Fifty years of change in Central European grassland vegetation: Large losses in species richness and animal-pollinated plants

There is growing concern that biodiversity loss in European agricultural landscapes is having negative effects on functional trait diversity. Long-term studies examining vegetation changes from the period before agricultural industrialisation are however rare. Here, we ask how management intensification and increased nutrient input initiated in the 1950/1960s have altered grassland plant community composition, species diversity and functional trait composition using comprehensive datasets from five floodplain regions (plus one protected reference region) in northern Germany. Sites with available historical relevés and vegetation maps (1950/1960s, 1990s) were resampled in 2008 to facilitate the analysis of a period spanning four to five decades.Plant community composition changed tremendously in all study regions during the 50 year period, which was related to increasing Ellenberg indicator values for nutrient availability. Species richness at the plot-level fell by 30–50% over the period, and losses in functional diversity were equally large. A non-formal comparison with the results from the protected reference study region indicates that the changes may mostly be attributable to local nutrient input rather than to supra-regional climate change. Our results indicate a consistent trend towards much more species-poor communities dominated by mow-tolerant, N-demanding competitive grasses, whereas species with more ruderal strategies, species flowering early in the season and, in particular, insect-pollinated herbs have all decreased. The substantial loss of nectar-producing grassland herbs is likely to have negative effects on the abundance of pollinating insects, with consequences for the grassland animal communities. This highlights the growing need for adequate grassland management schemes with low N input to preserve high-nature-value grassland.     

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.     

Changes in the community structure and diversity of soil invertebrates across the Franz Josef Glacier chronosequence

Following the creation of new land surfaces, there is an initial build-up phase of ecosystem development, but after a prolonged absence of major disturbance a retrogressive (decline) phase often follows due to reduced nutrient availability over time. Although many studies have considered how the soil community changes during the build-up phase, the response of this community to the retrogressive phase is poorly known. We measured litter and soil communities of microfauna and macrofauna along the Franz Josef Glacier chronosequence in New Zealand that spans ca. 120,000 years, and includes well-established build-up and retrogressive stages. We aimed to assess whether the abundances, community structure and diversity of these groups show the same pattern across the sequence as that for vegetation. With regard to microfaunal abundances, litter-dwelling microbe-feeding nematodes were most abundant in the first stage of the chronosequence, but several other groups of microfauna in both the soil and litter increased sharply during the first few stages and declined sharply during the last (retrogressive) stages. The ratios of bacterial- to fungal-feeding nematodes in both soil and litter were lowest for the final stages of the chronosequence, and (in the case of soil) for some of the early stages, pointing to domination by the fungal-based energy channel at those stages for which soil organic matter content or quality were lowest. This is consistent with the fungal-based energy channel being better adapted than the bacterial-based channel for resource-poor conditions. The main groups of macroinvertebrates typically had their lowest abundances at the very early and late stages of the chronosequence, although the relative abundances of different taxa differed during the intermediate stages. Taxonomic diversity of nematodes and macroinvertebrates in both litter and soil varied strongly with chronosequence stage but differed among taxa; diversity of only one group (macroinvertebrates in litter) declined significantly during retrogression. Diversity of nematodes and macroinvertebrates along the sequence did not closely match tree diversity or soil chemical properties, but community composition of these groups was often related to tree community composition and ratios of soil C to N, C to P and N to P. Different groups of soil invertebrates show contrasting responses to chronosequence stage, probably because they differ in their relative response to bottom-up and top-down controls. However, the abundance of most groups increased during the build-up phase and declined during retrogression. As such, the build-up and decline phases observed for plant communities and ecosystem processes across long-term chronosequences also apply to soil communities, pointing to the importance of resource availability as a major driver of soil biota during long-term ecosystem change.     

