing annuals, suggesting priority effect. RCIavg of wetland plants was significantly greater in mixture than in monoculture mainly due to complementarity–diversity effect among functional groups. In diversity–interaction models, species interaction patterns in mixtures were described best by interactions between functional groups when fitted to RCIavg or biomass, implying niche partitioning. Synthesis. Functional group identity and diversity of resident plant communities are good indicators of biotic resistance to invasion by introduced Phragmites australis, suggesting niche pre‐emption (priority effect) and niche partitioning (diversity effect) as underlying mechanisms. Guiding principles to understand and/or manage biological invasion could emerge from advances in community theory and the use of a functional framework. Targeting widely distributed invasive plants in different contexts and scaling up to field situations will facilitate generalization.

Collembola species composition and diversity effects on ecosystem functioning vary with plant functional group identity

The relationship between decomposer diversity and ecosystem functioning is little understood although soils accommodate a significant proportion of worldwide biodiversity. Collembola are among the most abundant and diverse decomposers and are known to modify plant growth. We examined the effects of Collembola species diversity (one, two and three species belonging to different life history groups) and composition on litter decomposition and the performance of plant communities (above- and belowground productivity) of different functional groups (grasses, forbs and legumes). Collembola densities did not increase with diversity indicating niche overlap. Generally, Collembola species composition was a better predictor for ecosystem functioning than Collembola species number with the impacts of Collembola diversity and composition on ecosystem functioning strongly depending on plant functional group identity. Non-linear effects of Collembola diversity on litter decomposition and plant productivity suggest pronounced and context dependent species interactions and feeding habits. Net surface litter decomposition was decreased by Collembola, whereas root litter decomposition was at maximum in the highest Collembola diversity treatment. Forbs benefitted most from the presence of three Collembola species. Similarly, Collembola diversity influenced root depth distribution in a plant functional group specific way: while grass root biomass decreased with increasing Collembola diversity in the upper and lower soil layer, legume root biomass increased particularly in the lower soil layer. Idiosyncratic and context dependent effects of Collembola diversity and composition even in rather simple assemblages of one to three species suggest that changes in Collembola diversity may have unpredictable consequences for ecosystem functioning. The finding that changes in Collembola performance did not directly translate to alterations in ecosystem functioning indicates that response traits do not necessarily conform to effect traits. Distinct plant functional group specific impacts of Collembola diversity on root depth distribution are likely to modify plant competition in complex plant communities and add a novel mechanism how decomposers may affect plant community assembly.

Plant species richness drives the density and diversity of Collembola in temperate grassland

Declining biodiversity is one of the most important aspects of anthropogenic global change phenomena, but the implications of plant species loss for soil decomposers are little understood. We used the experimental grassland community of the Jena Experiment to assess the response of density and diversity of Collembola to varying plant species richness, plant functional group richness and plant functional group identity. We sampled the experimental plots in spring and autumn four years after establishment of the experimental plant communities. Collembola density and diversity significantly increased with plant species and plant functional group richness highlighting the importance of the singular hypothesis for soil invertebrates. Generally, grasses and legumes beneficially affected Collembola density and diversity, whereas effects of small herbs usually were detrimental. These impacts were largely consistent in spring and autumn. By contrast, in the presence of small herbs the density of hemiedaphic Collembola and the diversity of Isotomidae increased in spring whereas they decreased in autumn. Beneficial impacts of plant diversity as well as those of grasses and legumes were likely due to increased root and microbial biomass, and elevated quantity and quality of plant residues serving as food resources for Collembola. By contrast, beneficial impacts of small herbs in spring probably reflect differences in microclimatic conditions, and detrimental effects in autumn likely were due to low quantity and quality of resources. The results point to an intimate relationship between plants and the diversity of belowground biota, even at small spatial scales, contrasting the findings of previous studies. The pronounced response of soil animals in the present study was presumably due to the fact that plant communities had established over several years. As decomposer invertebrates significantly impact plant performance, changes in soil biota density and diversity are likely to have major feedbacks on plant community productivity and composition.

Soil nutrient heterogeneity modulates ecosystem responses to changes in the identity and richness of plant functional groups.

