The effects of earthworm functional diversity on microbial biomass and the microbial community level physiological profile of soils

A novel experiment design based on the axial simplex design was used to study the effects of earthworm functional diversity on microbial biomass and microbial community level physiological profiles in soils. This mesocosm experiment was carried out at two densities of earthworm and at two levels of food supply. Soil mesocosms were maintained for 7 months and watered each week with 300 ml of distilled water. At the end of this period the soil columns were divided into upper and lower halves and microbial biomass carbon and functional diversity assessed. Greater microbial biomass and activity was found in the upper layer of soil. Food supply was the major driver leading to increased microbial activity and biomass. Increased earthworm biomass had the opposite effect. In the lower soil layer, the presence of anecic worms had the greatest effect on microbial biomass and activity. This effect was enhanced with increased food supply. Overall, the relationship between earthworm functional diversity and microbial community level physiological profile appeared to be idiosyncratic and the effect of the earthworms varied with soil layer, initial biomass and food supply. The loss of any earthworm functional group might lead to significant alterations in the microbial biomass, distribution and physiological profile.     

Efficiency of two widespread non-destructive extraction methods under dry soil conditions for different ecological earthworm groups

Reliable non-destructive extraction methods are required for the assessment of the size and composition of earthworm communities where physical disturbances are not acceptable. The aim of the present study was to investigate the efficiency of the electrical octet method and the mustard extraction method for sampling of different ecological groups of earthworms (anecics, endogeics and epigeics) under dry soil conditions. We hypothesized that: (1) the extraction efficiency of the mustard method and the octet method will vary with ecological earthworm group; and (2) beforehand water addition to dry soil will increase the extraction efficiency of the octet method but not that of the mustard method.Endogeic earthworm species were extracted in low numbers irrespective of the extraction method indicating their inactivity during dry periods. The mustard method was more efficient for the extraction of anecic earthworms even under dry soil conditions, whereas the octet method was inappropriate in reflecting the actual earthworm community structure. Surprisingly, the efficiency of both methods was not improved by beforehand water addition. These findings are essential to be considered when working under dry soil conditions e.g. in the context of environmental monitoring.     

Plant community impacts on the structure of earthworm communities depend on season and change with time

Declining plant diversity potentially threatens essential ecosystem functions driven by the decomposer community, such as litter decomposition and nutrient cycling. Currently, there is no consensus on the interrelationships between plant diversity and decomposer performance and previous studies highlighted the urgent need for long-term experiments.In the Jena Experiment we investigated the long-term impacts of plant community characteristics on the structure of earthworm communities representing key decomposers in temperate grassland. We repeatedly sampled plots varying in plant species richness (1-16 species), plant functional group richness (1-4 groups), and presence of certain plant functional groups (grasses and legumes) three, four, and six years after establishment of the experiment in spring and autumn.The results show that earthworm performance is essentially driven by the presence of certain plant functional groups via a variety of mechanisms. Plant productivity (root biomass) explained most of the detrimental grass impacts (decrease in earthworm performance), while beneficial legume effects likely were linked to high quality inputs of plant residues (increase in earthworm performance). These impacts depended on the functional group of earthworms with the strongest effects on surface feeding anecic earthworms and minor effects on soil feeding endogeic species. Remarkably, effects of plant community characteristics on the composition and age structure of earthworm communities varied between seasons. Moreover, plant diversity effects reported by a former study decreased and detrimental effects of grasses increased with time.The results indicate that plant community characteristics, such as declining diversity, indeed affect the structure of earthworm communities; however, loss of key plant functional groups is likely to be more important than plant species number per se. However, in frequently disturbed ecosystems plant species richness might be important for the recovery and resilience of belowground functions. Moreover, the results accentuate the importance of long-term repeated measurements to fully appreciate the impacts of plant community composition and diversity on ecosystem properties. Single point observations may be misleading and potentially mask the complexity of above-belowground interrelationships.     