Relating microarthropod community structure and diversity to soil fertility manipulations in temperate grassland

We aimed to identify patterns of diversity in a below-ground community of microarthropods (mites and Collembola) after 15 months of a nutrient (calcium and nitrogen) manipulation experiment, located at the Natural Environment Research Council (NERC) Soil Biodiversity Site in Scotland, UK. We found that microarthropod densities increased with elevated soil fertility, but we detected no concurrent change in the diversity of soil microarthropods (mites and Collembola combined). That microarthropod density increased concurrently with improvements in soil fertility and plant productivity suggests that soil microarthropod communities are predominately regulated by bottom-up forces, driven by increased energy transfer via plant inputs to soil, providing increased food resources for fauna. However, that we found no concurrent change in the diversity of soil microarthropods provides little support for the idea that the diversity of soil fauna is positively related to their population density, primary productivity or improvements in soil conditions resulting from nutrient manipulations. However, we did find that microarthropod communities of more fertile sites contained a greater proportion of predators suggesting that more energy was transferred to higher trophic levels under elevated soil fertility. Our findings suggest that unlike plant communities, soil faunal diversity may not be strongly regulated by competition in productive situations, since competitive exclusion might not occur due to increased predation. Whilst we conclude that soil microarthropod diversity at our study site has not been affected by the nutrient additions to date, in the longer term we predict that changes in community composition and diversity could arise, most likely through top-down regulation of the soil food web.     

The restoration and re-creation of species-rich lowland grassland on land formerly managed for intensive agriculture in the UK

Intensive agriculture has resulted in the loss of biodiversity and the specialist flora and fauna associated with the semi-natural grasslands of low-intensity pastoral systems throughout northwest Europe. Techniques employed to restore and re-create these grasslands on agricultural land in the UK are reviewed. Extensive cutting and grazing management have been shown to diversify improved swards and facilitate re-colonisation on ex-arable soils, although rates of re-assembly of plant communities with affinity to existing semi-natural grasslands have generally been slow. On former agriculturally improved swards, nutrient depletion has accelerated this process, especially where “gaps” for establishment have been created. Similarly, on ex-arable soils “nutrient stripping” and sowing with diverse seed mixtures has led to the rapid development of species-rich swards. On free draining brown earths such an approach may be required to restore grassland communities where soil phosphorous concentrations exceed semi-natural levels by more than 10 mg/l (using Olsen's bicarbonate extractant). However, the appropriateness of this threshold for other soil types requires further sampling. Although restored grasslands are likely to contribute to national biodiversity targets success will ultimately depend on the reinstatement of the communities and ecological functions of semi-natural references. Although this is technically feasible for a few plant assemblages, less is known about the re-assembly of microbial and faunal communities, or the importance of trophic interactions during grassland succession. As a consequence, more research is required on the functional attributes of semi-natural grasslands, as well as the methods required to restore localised types, novel nutrient depletion techniques, the “phased” introduction of desirable but poor-performing species and the performance of different genotypes during grassland restoration.     

High colonization by native arbuscular mycorrhizal fungi (AMF) of rubber trees in small-holder plantations on low fertility soils in North East Thailand

Rubber tree is a very important crop in Thailand, representing an essential source of income for farmers. In the past two decades, rubber tree plantations have been greatly expanding in unfavorable areas, where climate conditions are difficult and soil fertility is very poor. To optimize latex yields, mineral fertilizers have been widely used. A better understanding of the roles of the biological compartment in soil fertility is essential to determine alternative management practices to sustain soil fertility and optimize latex yields. Arbuscular mycorrhizal fungi (AMF) are widely recognized as beneficial for plants, mainly through their role in improving plant nutrient uptake. The objective of this study was to assess the AMF populations in rubber tree plantations and the impact of both soil characteristics and plantation age on these communities. Our results showed that all rubber trees were highly colonized, regardless of the soil structure and nutrient contents. AMF colonization was not affected by the age of the trees, suggesting that maintaining the symbiosis is likely to be beneficial at all stages. A better understanding and management of the microbial communities would contribute to maintaining or restoring soil fertility, leading to a better tree growth and optimized latex yield.     