Recent research has shown that biodiversity may has its greatest impact on ecosystem functioning in heterogeneous environments. However, the role of soil heterogeneity as a modulator of ecosystem responses to changes in biodiversity remains poorly understood, as few biodiversity studies have explicitly considered this important ecosystem feature.We conducted a microcosm experiment over two growing seasons to evaluate the joint effects of changes in plant functional groups (grasses, legumes, non-legume forbs and a combination of them), spatial distribution of soil nutrients (homogeneous and heterogeneous) and nutrient availability (50 and 100 mg of nitrogen [N] added as organic material) on plant productivity and surrogates of carbon, phosphorous and N cycling (β-glucosidase and acid phosphatase enzymes and in situ N availability, respectively).Soil nutrient heterogeneity interacted with nutrient availability and plant functional diversity to determine productivity and nutrient cycling responses. All the functional groups exhibited precise root foraging patterns. Above- and belowground productivity increased under heterogeneous nutrient supply. Surrogates of nutrient cycling were not directly affected by soil nutrient heterogeneity. Regardless of their above- and belowground biomass, legumes increased the availability of soil inorganic N and the activity of the acid phosphatase and β-glucosidase enzymes.Our study emphasizes the role of soil nutrient heterogeneity as a modulator of ecosystem responses to changes in functional diversity beyond the species level. Functional group identity, rather than richness, can play a key role in determining the effects of biodiversity on ecosystem functioning.Synthesis. Our results highlight the importance of explicitly considering soil heterogeneity in diversity-ecosystem functioning experiments, where the identity of the plant functional group is of major importance. Such consideration will improve our ability to fully understand the role of plant diversity on ecosystem functioning in ubiquitous heterogeneous environments.

Soil microbial communities resistant to changes in plant functional group composition

Abstract The soil community is an often ignored part of research which links plant biodiversity and ecosystem functioning despite their influence on numerous functions such as decomposition and nutrient cycling. Few consistent patterns have been detected that link plant and soil community composition. We used a removal experiment in a northern Canadian grassland to examine the effects of plant functional group identity on soil microbial community structure and function. Plant functional groups (graminoids, legumes and forbs) were removed independently from plots for five growing seasons (2003–2007) and in the fifth year effects on the soil microbial community were examined using substrate-induced respiration (SIR – a measure of metabolic diversity) and phospholipid fatty acid analysis (PLFA – a measure of microbial community composition). Removal treatments were also crossed with both a fertilizer treatment and a fungicide treatment to determine if effects of functional group identity on the soil community were context dependent. Plant functional group identity had almost no effect on the soil microbial community as measured by either SIR or PLFA. Likewise, soil properties including total carbon, pH, moisture and nutrients showed a limited response to plant removals in the fifth year after removals. We found a direct effect of fertilizer on the soil community, with fertilized plots having decreased metabolic diversity, with a decreased ability to metabolize amino acids and a phenolic acid, but there was no direct soil microbial response to fungicide. We show that in this northern Canadian grassland the soil microbial community is relatively insensitive to changes in plant functional group composition, and suggest that in northern ecosystems, where plant material is only slowly incorporated into the soil, five growing seasons may be insufficient to detect the impact of a changing plant community on the soil microbes.

Nematicide impacts on nematodes and feedbacks on plant productivity in a plant diversity gradient

A major issue in current ecological research is the effect of biodiversity on ecosystem functioning. Although several studies reported a positive diversity – productivity relationship, the role of soil animals has been largely neglected. Nematodes are among the most widespread and important herbivores causing substantial yield losses in agriculture; however, impacts of nematodes on the diversity – productivity relationship in semi-natural plant communities have not been investigated until today. In the framework of the Jena Experiment (Thuringia, Germany) we established control and nematicide treated subplots to manipulate nematode densities on plots varying in plant species (1–16) and functional group richness (1–4). We explored the interacting effects of nematicide application and plant diversity on the main trophic groups of nematodes and on aboveground plant productivity. Nematicide application reduced the number of nematodes significantly, particularly that of plant feeders and predators. The negative impact of nematicide application on plant and bacterial feeders depended however on the diversity of the plant community. Total plant shoot biomass tended to decrease in the presence of ambient nematode densities. In detail, nematode effects varied however with plant functional group identity by reducing only the shoot biomass of herbs significantly but not that of legumes. Furthermore, the shoot biomass of grasses tended to decrease in the presence of ambient nematode densities. In contrast to total shoot biomass, nematodes decreased grass shoot biomass only in high diverse but not in low diverse plant communities. Thus, the present study for the first time highlights that nematodes likely modify the community structure und functions of semi-natural plant communities by altering the competition between plant functional groups and by attenuating the diversity – productivity relationship.

