Soil community changes during secondary succession to naturalized grasslands

Succession to a naturalized grassland from former agricultural land and pastures is accompanied by changes in plant biodiversity and in the soil community. These changes are the result of a reduction or elimination of management, fertilizer applications and of grazing by large herbivores. We review soil biology studies on agricultural land that are in various successional stages towards naturalized grasslands, where interactions between plant species composition changes and the soil ecology affect each other. In many chronosequence studies, the soil microbial community tends to shift towards a less bacterial, and more fungal dominated food web energy channel following a reduction in fertilizer inputs and grazing intensity. Whereas changes in microarthropod communities are obscured, nematode trophic functional group (ecological guild) changes respond to both plant and soil community changes. There are opportunities to further study the feedback interactions between roots and soil organisms in grasslands. A better understanding of the molecular feedback mechanisms would be beneficial in long-term grassland management.     

Soil microbial community composition as affected by restoration practices in California grassland

Agricultural practices have strong impacts on soil microbes including both the indices related to biomass and activity as well as those related to community composition. In a grassland restoration project in California, where native perennial bunchgrasses were introduced into non-native annual grassland after a period of intensive tillage, weeding, and herbicide use to reduce the annual seed bank, microbial community composition was investigated. Three treatments were compared: annual grassland, bare soil fallow, and restored perennial grassland. Soil profiles down to 80 cm in depth were investigated in four separate layers (0-15, 15-30, 30-60, and 60-80 cm) using both phospholipid ester-linked fatty acid (PLFAs) and ergosterol as biomarkers in addition to microbial biomass C by fumigation extraction. PLFA fingerprinting showed much stronger differences between the tilled bare fallow treatment vs. grasslands, compared to fewer differences between restored perennial grassland and annual grassland. The presence or absence of plants over several years clearly distinguished microbial communities. Microbial communities in lower soil layers were little affected by management practices. Regardless of treatment, soil depth caused a strong gradient of changing habitat conditions, which was reflected in Canonical Correspondence Analysis of PLFAs. Fungal organisms were associated with the presence of plants and/or litter since the total amount and the relative proportion of fungal markers were reduced in the tilled bare fallow and in lower layers of the grassland treatments. Total PLFA and soil microbial biomass were highly correlated, and fungal PLFA biomarkers showed strong correlations to ergosterol content. In conclusion, microbial communities are resilient to the grassland restoration process, but do not reflect the change in plant species composition that occurred after planting native bunchgrasses.     

The influence of soil properties on the structure of bacterial and fungal communities across land-use types

Land-use change can have significant impacts on soil conditions and microbial communities are likely to respond to these changes. However, such responses are poorly characterized as few studies have examined how specific changes in edaphic characteristics do, or do not, influence the composition of soil bacterial and fungal communities across land-use types. Soil samples were collected from four replicated (n = 3) land-use types (hardwood and pine forests, cultivated and livestock pasture lands) in the southeastern US to assess the effects of land-use change on microbial community structure and distribution. We used quantitative PCR to estimate bacterial–fungal ratios and clone libraries targeting small-subunit rRNA genes to independently characterize the bacterial and fungal communities. Although some soil properties (soil texture and nutrient status) did significantly differ across land-use types, other edaphic factors (e.g., pH) did not vary consistently with land-use. Bacterial–fungal ratios were not significantly different across the land-uses and distinct land-use types did not necessarily harbor distinct soil fungal or bacterial communities. Rather, the composition of bacterial and fungal communities was most strongly correlated with specific soil properties. Soil pH was the best predictor of bacterial community composition across this landscape while fungal community composition was most closely associated with changes in soil nutrient status. Together these results suggest that specific changes in edaphic properties, not necessarily land-use type itself, may best predict shifts in microbial community composition across a given landscape. In addition, our results demonstrate the utility of using sequence-based approaches to concurrently analyze bacterial and fungal communities as such analyses provide detailed phylogenetic information on individual communities and permit the robust assessment of the biogeographical patterns exhibited by soil microbial communities.     