Functional aspects of soil animal diversity in agricultural grasslands

There has been recent interest in the characterization of soil biodiversity and its function in agricultural grasslands. Much of the interest has come from the need to develop grassland management strategies directed at manipulating the soil biota to encourage a greater reliance on ecosystem self-regulation. This review summarises information on selected groups of soil animals in grasslands, the factors influencing their abundance, diversity and community structure and their relationships to the functioning and stability of grassland ecosystems. Observations on the impacts of agricultural managements on populations and communities of soil fauna and their interactions confirm that high input, intensively managed systems tend to promote low diversity while lower input systems conserve diversity. It is also evident that high input systems favour bacterial-pathways of decomposition, dominated by labile substrates and opportunistic, bacterial-feeding fauna. In contrast, low-input systems favour fungal-pathways with a more heterogeneous habitat and resource leading to domination by more persistent fungal-feeding fauna. In view of this, we suggest that low input grassland farming systems are optimal for increasing soil biotic diversity and hence self-regulation of ecosystem function. Research is needed to test the hypothesis that soil biodiversity is positively associated with stability, and to elucidate relationships between productivity, community integrity and functioning of soil biotic communities.     

Grassland management influences spatial patterns of soil microbial communities

Soil micro-organisms play a vital role in grassland ecosystem functioning but little is known about the effects of grassland management on spatial patterns of soil microbial communities. We compared plant species composition with terminal restriction fragment length polymorphism (T-RFLP) fingerprints of soil bacterial and fungal communities in unimproved, restored and improved wet grasslands. We assessed community composition of soil micro-organisms at distances ranging from 0.01 m to 100 m and determined taxa–area relationships from field- to landscape level. We show that land management type influenced bacterial but not fungal community composition. However, extensive grassland management to restore aboveground diversity affected spatial patterns of soil fungi. We found distinct distance–decay and small-scale aggregation of fungal populations in extensively managed grasslands restored from former arable use. There were no clear spatial patterns in bacterial communities at the field-scale. However, at the landscape level there was a moderate increase in bacterial taxa and a strong increase in fungal taxa with the number of sites sampled. Our results suggest that grassland management affects soil microbial communities at multiple scales; the observed small-scale variation may facilitate plant species coexistence and should be taken into account in field studies of soil microbial communities.     

Habitat use and conservation of bumblebees (Bombus spp.) under different grassland management regimes

Declines in the natural populations of several bumblebee species across Britain and Europe are an increasing cause for concern. In this study the habitat use of bumblebees was investigated on Salisbury Plain Training Area, the largest remaining area of unimproved chalk grassland in north-west Europe. Habitat characteristics influencing the overall abundance, species richness and foraging activity of bumblebees included the diversity and abundance of flowering plant species (particularly of favoured forage plants such as Trifolium pratense), vegetation structure and height. It is suggested that different Bombus species respond to these habitat characteristics depending on their specific foraging and nesting requirements, the case of Bombus humilis being especially relevant. The effects of several grassland management practices were considered in terms of their suitability for the conservation of bumblebee habitats. Cattle grazing was shown to be preferable to both sheep grazing and the absence of any management, although the timing and intensity of such grazing was important. Small-scale disturbances caused by vehicle activity were also of value in producing locally abundant forage resources in less intensively managed grasslands.     

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     

Recovery of soil macronutrients following shifting cultivation and ethnopedology of the Adi community in the Eastern Himalaya

Shifting cultivation involves a cycle of forest clearing, cultivation and a fallow phase. As the practice involves clearing forest, it is considered unsustainable and leads to soil fertility loss and erosion. While several variations of the practice exist, traditional communities undertake the practice systematically with relatively long fallow periods and are often knowledgeable about their landscape in terms of soil and its management. To better understand one such system, we quantified soil recovery following cultivation in terms of macronutrients and documented the traditional knowledge of the Adi community in a remote site in the Eastern Himalaya. We collected soil samples from three replicates, each from currently cultivated sites, uncut forest sites and successional sites 3, 12 and 25 yr following cultivation. Available nitrogen and phosphorus significantly increased, and there was an increasing trend in soil organic matter following cultivation. The Adi differentiated nine types of soil and preferred specific soil types for shifting and settled cultivation. We documented soil management and methods of soil fertility retention practised by the Adi. Their location of different crops in the field was based on the effect of the crop on soil fertility. Our research indicated that soil nutrient recovery was considerable following cultivation and that traditional shifting cultivators in the landscape were knowledgeable about their landscape in terms of soil diversity, undertaking practices to manage soil erosion and fertility. Future policies that will affect shifting cultivation in the region should acknowledge such systematic use of a landscape by traditional farming communities.     