Evenness and plant species identity affect earthworm diversity and community structure in grassland soils

Diversity of plant species and the species composition (identity) are known to influence below-ground diversity. In this paper we examine the effects of plant species diversity (richness and evenness), rates of nitrogen application and planting density, on earthworm community structure in grassland. The study was carried out at three sites in Ireland using a Simplex experimental design to define the compositions of the experimental plant communities used.A negative relationship was detected between diversity (evenness) of plant species and diversity of earthworms in the soils. However, plant species identity also affected the structure of the earthworm assemblage. In particular, the legume, Trifolium repens had a strong effect but this was conditional on the rate of nitrogen application. No earthworm species favoured communities dominated by slow growing grasses (Phleum pratense and Dactylis glomerata) (P = 0.02).Higher N inputs reduced earthworm abundance and biomass under T. repens. Earthworm richness, was negatively influenced by elevated amounts of N inputs. No effect of planting density was detected but this factor also did not affect plant biomass production.     

Earthworm communities in alluvial forests: Influence of altitude,vegetation stages and soil parameters

In many terrestrial ecosystems, soil parameters usually regulate the distribution of earthworm communities. In alluvial ecosystems, few studies have investigated the impact of periodic floods and alluvium deposition on soil fauna. In this context, we assumed that earthworm communities may vary depending on altitude (alpine, subalpine, mountain and hill levels), forest successional stage (post-pioneer to mature forests) and some soil parameters. Our results demonstrated that the composition of earthworm communities differed depending on altitudinal gradients. No earthworm was found at the alpine level while maximum density and biomass were observed at the hill level mainly due to the contribution of anecic species. A total of 27 species and subspecies were found over the three sampling sites, and Lumbricus moliboeus was discovered for the first time in carbonated soils. Soil texture had a major effect on epigeics that were often associated with coarse sandy texture in contrast to anecics which preferred deep soils and mature forest stages, which in combination provided the highest carbon content and the finest soil texture. In our study, carbonated fluviosols (Fluvisols according to the World Reference Base) were recorded; fluviosols typiques with well-structured A layers were generally found in mature or intermediate forest stages while most of fluviosols juveniles with heterogeneous texture were observed principally in post-pioneer forests. We conclude that in alluvial ecosystems, earthworm communities were highly dependent first on soil parameters, then altitude and to a lesser extent forest successional stages. Changes in earthworm communities tend to reflect a gradient of alluvial dynamics thus reinforcing the potential role of earthworms as bioindicators in natural and/or semi natural alluvial ecosystems.     

The effects of earthworm functional group diversity on nitrogen dynamics in soils

A novel experimental design, based on the simplex model, was used to study the effects of earthworm functional group diversity on nitrogen dynamics in soils. This mesocosm experiment was carried out at two densities of earthworm and at two levels of food supply. Leachate was collected and analysed over a 20-week period. Soil nitrogen content was measured when the soil columns were destructively sampled on week 30.Results showed the presence of variation among the functional groups in their effects on N dynamics and that both population densities and levels of resource availability were significant. Ammonium concentrations in leachate were generally higher with anéciques and endogées; the opposite was true for nitrate-N, which under certain circumstances was higher with epigées. A significant synergy between the endogées and anéciques was shown in terms of nitrate in soil. Earthworm biomass was important in some instances. For example, larger amounts of soil nitrate were present at higher densities of endogées. A varying effect of food supply was seen for the three functional groups. For example, there were reduced concentrations of nitrate-N in the leachate from the anecic monocultures at low levels of food supply, while increased amounts were leached from the epigées at low food supply. Greater concentrations of ammonium-N were leached from anéciques monocultures at low levels of food supply. Increased food supply resulted in increased amounts of soil nitrate-N in monocultures of both epigées and endogées. It was apparent that nitrogen transformations and amounts available in soil water are dependant on the composition of the earthworm community.     

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.     

Impact of soil stockpiling and mining rehabilitation on earthworm communities

As key ‘ecosystem engineers’, earthworms improve mineralization of organic matter, plant growth, soil quality, and are an important component of many terrestrial food webs. Under appropriate conditions, they are therefore likely to accelerate the restoration of soil ecosystem function after mining.Conserving naturally occurring populations and facilitating their recolonisation appears as the most efficient way to increase earthworms’ overall effect. The impact of mining activities and restoration measures on New Zealand endemic earthworm communities was tested. Earthworm biomass and diversity were compared in four different habitat types.Mining activities, not surprisingly, are shown here to have a detrimental impact on earthworm communities. Soil stockpiling induces anaerobic conditions at and below a depth of 1 m, where earthworms do not survive. The use of stockpiled soil for vegetation replanting therefore leads to low diversity and low abundance of earthworms. An alternative restoration technique consisting in transferring vegetation and soil units (the vegetation direct transfer) was efficient in preserving earthworm populations with earthworm biomass and diversity not significantly different from those observed in undisturbed areas. Based on these results, we recommend vegetation direct transfer (VDT) to be prioritised whenever it is logistically and economically feasible. When VDT is not applicable, low stockpiles should be prioritised as they will comprise a higher proportion of good quality soil (at the surface) and a lower proportion of anaerobic and compacted soil (below 1 m depth at the studied site).     