Influence of grass species and soil type on rhizosphere microbial community structure in grassland soils

A number of studies have reported species specific selection of microbial communities in the rhizosphere by plants. It is hypothesised that plants influence microbial community structure in the rhizosphere through rhizodeposition. We examined to what extent the structure of bacterial and fungal communities in the rhizosphere of grasses is determined by the plant species and different soil types. Three grass species were planted in soil from one site, to identify plant-specific influences on rhizosphere microbial communities. To quantify the soil-specific effects on rhizosphere microbial community structure, we planted one grass species (Lolium perenne L.) into soils from three contrasting sites. Rhizosphere, non-rhizosphere (bulk) and control (non-planted) soil samples were collected at regular intervals, to examine the temporal changes in soil microbial communities. Rhizosphere soil samples were collected from both root bases and root tips, to investigate root associated spatial influences. Both fungal and bacterial communities were analysed by terminal restriction fragment length polymorphism (TRFLP). Both bacterial and fungal communities were influenced by the plant growth but there was no evidence for plant species selection of the soil microbial communities in the rhizosphere of the different grass species. For both fungal and bacterial communities, the major determinant of community structure in rhizospheres was soil type. This observation was confirmed by cloning and sequencing analysis of bacterial communities. In control soils, bacterial composition was dominated by Firmicutes and Actinobacteria but in the rhizosphere samples, the majority of bacteria belonged to Proteobacteria and Acidobacteria. Bacterial community compositions of rhizosphere soils from different plants were similar, indicating only a weak influence of plant species on rhizosphere microbial community structure.     

Development of soil microbial communities during tallgrass prairie restoration

Soil microbial communities were examined in a chronosequence of four different land-use treatments at the Konza Prairie Biological Station, Kansas. The time series comprised a conventionally tilled cropland (CTC) developed on former prairie soils, two restored grasslands that were initiated on former agricultural soils in 1998 (RG₉₈) and 1978 (RG₇₈), and an annually burned native tallgrass prairie (BNP), all on similar soil types. In addition, an unburned native tallgrass prairie (UNP) and another grassland restored in 2000 (RG₀₀) on a different soil type were studied to examine the effect of long-term fire exclusion vs. annual burning in native prairie and the influence of soil type on soil microbial communities in restored grasslands. Both 16S rRNA gene clone libraries and phospholipid fatty acid analyses indicated that the structure and composition of bacterial communities in the CTC soil were significantly different from those in prairie soils. Within the time series, soil physicochemical characteristics changed monotonically. However, changes in the microbial communities were not monotonic, and a transitional bacterial community formed during restoration that differed from communities in either the highly disturbed cropland or the undisturbed original prairie. The microbial communities of RG₉₈ and RG₀₀ grasslands were also significantly different even though they were restored at approximately the same time and were managed similarly; a result attributable to the differences in soil type and associated soil chemistry such as pH and Ca. Burning and seasonal effects on soil microbial communities were small. Similarly, changing plot size from 300 m² to 150 m² in area caused small differences in the estimates of microbial community structure. In conclusion, microbial community structure and biochemical properties of soil from the tallgrass prairie were strongly impacted by cultivation, and the microbial community was not fully restored even after 30 years.     

川西北不同沙化程度草地土壤细菌群落特征

研究川西北不同沙化程度草地（未沙化草地、轻度沙化草地、中度沙化草地、重度沙化草地）土壤细菌多样性和群落结构特征,利用Illumina二代高通量测序技术MiSeq对土壤细菌的16 SrRNA V₃—V₄可变区进行测序,研究土壤细菌多样性、物种组成和丰富度,并结合土壤理养分探讨影响细菌群落结构的环境因素,对发挥土壤潜在肥力,了解土壤健康状况,实现该区植被的管理与可持续利用有着重要的意义。结果表明:（1）不同沙化草地土壤养分具有明显差异,依次表现为:随着沙化程度的增加,土壤pH值逐渐增加,而土壤有机碳、全氮、全钾、碱解氮和速效磷逐渐降低;（2）不同沙化程度草地土壤样品中共检测到细菌的32个门,65个纲,169个目,优势菌门为变形菌门（Proteobacteria）、放线菌门（Actinobacteria）、酸杆菌门（Acidobacteria）、绿弯菌（Chloroflexi）、浮霉菌门（Planctomycetes）,主要的优势菌纲为放线杆菌纲（Actinobacteria）、α-变形菌纲（α-Proteobacteria）、酸杆菌纲（Acidobacteria）、β-变形菌纲（β-Proteobacteria）、浮霉菌纲（Planctomycetacia）,与沙化草地相比,未沙化草地优势菌主要是变形菌门（Proteobacteria）和放线菌门（Actinobacteria）;（3）随着沙化程度的增加,OUT数目、Chao指数、Ace指数、Shannon指数逐渐减小,其中不同沙化草地土壤细菌覆盖率和Simpson指数差异不显著（p > 0.05）;（4）冗余分析和Pearson相关性分析表明,土壤pH值、土壤有机碳（SOC）和全氮（TN）是土壤细菌群落结构和多样性的主要影响因子。     