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.     

Shrubs influence arbuscular mycorrhizal fungi communities in a semi-arid environment

Interactions between arbuscular mycorrhizal fungi (AMF) and plants are essential components of ecosystem functioning; however, they remain poorly known in dry ecosystems. We examined the relationship between seven shrub species and their associated AMF community in a semi-arid plant community in southern Spain. Soil characteristics and plant physiological status were measured and related to AMF community composition and genetic diversity by multivariate statistics. We found differences in AMF communities in soils under shrubs and in gaps among them, whereas no differences were detected among AMF communities colonizing roots. Soil nutrients content drove most of the spatial variations in the AMF community and genetic diversity. AMF communities were more heterogeneous in fertile islands with low nitrogen-to-phosphorus ratio and vice versa. AMF genetic diversity increased in soils limited by phosphorus and with high soil organic matter content, while AMF genetic diversity increased in roots growing in soil not limited by phosphorus. Overall, we could not find a clear link between plant performance and the associated AMF community. Our findings show that different shrub species generate islands of fertility which differ in nutrient content and, therefore, support different AMF communities, increasing AMF diversity at the landscape level.     

放牧管理对青藏高原东缘高寒草甸土壤线虫的影响及作用机制

不同强度、季节的放牧是草地生态系统中主要的放牧管理措施,在生物多样性维持以及生态过程发挥中起着重要的作用,然而,关于青藏高原东缘高寒草甸地区放牧对土壤线虫群落的研究甚少。本文调查了放牧（轻度、中度和重度）对植物群落、土壤理化性质和线虫群落的影响。结果表明：植物群落、土壤理化性质和线虫群落受到放牧、时间以及放牧和时间共同作用的影响;食细菌线虫、植物凋落物生物量、根生物量、土壤含水量、土壤有机碳含量在轻度放牧草地内最高（p＜0.05）;杂类草生物量在中度放牧草地内最高;杂食线虫数量在重度放牧地内最低（p＜0.05）。植物群落和土壤特性与土壤线虫群落有明显的相关关系（p＜0.05）。结构方程模型显示食草动物主要通过植物群落影响植食和食细菌线虫,进一步的研究应针对植物群落多样性以及个体特征对线虫群落的影响。     

Changes in agricultural management drive the diversity of Burkholderia species isolated from soil on PCAT medium

In order to assess the diversity of culturable Burkholderia populations in rhizosphere and bulk soil and to evaluate how different agricultural management regimes and land use history affect this diversity, four treatments were evaluated: permanent grassland; grassland converted into maize monoculture; arable land and arable land converted into grassland. Burkholderia isolates obtained on PCAT medium were grouped in 47 clusters using 16S ribosomal RNA gene based PCR-DGGE combined with BOX genomic fingerprinting (DGGE-BOX). The distribution of the isolates in the DGGE-BOX clusters was used to calculate the Shannon diversity index per treatment. Interestingly, we observed that the Burkholderia diversity was affected by changes in the agricultural management, since the highest diversity was observed in permanent grassland and in continuous arable land. In addition, the diversity tended to be higher in the rhizosphere than in the corresponding bulk soil. The use of species abundance models indicated that rhizosphere communities had more even distributions than communities collected from the bulk soil. Identification of isolates revealed that only 2% of these belonged to the B. cepacia complex and that the majority was assigned to either (1) new Burkholderia species or (2) Burkholderia species that had originally been isolated from soil. Isolates classified as B. hospita, B. caledonica and Burkholderia sp. ‘LMG 22934’ and ‘LMG 22936’ were found mainly in the arable land, while isolates belonging to Burkholderia sp. ‘LMG 22929’ and B. phytofirmans were associated with the grassland area. Another potentially new Burkholderia species, ‘LMG 22932’, was found in both areas, in close association with the maize rhizosphere.     