Ecotoxicological Assessment of Chemical Fumigants Utilising an Earthworm (<Emphasis Type="Italic">Eisenia andrei</Emphasis>) Bioassay and Soil Microbial Communities

Fumigation is an important crop protection practice employed to control soil pathogens and diseases. Metham sodium and cadusafos are two commonly used soil fumigants for this purpose. However, little information is available on their effects on non-target soil organisms. The aim of the study was to determine the ecotoxicity of these chemical fumigants on earthworms (organismal responses and DNA damage) and soil microbial communities. Changes in soil microbial community function and structure were evaluated by means of Biolog? Ecoplates and phospholipid fatty acid (PLFA) analyses, respectively. Both fumigants had a significant (p < 0.05) negative impact on all earthworm endpoints. Earthworms did not reproduce; biomass was affected negatively and manifested significant DNA damage with metham sodium causing more pronounced effects in comparison to cadusafos. The fumigants had an inhibitory effect on microbial growth. No lasting effects were observed in the community structure but cadusafos had a pronounced effect on the microbial community functional diversity. Metham sodium and cadusafos had varying effects on earthworm and microbial endpoints. This illustrates the importance of using different bioindicators to get a better understanding of the overall effects on the soil ecosystem.     

The action of an animal ecosystem engineer: Identification of the main mechanisms of earthworm impacts on soil microarthropods

Non-trophic interactions are shaping soil food web structure and functions. Particularly, the action of ecosystem engineers, such as earthworms, are likely to fundamentally impact the abiotic and biotic properties of their environment. The present study aimed to identify the main mechanisms through which earthworms belonging to varying ecological groups - epigeic, endogeic and anecic species - affect soil microarthropods by reviewing the literature on this topic and by performing meta-analyses.Earthworm ecological groups differed considerably in their impacts on microarthropods, whereas effects did not vary significantly between microarthropod taxa at the habitat scale. Inconsistent impacts of epigeic species on soil microarthropods are most likely due to differences in earthworm densities. Effects can thus be positive in the case of moderate densities or negative in the case of high densities and associated distinct changes in the physical structure of the upper soil organic layers. By contrast, impacts of endogeic earthworms appeared to be mainly negative and were primarily due to competition with microarthropods for food resources. Consequently, negative impacts on soil microarthropods intensified with increasing earthworm density and biomass. This interaction between endogeic earthworms and microarthropods is better referred to as amensalism due to the competitive predominance of earthworms. Impacts of anecic earthworm species differed significantly from that of endogeic ones; they were neutral at the habitat scale and positive on the microhabitat scale. Moreover, impacts were independent of earthworm densities due to the quasi-territorial behaviour of anecic earthworms. Positive effects were mainly attributed to the formation of stable microhabitats by anecic species; namely burrows/middens, rich in nutrients and microorganisms.The present study points to the relevance of the non-trophic biotic interactions that drive the composition of belowground food webs by identifying the most essential mechanisms underlying the impacts of animal ecosystem engineers on soil microarthropods. Moreover, as earthworms emerge as important biological invaders, the results of the present study may help to fully appreciate, estimate and model the consequences of this momentous global change phenomenon. Particularly, the spread of exotic epigeic and endogeic earthworm species likely threatens soil microarthropod density, diversity and functions.     