The measurement of soil fungal:bacterial biomass ratios as an indicator of ecosystem self-regulation in temperate meadow grasslands

 There is much interest in the development of agricultural land management strategies aimed at enhancing reliance on ecosystem self-regulation rather than on artificial inputs such as fertilisers and pesticides. This study tested the usefulness of measures of soil microbial biomass and fungal:bacterial biomass ratios as indicators of effective conversion from an intensive grassland system, reliant mainly on fertilisers for crop nutrition, to a low-input system reliant mainly on self-regulation through soil biological pathways of nutrient turnover. Analysis of soils from a wide range of meadow grassland sites in northern England, along a gradient of long-term management intensity, showed that fungal:bacterial biomass ratios (measured by phospholipid fatty acid analysis; PLFA) were consistently and significantly higher in the unfertilised than the fertilised grasslands. There was also some evidence that microbial biomass, measured by chloroform fumigation and total PLFA, was higher in the unfertilised than in the fertilised grasslands. It was also found that levels of inorganic nitrogen (N), in particular nitrate-N, were significantly higher in the fertilised than in the unfertilised grasslands. However, microbial activity, measured as basal respiration, did not differ between the sites. A field manipulation trial was conducted to determine whether the reinstatement of traditional management on an improved mesotrophic grassland, for 6 years, resulted in similar changes in the soil microbial community. It was found that neither the cessation of fertiliser applications nor changes in cutting and grazing management significantly affected soil microbial biomass or the fungal:bacterial biomass ratio. It is suggested that the lack of effects on the soil microbial community may be related to high residual fertility caused by retention of fertiliser N in the soil. On the basis of these results it is recommended that following the reinstatement of low-input management, the measurement of a significant increase in the soil fungal:bacterial biomass ratio, and perhaps total microbial biomass, may be an indicator of successful conversion to a grassland system reliant of self-regulation. Received: 4 May 1998     

Evaluating patterns of biodiversity in managed grasslands using spatial turnover metrics

Market and policy incentives that encourage agricultural intensification, such as incentives for bioenergy, may contribute to biodiversity decline when they encourage a large-scale conversion of native and semi-natural ecosystems to production fields. In order to appreciate the impact of these incentives on biodiversity, it is imperative to better understand how native and semi-natural ecosystems contribute to plant diversity and composition. We studied the five most common types of managed grasslands in Northeastern Kansas, a region undergoing agricultural intensification. We analyzed plant community data recorded at three spatial scales in 98 managed grassland sites, and examined patterns of diversity and composition of plant species and functional groups; and spatial turnover of species and functional groups measured at different spatial scales. We found reduced soil quality and plant diversity as well as a lack of scale-dependent community patterns on sites that were historically cultivated. Forage management practices (haying or grazing) altered plant diversity and composition in native grassland remnants but not in non-native grasslands. We identified several opportunities where changes to existing management practices could benefit both conservation and bioenergy production objectives. Recommendations to conserve biodiversity include increasing the use of hay management or other biomass collection on native grassland remnants and improving the Conservation Reserve Program by increasing enrollment, adding more native species to seed mixes, and incorporating a periodic biomass collection. We also found that using measures of spatial turnover in community composition added important insights in understanding the effects of management decisions on biodiversity.     

Soil carbon content drives the biogeographical distribution of fungal communities in the black soil zone of northeast China