Functional identity versus species richness: herbivory resistance in plant communities.

The resistance of a plant community against herbivore attack may depend on plant species richness, with monocultures often much more severely affected than mixtures of plant species. Here, we used a plant–herbivore system to study the effects of selective herbivory on consumption resistance and recovery after herbivory in 81 experimental grassland plots. Communities were established from seed in 2002 and contained 1, 2, 4, 8, 16 or 60 plant species of 1, 2, 3 or 4 functional groups. In 2004, pairs of enclosure cages (1 m tall, 0.5 m diameter) were set up on all 81 plots. One randomly selected cage of each pair was stocked with 10 male and 10 female nymphs of the meadow grasshopper, Chorthippus parallelus. The grasshoppers fed for 2 months, and the vegetation was monitored over 1 year. Consumption resistance and recovery of vegetation were calculated as proportional changes in vegetation biomass. Overall, grasshopper herbivory averaged 6.8%. Herbivory resistance and recovery were influenced by plant functional group identity, but independent of plant species richness and number of functional groups. However, herbivory induced shifts in vegetation composition that depended on plant species richness. Grasshopper herbivory led to increases in herb cover at the expense of grasses. Herb cover increased more strongly in species-rich mixtures. We conclude that selective herbivory changes the functional composition of plant communities and that compositional changes due to selective herbivory depend on plant species richness.

Ecosystem properties determined by plant functional group identity

Summary 1. Ecosystem properties may be determined by the number of different species or groups of species in a community, the identity of those groups, and their relative abundance. The mass ratio theory predicts that the effect of species or groups of species on ecosystem properties will be dependent on their proportional abundance in a community. 2. Single plant functional groups (graminoids, legumes, non‐leguminous forbs) were removed from a natural grassland in northern Canada to examine the role of group identity in determining both ecosystem properties and biomass compensation by remaining species. Removals were conducted across two different environmental treatments (fertilization and fungicide) to examine the context dependency of functional group identity effects. 3. The degree of biomass compensation in the first 4 years after removal was influenced by the identity of the functional group removed and also of those remaining. When graminoids were removed, none of the remaining functional groups compensated for the loss of biomass. Graminoids partially compensated for the removal of forbs or legumes, with the degree of compensation depending on environmental treatments. 4. Light interception, soil moisture and soil nutrients were all influenced by functional group identity, with graminoids having a greater impact than expected based on their biomass contribution to the community. Legumes, in contrast, had very little effect on any of the ecosystem properties measured. 5. For most ecosystem properties measured, the role of plant functional groups was not context dependent; functional groups had the same effect on ecosystem properties regardless of fertilization or fungicide treatments. 6. Synthesis. We have shown that the effects of losing a functional group do not solely depend on the group’s dominance. In this northern grassland, there are greater effects of losing graminoids than one would predict based on their biomass contributions to the community, and functional group identity plays a critical role in determining the effects of diversity loss.

Plant functional group identity differentially affects leaf and root decomposition

AbstractLosses of species and changes in the composition of plant communities are likely to influence numerous ecosystem functions. Changes in the plant‐soil interactions that control decomposition, in particular, could alter carbon and nutrient cycling in soils and further alter other ecosystem functions. The effects of plant communities on decomposition may depend both on the type of tissue being decomposed and also on the different stages of the decomposition process. We used an experimental design where single plant functional groups were removed from a northern grassland to examine the role of plant identity in determining both short‐term and long‐term above‐ and belowground decomposition rates. Plant removals were conducted across fertilization and fungicide treatments to examine environmental context‐dependency of functional group identity effects on decomposition. There were significant effects of plant functional group identity on aboveground decomposition, with the loss of grasses and forbs slowing decomposition, whereas the effects on belowground decomposition were rare and transient. Effects of plant identity on decomposition were consistent in both short‐ and long‐term decomposition studies indicating that the influences of identity on the decomposition environment remained consistent throughout the different stages of the decomposition process. Both fertilizer and fungicide treatments affected overall decomposition rate, but there were few interactions between these treatments and plant removals. Although current species loss is likely to be happening in concert with environmental changes, the role a species plays in determining ecosystem functions such as decomposition may not be context‐dependent in these northern environments, and this may provide greater predictive power in determining the effects of species loss with changing environments. Further, as plant identity shows significant effects on litter decomposition rates, the effects of current and predicted future biodiversity losses may depend specifically on which species are lost.