蚯蚓对土壤温室气体排放的影响及机制研究进展

土壤是温室气体的重要源和汇。蚯蚓是土壤物质循环的重要参与者,能够直接或间接影响土壤CO2、N2O和CH4等温室气体的产生和释放。蚯蚓呼吸产生的CO2,是土壤呼吸的重要组成部分;蚯蚓自身肠道、分泌液、消化物和排泄物等微环境促进反硝化过程释放N2O。蚯蚓还通过取食、掘穴、排泄等活动,改变土壤理化性质、微生物组成和活性及其他土壤动物的组成,影响地上植物生长,调节土壤分解、矿化、硝化、反硝化和甲烷生成及氧化等生态过程,间接影响土壤温室气体的排放。蚯蚓对土壤温室气体排放的影响逐渐受到重视,但目前研究仍以室内培养和单因子环境条件的模拟为主,缺少野外原位实验和多环境因子的交互实验研究。长期监测和同位素示踪技术,是深入探讨蚯蚓影响温室气体排放机制的重要手段。温室气体类型上,CO2和N2O是研究热点,CH4研究比较罕见。未来研究,应重视不同生态类群蚯蚓与土壤理化特征、微生物组成、其他类群土壤动物和地上植物间的交互作用,加强机制研究,并关注土壤污染环境下蚯蚓功能性状的变化;综合评价蚯蚓对土壤温室气体排放和土壤碳氮固定的影响,科学评估蚯蚓活动对土壤碳氮释放的促进或减缓作用。     

Species effects on earthworm density in tropical tree plantations in Hawaii

Summary Tree species differ in the quantity and quality of litter produced, and these differences may significantly affect ecosystem structure and function. I examined the importance of tree species in determining earthworm densities in replicated stands of Eucalyptus saligna Sm. and Albizia falcataria (L.) Fosberg, and in mixed stands (25% albizia and 75% eucalyptus). Mean earthworm densities ranged from 92 m^-2 in the pure eucalyptus, to 281 m^-2 in the mixture, and a maximum of 469 m^-2 in the pure albizia stands. Only two earthworm species were present, Pontoscolex corethrurus and Amynthas gracilis. Leaf biomass on the forest floor was highest in the pure eucalyptus and lowest in the pure albizia stands, whereas the annual fine litterfall production was lowest in the pure eucalyptus and highest in the albizia stands. The N content of fine litterfall was correlated positively with earthworm density, and the fine litterfall biomass: N ratio was correlated negatively with earthworm density. Greater leaf biomass on the forest floor under eucalyptus stands despite lower rates of litterfall suggests that litter quality, rather than litter quantity, was primarily responsible for the greater earthworm density in the albizia stands. Some biogeochemical effects of tree species in the tropics may be mediated through effects on earthworm populations.     

Impact of microarthropod biomass on the composition of the soil fauna community and ecosystem processes

The effect of soil organisms on ecosystem processes strongly depends on the composition of the overall community. Community composition however undergoes constant shifts due to pronounced spatio-temporal patterns in biomass and abundance of individual fauna groups. On this background the present experiment aimed to assess the potential impacts of shifts in the biomass of a dominant soil fauna group (microarthropods) on total community composition and on ecosystem processes mediated by fauna or microbes (e.g. decomposition, nitrogen mineralization).Microcosms, filled with spruce litter, hosted soil fauna communities that either contained ambient microarthropod biomass (control) or two elevated levels of microarthropod biomass (1.5 and 2 fold increase), while initial microbial biomass and that of other faunal groups remained unaltered. After an incubation period of 2 months, the biomass of microorganisms and fauna groups as well as ecosystem process variables were determined.The increase in microarthropod biomass at the investigated levels induced changes in the faunal community; mainly via negative or positive feeding interactions between microarthropods and the affected animal groups (Enchytraeidae, Nematoda). The abundance and activity of microorganisms at lower trophic levels however remained unaffected by these alterations; buffering the effect of shifts in the community structure on ecosystem processes.     

Indirect gradient analysis at different spatial scales of prorated and non-prorated earthworm abundance and biomass data in temperate agro-ecosystems

We investigated how earthworm communities of agricultural systems vary in abundance, biomass and species composition at different spatial scales. In four farms representative of the main agro-ecological regions of Belgium 14 parcels were sampled using a combined method (hand sorting after formalin extraction). Parcels in both grassland and arable land were studied. In each parcel two or three sample plots (1 m²) were randomly selected; these were further split up in two, resulting in four or six 0.5 m² subplots per parcel. Principal component analysis was applied to earthworm numbers and biomass. Innovative was that we compared the results from data matrices with raw data with prorated matrices, where unidentified juveniles were added to species following certain rules. To test the degree of variability between the different spatial levels, average distances between subplots, plots and parcels based on ordination scores were compared. A general linear model with a nested structure for subplots, plots, and parcels was built to find absolute differences between farms. Earthworm numbers ranged from 10 to 463 individuals m^–2, biomass varied between 3 and 186 g m^–2 and species numbers ranged from one to seven per parcel. Ordinations separated earthworm communities along a geographical—mainly soil—gradient on the first axis and distinguished between land uses along the second axis. Both in numbers of individuals and in biomass differences in earthworm populations increased from subplot to parcel level. As expected, differences within a pair of subplots were small. Plots within the same parcel tended to be highly similar, but in some cases large dissimilarities were encountered, reflecting a clustered population structure of earthworm communities at the field scale, and, possibly, abiotic gradients. The largest differences were observed at the parcel level. The latter may be attributed to differences in land use. Prorating had a narrowing effect on the data through which differences became less pronounced.     