Black soils (Mollisols) are one of the most important soil resources for maintaining food security in China, and they are mainly distributed in northeast China. A previous comprehensive study revealed the biogeographical distribution patterns of bacterial communities in the black soil zone. In this study, we used the same soil samples and analyzed the 454 pyrosequencing data for the nuclear ribosomal internal transcribed spacer (ITS) region to examine the fungal communities in these black soils. A total of 220,812 fungal ITS sequences were obtained from 26 soil samples that were collected across the black soil zone. These sequences were classified into at least 5 phyla, 20 classes, greater than 70 orders and over 350 genera, suggesting a high fungal diversity across the black soils. The diversity of fungal communities and distribution of several abundant fungal taxa were significantly related to the soil carbon content. Non-metric multidimensional scaling and canonical correspondence analysis plots indicated that the fungal community composition was most strongly affected by the soil carbon content followed by soil pH. This finding differs from the bacterial community results, which indicated that soil pH was the most important edaphic factor in determining the bacterial community composition of these black soils. A variance partitioning analysis indicated that the geographic distance contributed 20% of the fungal community variation and soil environmental factors that were characterized explained approximately 35%. A pairwise analysis revealed that the diversity of the fungal community was relatively higher at lower latitudes, which is similar to the findings for the bacterial communities in the same region and suggests that a latitudinal gradient of microbial community diversity might occur in the black soil zone. By incorporating our previous findings on the bacterial communities, we can conclude that contemporary factors of soil characteristics are more important than historical factor of geographic distance in shaping the microbial community in the black soil zone of northeast China.     

煤矿复垦区不同恢复年限沙棘人工林土壤真菌群落特征

  目的  探讨矿区不同恢复年限复垦地沙棘人工林土壤真菌群落组成、多样性及其影响因子。  方法  以鄂尔多斯聚鑫龙煤矿复垦区不同恢复年限沙棘人工林为研究对象,撂荒草地为对照,对研究区土样进行采集,通过高通量测序对土样真菌群落测序分析。  结果  研究区共获得真菌8门、30纲、69目、164科和285属,其中子囊菌门和担子菌门为主要菌群,占样地所有菌门的98.2%,不同恢复年限不同土层真菌群落组成存在差异。随着恢复年限的增加,研究区土壤真菌门分类水平上主要类群的个体数及类群数整体呈现增长趋势,整体高于对照样地草地。另外,恢复7年的样地土壤真菌群落丰富度及多样性显著高于其他年限和撂荒草地。冗余分析表明,研究区不同恢复年限沙棘林土壤微生物群落组成主要由土壤pH、土壤有机质含量决定,同时也与土壤氮、磷、钾元素有关。  结论  不同恢复年限沙棘人工林土壤理化性质存在差异,恢复年限的增加提高了真菌的物种丰富度及多样性。     

Exotic annuals reduce soil heterogeneity in coastal sage scrub soil chemical and biological characteristics

Exotic plant invasions alter ecosystem structure and function above- and below-ground through plant–soil feedbacks. The resistance of ecosystems to invasion can be measured by the degree of change in microbial communities and soil chemical pools and fluxes, whereas their resilience can be measured by the ability to recover following restoration. Coastal sage scrub (CSS) is one of the most highly invaded ecosystems in the US but the response of CSS soils to exotic plant invasion is little known. We examined resistance and resilience of CSS soil chemical and biological characteristics following invasion of exotic annual grasses and forbs and restoration of the native plant community. We hypothesized that invasion of exotic plant species would change biological and chemical characteristics of CSS soils by altering soil nutrient inputs. Additionally, we expected that if exotic plants were controlled and native plants were restored, native soil characteristics would recover. We sampled two locations with invaded, restored and native CSS for plant community composition, soil chemistry and microbial communities, and phospholipid fatty acid (PLFA) profiles. Communities invaded by exotic annuals were resistant to some measured parameters but not others. Extractable nitrogen pools decreased, nitrogen cycling rates increased, and microbial biomass and fungal:bacterial ratios were altered in invaded soils, and these effects were mediated by the phenological stage of the dominant plant species. The largest impact of invasion on soils was an overall reduction of spatial heterogeneity in soil nutrients, nutrient cycling and microbial communities. Restored plots tended to recover in most biotic and chemical parameters including increased resource heterogeneity compared to invaded plots, suggesting that CSS soils are resilient but not resistant to invasion.     