Management effects on the vegetation and soil seed bank of calcareous grasslands: An 11-year experiment

Calcareous grasslands, most of which are man made and therefore depend on some kind of human interference for their maintenance, are among the most species-rich communities on Earth at a small scale. For many centuries, most of these grasslands have been used as extensive pasture. However, after 1900, and particularly from 1940 onwards, livestock grazing has declined throughout Europe leading to the abandonment of low intensity grasslands over large areas. To conserve the remaining grasslands or to restore recently abandoned grasslands, better insights about the effects of grassland management on above and belowground species diversity are needed. Here, we describe the results of an 11-year experiment to investigate the role of grassland management (grazing, mowing and abandonment) in determining species composition and diversity both in the aboveground vegetation and the seed bank of a calcareous grassland in Belgium. Species diversity declined by about 60% 11 years after abandonment, from 29 species m⁻² to as few as 12 species m⁻². Plots that were grazed remained constant in species richness, whereas mown sites lost about 20% of their original species. Abandoned plots were largely dominated by a few grass species, in particular Festuca rubra. Concomitant with changes in the aboveground vegetation, both the number of species found in the seed bank and seed density (number of seeds m⁻²) had changed significantly 11 years after abandonment. Species diversity and seed density were significantly lower in abandoned plots than in grazed or mown plots. We conclude that abandonment of calcareous grasslands may lead to rapid decline of plant species diversity both in the aboveground vegetation and in the seed bank. As a result, seed banks probably have a limited role to play in the restoration of recently abandoned grasslands.     

Can a mesotrophic grassland community be restored on a post-industrial sandy site with compost made from waste materials?

Restoration of sites degraded by industry to species-rich semi-natural vegetation communities is difficult; it usually involves the addition of soil ameliorants but excessive fertility may favour dominance by competitive species. In a field-experiment we tested the establishment of a biodiverse mesotrophic grassland community using different compost types (comprising of mixtures of waste materials), application rates and seeding (with species in the target community). Compost addition to the alkaline sandy substrate increased soil organic matter, nutrient content and water holding capacity (WHC), whilst decreasing pH. Over the first two growing seasons compost addition, (especially at a higher rate) increased total vegetation cover (from <20% to a maximum of 67%), although the cover of the target community remained below 20%. Seeding with target species greatly increased their establishment on compost-treated plots, demonstrating its value for restoration of mesotrophic grassland communities in such sites lacking a local seed source. Five soil properties accounted for 46% of the variation in target species density: negative correlations with soil pH and %N, and positive correlations with electrical conductivity (EC), %C, and WHC. For this mesotrophic grassland community, high EC and WHC and low pH were most important for forb species and high %C for grasses. Overall, %C was the soil property that best explained variation in the early restoration success of different compost types and application rates; pH and EC were also correlated with the rate of vegetation establishment and available-P was linked to plant community composition. While a longer time period is needed to judge the sustainability of the outcome, this demonstrates the potential to refine compost properties for restoration of biodiversity.     

Increasing dominance of Brachypodium pinnatum (L.) beauv. in chalk grasslands: A threat to a species-rich ecosystem

Chalk grassland is a semi-natural, species-rich vegetation which was once widespread in West Europe. Due to the high species diversity a number of remnants of this vegetation have been set aside as nature reserves and managed by man to prevent natural succession into woodland.

During the last decade an increase of dominance of the grass Brachypodium pinnatum has been observed in chalk grassland within the greater part of the distribution area. Increasing dominance is strongly negatively correlated with the species diversity. Increase of this species is observed in grassland with different management regimes, e.g. sheep grazing, mowing or burning. Brachypodium can gain 80% or more of the relative above-ground phytomass. In such vegetation the species number decreases to less than 50% of the original number within a few years—the low growing forbs especially disappear almost completely. It is argued that the impact of nutrients by air pollution, especially of nitrogen, causes the increasing dominance of Brachypodium. Further research will focus on the effects of this extra nutrient supply.

The aim of the present research is to formulate management strategies in order to prevent degradation of the species diversity in chalk grasslands.