Microbial communities and their relevance for ecosystem models: Decomposition as a case study

Soil microbial communities are so vastly diverse that complex interactions, which alter ecosystem functions, may occur among microbial species and functional groups. In this review, we explore the empirical evidence for situations when shifts in the community structure of microbes would elicit a change in ecosystem process rates, specifically decomposition, even when microbial biomass remains constant. In particular, we are interested in a subset of these scenarios in which knowledge of microbial community structure would improve model predictions for ecosystem functions. Results from microcosm and field studies indicate that microbial species diversity, functional group diversity, and community composition can all influence ecosystem process rates. The underlying mechanisms that may elicit changes in ecosystem functions from shifts in microbial community structure include evolutionary constraints on microbial trait adaptation, trait correlations, dispersal limitation, and species interactions. The extent of microbial diversity in soils is not known, so it is presently not possible to model all scenarios of microbial community structure shifts. However, by incorporating documented patterns in functional groups that are relevant for a particular ecosystem process and potential relationships between microbial phylogeny and function, the predictive power of process models will be significantly improved. The inclusion of this information into process models is critical for predicting and understanding how ecosystem functions may shift in response to global change.     

Population dynamics of earthworm communities in corn agroecosystems receiving organic or inorganic fertilizer amendments

 The dynamics of earthworm populations were investigated in continuously-cropped, conventional disk-tilled corn agroecosystems which had received annual long-term (6 years) amendments of either manure or inorganic fertilizer. Earthworm populations were sampled at approximately monthly intervals during the autumn of 1994 and spring and autumn of 1995 and 1996. The dominant earthworm species were Lumbricus terrestris L. and Aporrectodea tuberculata (Eisen), which comprised 50–60% and 8–13%, respectively, of the total annual earthworm biomass. Lumbricus rubellus (Hoffmeister) and Aporrectodea trapezoides (Dugés) were much less abundant and contributed a small fraction of total earthworm biomass. Earthworm numbers and biomass were significantly greater in manure-amended plots compared to inorganic fertilizer-treated plots during the majority of the study period. Seasonal fluctuations in earthworm numbers and biomass were attributed to changes in soil temperature and moisture, and cultivation. Unfavorable climatic conditions in the summer and autumn of 1995 caused earthworm abundance and biomass to decline significantly. Mature L. terrestris, L. rubellus and A. tuberculata were most abundant in May and June of 1995 and 1996, and cocoon production was greatest in June and July 1995 and June 1996. Recruitment of juveniles of Lumbricus spp. and Aporrectodea spp. into earthworm communities occurred primarily in the autumn. Long-term amendments of manure or inorganic fertilizer did not change the species composition of earthworm communities in these agroecosystems. The earthworm populations in both manure and inorganic fertilizer plots have declined significantly after 5 years of continuously-cropped corn. Received: 24 August 1997     

Reducing tillage in cultivated fields increases earthworm functional diversity

Alternative cropping systems such as conservation agriculture have been implemented to limit the harmful effects of intensive conventional cropping systems. Conservation agriculture is known to modify the structural diversity of earthworm communities, but no data have been reported so far on their functional diversity. Structural and functional indices of community were used to study the effects of different soil tillage intensity on earthworm diversity in arable soils.Field data were collected in four agricultural trials across France representing different soiland climatic conditions. Three types of soil tillage were assessed: plowing, superficial tillage and direct seeding. Earthworm abundance, species richness and ecomorphological group abundance were investigated. Seven functional traits, i.e. body length, body mass/length ratio, epithelium type, cocoon diameter, typhlosolis type, carbon preferences and vertical distribution, were selected according to their hypothesized link with mechanisms of tillage impact. Functional diversity indices were then computed. Soil tillage intensity decreased functional diversity and modified the functional trait profile within the earthworm community whereas neither structural diversity (species number) nor abundance changed with tillage intensity. Differences between plowing and direct seeding were significant in each trial, and superficial tillage often showed intermediate trait values. Regarding ecomorphological groups, anecic abundance was positively influenced by a decrease in soil tillage, contrary to epigeic and endogeic earthworms that showed no response. Tillage acts as an environmental filter, and decreasing its intensity caused a lesser convergence of traits and thus higher functional trait diversity. We demonstrated that a trait-based approach better permitted comparisons of community responses across sites than species number or abundance.     