Impact of cattle grazing and inorganic fertiliser additions to managed grasslands on the microbial community composition of soils

A study was undertaken to determine if cattle grazing on managed grasslands had an impact on the microbial community composition of soils. Microbial community molecular profiles of bacteria, actinomycetes, pseudomonads and fungi were generated by polymerase chain reaction (PCR) amplification of rDNA sequences from community DNA isolated from soils. PCR products were profiled using denaturing gradient gel electrophoresis (DGGE) and analysed by principal co-ordinate analysis. PCR–DGGE profiles indicated that cattle grazing had an impact on the pseudomonad community structure only, and that the addition of inorganic nitrogen (N) fertiliser impacted on bacterial, actinomycete and pseudomonad community structure. There was no difference in the community profiles of fungi from grazed and N fertilised grassland plots. Analysis of phospholipid fatty acid (PLFA) profiles revealed that both cattle grazing and N fertiliser impacted on microbial community structure. The abundance of individual PLFAs differed between treatments, with bacterial (15:0), actinomycete (10Me18:0) and fungal (18:2ω6) PLFAs not affected directly by grazing cattle and N fertiliser, however, there were significant grazing–fertiliser interactions. Bacterial plate counts were highest in the N fertilised plots and fungal plate counts were highest in the cattle grazed plots. Analysis of molecular microbial community profiles with PLFA and background soil data revealed several significant correlations. Notably, soil pH was positively correlated with PCO1 of the pseudomonad community profiles and negatively correlated with the fungal PLFA 18:2ω6. Fungal DGGE profiles were negatively correlated with the fungal PLFA 18:2ω6, and bacterial and fungal plate counts positively correlated with each other. Correlation analysis using PC1 from PLFA profile data showed no significant relationship with soil organic matter, pH, total C and total N. The results indicate that cattle grazing and N fertiliser addition to grasslands impact on the community composition of specific groups of micro-organisms. The consequences of such changes in population structure may have implications regarding the dynamics of nutrient turnover in soils.     

Diversity and complexity of microbial communities from a chlor-alkali tailings dump

Revegetation of the tailings dumps produced by various industrial activities is necessary to prevent dust storms and erosion and represents a great challenge for ecological restoration. Little is known about the microbial colonisation and community structure of revegetated tailings following site exploitation. Here, we report the sequencing of 16S rRNA and internal transcribed spacer (ITS) fungal RNA gene amplicons from chlor-alkali residue and from an adjacent undisturbed soil to define the composition and assembly of the rhizosphere microbial communities. After quality filtering, a total of 72,373 and 89,929 bacterial sequences and 122,618 and 111,209 fungal sequences remained for community analysis from undisturbed soil and tailings dump samples, respectively. These reads were affiliated with 45 bacterial and 9 fungal phyla and 113 bacterial and 35 fungal classes. We observed a clear dominance of Gammaproteobacteria at our study site (24% of total sequences), especially of the Pseudomonas genera (72% of Gammaproteobacteria sequences), together with the dominance of a few fungal taxa, such as Hebeloma and Geopora. However, we also noticed that the core microbiome comprised 64.4% and 62.4% of the bacterial and fungal genera, respectively, despite marked differences in soil physico-chemical properties. A heatmap of correlations between soil parameters and taxa confirmed that approximately 50% of the 33 dominant taxa colonised both types of soil. We further demonstrated that the global bacterial-fungal network topology of the dump approximated that of the undisturbed soil. Our approach illuminates the importance of studying more than just a single component of the microbial community and represents a step forward in uncovering the microbial ecology of disturbed environments beyond what is generally found in conventional studies. Our study also provides novel global community proxies that have led us to conclude that environmental filtering is more likely to occur through the activity of tree roots rather than as a result of specific soil characteristics and could be an important force in the assembly of at least some microbial communities.     

Soil microbial community composition and land use history in cultivated and grassland ecosystems of coastal California

Phospholipid ester-linked fatty acid (PLFA) profiles were used to evaluate soil microbial community composition for 9 land use types in two coastal valleys in California. These included irrigated and non-irrigated agricultural sites, non-native annual grasslands and relict, never-tilled or old field perennial grasslands. All 42 sites were on loams or sandy loams of similar soil taxa derived from granitic and alluvial material. We hypothesized that land use history and its associated management inputs and practices may produce a unique soil environment, for which microbes with specific environmental requirements may be selected and supported. We investigated the relationship between soil physical and chemical characteristics, management factors, and vegetation type with microbial community composition. Higher values of total soil C, N, and microbial biomass (total PLFA) and lower values of soil pH occurred in the grassland than cultivated soils. The correspondence analysis (CA) of the PLFA profiles and the canonical correspondence analysis (CCA) of PLFA profiles, soil characteristics, and site and management factors showed distinct groupings for land use types. A given land use type could thus be identified by soil microbial community composition as well as similar soil characteristics and management factors. Differences in soil microbial community composition were highly associated with total PLFA, a measure of soil microbial biomass, suggesting that labile soil organic matter affects microbial composition. Management inputs, such as fertilizer, herbicide, and irrigation, also were associated with the distinctive microbial community composition of the different cultivated land use types.     

Mycorrhizal fungi associated with high soil N:P ratios are more likely to be lost upon conversion from grasslands to arable agriculture

Agriculture often leads to altered composition and reduced diversity of arbuscular mycorrhizal fungal (AMF) communities compared to natural grassland systems. However, the ecology of taxa that are lost in this transition has thus far not been well characterized. In this study we found that reduced or lost AMF taxa in farmlands were significantly stronger correlated with soil N:P ratio than a randomly sampled community; this indicates that taxa that prevail at high N:P ratio in grasslands are the ones most sensitive to agriculture. As a high N:P environment is also commonly argued to impose the highest AMF benefit to plants, the observation that those taxa are lost could indicate that agricultural fields are left with communities of reduced symbiotic quality.     

Interactive effects of elevation and land use on soil bacterial communities in the Tibetan Plateau

The Tibetan Plateau of China is uniquely vulnerable to the global climate change and anthropogenic disturbances. As soil bacteria exert a considerable influence on the ecosystem function, understanding their response to different climates and land-use types is important. Here, we characterized the bacterial community composition and diversity across three major ecosystems (cropland, forest, and grassland) in the Sygera Mountains of Tibet, along a typical elevational gradient (3 300-4 600 m). The abundance of taxa that preferentially inhabit neutral or weak alkaline soil environments (such as Actinobacteria, Thermoleophilia, and some non-acidophilus Acidobacteria) was significantly greater in the cropland than in the forest and grassland. Furthermore, the diversity of soil bacterial communities was also significantly greater in the cropland than in the forest and grassland. We observed a unimodal distribution of bacterial species diversity along the elevation gradient. The dominant phyla Acidobacteria and Proteobacteria exhibited consistent elevational distribution patterns that mirrored the abundance of their most abundant classes, while different patterns were observed for Acidobacteria and Proteobacteria at the class level. Soil pH was the primary edaphic property that regulated bacterial community composition across the different land-use types. Additionally, soil pH was the main factor distinguishing bacterial communities in managed soils (i.e., cropland) from the communities in the natural environments (i.e., forest and grassland). In conclusion, land use (particularly anthropogenic disturbances such as cropping) largely controlled soil environment, played a major role in driving bacterial community composition and distribution, and also surpassed climate in affecting bacterial community distribution.     

Community patterns of soil bacteria and nematodes in relation to geographic distance

Ecosystems consist of aboveground and belowground subsystems and the structure of their communities is known to change with distance. However, most of this knowledge originates from visible, aboveground components, whereas relatively little is known about how soil community structure varies with distance and if this variability depends on the group of organisms considered. In the present study, we analyzed 30 grasslands from three neighboring chalk hill ridges in southern UK to determine the effect of geographic distance (1–198 km) on the similarity of bacterial communities and of nematode communities in the soil. We found that for both groups, community similarity decayed with distance and that this spatial pattern was not related to changes either in plant community composition or soil chemistry. Site history may have contributed to the observed pattern in the case of nematodes, since the distance effect depended on the presence of different nematode taxa at one of the hill ridges. On the other hand, site-related differences in bacterial community composition alone could not explain the spatial turnover, suggesting that other factors, such as biotic gradients and local dispersal processes that we did not include in our analysis, may be involved in the observed pattern. We conclude that, independently of the variety of causal factors that may be involved, the decay in similarity with geographic distance is a characteristic feature of both communities of soil bacteria and nematodes.     

Soil properties, grassland management, and landscape diversity drive the assembly of earthworm communities in temperate grasslands