Molecular and morphological analyses of faeces to investigate the diet of earthworm predators: Example of a carnivorous land snail endemic to New Zealand

Ecosystem services provided by earthworms include increasing soil fertility, recycling organic material and provision of food for numerous predatory species. Estimating the role of earthworms in food webs is essential for a thorough evaluation of the ecosystem services they provide. Although their soft bodies are rapidly digested, earthworm remains can be detected via their chaetae in the faeces of predators.This study proposes a method for the identification of predated earthworm species using only predator faeces. We conducted morphological and molecular analyses of earthworm remains in the faeces of a highly endangered carnivorous land snail (Powelliphanta augusta) endemic to a restricted area of New Zealand.Microscopic examination of faecal strings from P. augusta showed the presence of earthworm chaetae in 47 of the 49 strings examined, suggesting an important role for earthworms in snail diets. Although earthworm tissue was not detectable in snail faeces, earthworm DNA was still present in sufficient quantity for molecular identification. Results showed that this land snail feeds on at least five species of New Zealand endemic earthworms.The method developed here is particularly appropriate for investigating the diet of potential invertebrate and vertebrate predators for which feeding behaviour is difficult to observe and gut content analysis is not possible. Such studies can help to develop a more complete understanding of the ecosystem services provided by earthworm communities. In this case it also helps the conservation of a rare and endangered predatory snail species.     

Earthworm communities in temperate beech wood forest soils affected by liming

To monitor the effects of liming on forest ecosystems, experimental plots were installed in forests in mid-western Germany. In addition to soil chemical indices, earthworm communities were investigated on these plots about 15 years after first lime applications took place. As a “natural reference”, communities were compared to earthworm records that derived from a beech forest on limestone. In the non-acidified plots that had never been limed only epigeic earthworms were detected in small numbers and low species richness. Forest liming caused higher pH and a higher base saturation in the mineral topsoils. To a large extent, epigeic earthworm species seemed to benefit from this and had increased in number and biomass at all three different locations selected for the investigations. The epigeic dominated communities were completed by anecic Lumbricus terrestris that was rarely found in some of the samples from one location and a number of endogeic species that showed a very patchy distribution in limed plots. In contrast to this, the soil of the beech forest on limestone showed a different community composition. It was dominated by endogeic species in abundance and by anecic species in biomass. On limestone the total biomass of earthworms clearly exceeded the biomass values from all other plots. In conclusion, a long-term support of forest earthworm fauna due to liming was detected. This support was mainly effective for epigeic species, but in some cases for endogeic and anecic species, too.     

黄土丘陵区撂荒群落地上生物量和物种多样性关系

研究黄土丘陵区撂荒群落生物量与物种多样性的关系及其影响机制,有助于明确群落演变的机理,可对该区植被恢复重建、群落演替、生物多样性保护和维持具有重要意义。为此,选取该区5个典型撂荒群落,在群落尺度上,分析地上生物量的月动态,重点研究了地上生物量与物种多样性的关系。结果表明:(1)5个撂荒群落地上生物量总体呈上升趋势,月份之间差异极显著,群落生物量由大到小为白羊草(Bothriochloa ischaemum)群落茭蒿(Artemisia giraldii)群落铁杆蒿(Artemisia sacrorum)群落猪毛蒿(Artemisia scoparia)群落达乌里胡枝子(Lespedeza dahurica)群落;(2)群落地上生物量与物种多样性关系表现为线性增加和线性减少两种形式。猪毛蒿群落中二者呈线性增加。铁杆蒿和白羊草群落地上生物量随丰富度的增加线性增加。白羊草和达乌里胡枝子群落地上生物量随均匀度的增加线性减少。空白生态位的占有和利用、生态位互补效应、种间相互作用是解释二者关系的内在机制,冗余种假说效应可能会导致不显著关系。