Earthworms are widespread soil organisms that contribute to a wide range of ecosystem services. As such, it is important to improve our knowledge, still scanty, of the factors that drive the assembly of earthworm communities. The aim of the present study was to conjointly evaluate the effects on the assembly of earthworm communities of i) soil properties (texture, organic matter content, and pH), ii) grassland management (grassland age, livestock unit, and type of fertilization), iii) landscape diversity (richness, diversity of surrounding habitats, and grassland plant diversity), and iv) presence of hedgerows. The study was conducted in temperate grasslands of Brittany, France. Earthworms were sampled in 24 grasslands and, in three of these grasslands, they were sampled near a hedgerow or near a ditch (control without a hedgerow). Soil properties explained the larger portion of the variation in the earthworm community parameters compared to grassland management or landscape diversity. The increase in soil organic matter content and pH were the most favorable factors for earthworm abundance and biomass, in particular for endogeic species. Regarding grassland management, the increase in the livestock unit was the most damaging factor for earthworm communities, in particular for the anecic earthworm biomass and endogeic species richness. Surprisingly, landscape diversity negatively affected the total earthworm abundance and epigeic earthworm biomass, but it was related to an increase in the epi-anecic species. At a finer scale, we also demonstrated that the presence of hedgerows surrounding grasslands enhanced earthworm species richness, especially within the epigeic and anecic ecological categories. This study highlights that the earthworm ecological categories respond specifically to environmental filters; further studies need to be conducted to elucidate the factors that drive the assembly of earthworm communities at this ecological category level. We recommend that policymakers should act on landscape management to favor earthworm diversity in order to improve the ecosystem services they drive.     

Rapid effects of plant species diversity and identity on soil microbial communities in experimental grassland ecosystems

Changes in plant community structure, including the loss of plant diversity may affect soil microbial communities. To test this hypothesis, plant diversity and composition were experimentally varied in grassland plots cultivated with monocultures or mixtures of 2, 3 or 4 species. We tested the effects of monocultures versus mixtures and of plant species composition on culturable soil bacterial activity, number of substrates used and catabolic diversity, microbial biomass N, microbial respiration, and root biomass. These properties were all measured 10 months after seeding the experiment. Soil bacterial activity, number of substrates used and catabolic diversity were measured in the different plant communities using BIOLOG GN and GP microplates, which are redox-based tests measuring capacity of soil culturable bacteria to use a variety of organic substrates. Microbial biomass N, microbial respiration, and root biomass were insensitive to plant diversity. Culturable soil microbial activity, substrates used and diversity declined with declining plant diversity. Their activity, number of substrates used and diversity were significantly higher in plots with 3 and 4 plant species than in monocultures and in plots with 2 species. There was also an effect of plant species composition. Culturable soil microbial activity and diversity was higher in the four-species plant community than in any of the plant monocultures suggesting that the effect of plant diversity could not be explained by the presence of a particular plant species. Our results showed that changes in plant diversity and composition in grassland ecosystems lead to a rapid response of bacterial activity and diversity.     

Diversity and distribution of nematode communities in grasslands from Romania in relation to vegetation and soil characteristics

The nematode communities of 36 grassland ecosystems in Romania, belonging to different plant associations and soil types, were studied. The abundance of nematodes, the species and trophic types present, as well as their distribution in relation to plant community and soil characteristics are analyzed and discussed.The abundance of nematodes from the 36 grasslands studied ranged between 0.41 × 10⁶ and 8.57 × 10⁶ individuals/m², and a total of 121 genera and 145 species of nematodes were found. The highest diversity was found in grasslands developed on brown earth soil (65–67 genera and 74–76 species), with least diversity in those evolving on podzol and lithosol (33–36 genera with 25–28 identified species). Most of the dominant taxa were found in specific soil layers; some obligate plant parasitic genera (e.g., Paratylenchus, Rotylenchus, Criconema) showed preference for deeper soil layers. The nematode diversity index (H′), with values ranging between 2.38 and 3.47, did not differ significantly between the different types of grasslands. Plant feeding, bacterial feeding, hyphal feeding and omnivorous nematodes were the main groups in mountainous grasslands developed on different soil types. Plant feeding and bacterial feeding nematodes dominated the trophic structure and more plant feeders (62–69%) were found in communities of subalpine and alpine grasslands developed on podzol and alpine meadow soil, than in those developed on rendzina and lithosol (27–33%). The ratio of hyphal feeding to bacterial feeding nematodes (Hf/Bf) is constantly in favour of the bacterial feeding group, the values being an indicator of good soil fertility for most studied grasslands. The nematode communities of grasslands are grouped into six main clusters according to their genera affinity and distinguished by different grassland and soil types. Communities from subalpine grasslands developed on rendzina, acid brown and lithosol have the greatest similarities. An ordination of nematode communities in relation to important environmental variables is presented. Environmental variables relevant in explaining the patterns of nematode composition in grasslands, using canonical correspondence analysis (CCA), are: humus, pH, total nitrogen, exchangeable bases and soil type. No single factor could be selected.