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Soil structure is closely linked to biological activities. However, identifying, describing and quantifying soil aggregates remain challenging. X‐ray micro computed tomography (X‐ray μCT) provides a detailed view of the physical structure at a spatial resolution of a few microns. It could be a useful tool to discriminate soil aggregates, their origin and their formation processes for a better comprehension of soil structure properties and genesis. Our study aims to (a) determine different X‐ray μCT‐based aggregate parameters for differentiating earthworm casts belowground (earthworm aggregates) from aggregates that are not formed by earthworms (non‐earthworm aggregates), and (b) to evaluate if these parameters can also serve as specific “tomographic signatures” for the studied earthworm species. For this purpose, we set up a microcosm experiment under controlled conditions during 8 weeks, including three species of earthworms tested separately: the epigeic Lumbricus rubellus, the anecic Lumbricus terrestris and the endogeic Allolobophora chlorotica. Our results show that X‐ray μCT analysis helps distinguish earthworm aggregates from non‐earthworm ones using (a) the relative volume of the components within aggregates and (b) the volumetric mass of aggregates and their global volume. In particular, the volume ratio of mineral grains within the aggregates is significantly different according to earthworm species. So, X‐ray μCT is a powerful and promising tool for studying the composition of earthworm casts and their formation. However, future research is needed to take into account the shapes and spatial distribution of the aggregates' components, in particular the different states of organic matter decomposition. HIGHLIGHTS: Can earthworm belowground casts be differentiated from other soil aggregates using X‐ray μCT? The use of X‐ray μCT for characterizing soil aggregates at a spatial resolution of a few microns. A combination of X‐ray μCT variables discriminates earthworm casts from non‐earthworm aggregates. X‐ray μCT, used alone, is relevant for defining species‐specific signatures of earthworm casts.

We carried out mesocosm experiments using either the anecic earthworm Lumbricus terrestris or the endogeic earthworm Allolobophora chlorotica and loam, silt loam and sandy loam soils to investigate the differing impact of these earthworm of different ecotypes on aggregate formation (percentage water stable aggregates, %WSA) and soil water holding capacity (WHC), two soil properties that underpin many of the ecosystem services provided by soils. Earthworms significantly increased %WSA (by 16–56% and 19–63% relative to earthworm-free controls for L. terrestris and A. chlorotica, respectively). For L. terrestris, this increase was significantly greater in the upper 6.5 cm of the soil where their casts were more obviously present. Allobophora chlorotica treatments significantly increased WHC by 7–16%. L. terrestris only caused a significant increase in WHC (of 11%) in the upper 6.5 cm of the sandy loam soil. Linear regression indicated a consistent relationship between increases in %WSA and WHC for both earthworm species. However, for a given %WSA, WHC was higher for A. chlorotica than L. terrestris likely due to the known differences in their burrow structure. Overall, earthworms increased soil %WSA and WHC but the significant species/ecotype differences need to be considered in discussions of the beneficial impacts of earthworms to soil properties.

Earthworms are an integral part of soil ecosystems, especially for their role in soil functions such as organic matter (OM) decomposition and nutrient cycling. Earthworms and microorganisms are interdependent, and a considerable portion of the contribution earthworms make to influencing OM fate is through interactions with microorganisms. However, the importance of the earthworm-associated microbiome is not fully understood, because it is difficult to separate the direct influence of the earthworms from the indirect influence of their microbiome. Here, we evaluated an antibiotic-based procedure to suppress the microbiome of individuals of ecologically-contrasting earthworm species (Eisenia fetida, Lumbricus terrestris, Allolobophora chlorotica) as the first step towards soil studies aimed at understanding the importance of the earthworm microbiome for host health and function. Individual earthworms were exposed to antibiotics: cycloheximide (150 μg ml−1), ampicillin (100 μg ml−1), ciprofloxacin (50 μg ml−1), nalidixic acid (50 μg ml−1), and gentamicin (50 μg ml−1) either singly or in a cocktail via culture (96 h) in a semi-solid agar carrier. Compared to the non-antibiotic treated control, the cocktail (for all three species) and ciprofloxacin (for E. fetida and A. chlorotica) treatments significantly reduced (P < 0.05) culturable microbial abundance on nutrient agar and potato dextrose agar. The microbial counts were reduced to below detection (

Sulfonated and hydroxy-sulfonated PCBs were recently discovered by our group as new PCB soil contaminants, constituting about 1% of their parent compounds in soil. Here we investigate for the first time the bioaccumulation of these metabolites as well as hydroxy-PCBs and native PCBs in earthworms. A sequence of three experiments, at increasing complexity and ecological realism, were performed with four different earthworm species (Eisenia foetida Savigny, Lumbricus terrestris L, Allolobophora chlorotica Savigny and Aporrectodea caliginosa Savigny) exposed to contaminated soils. The first experiment confirmed that when exposing earthworms to soil contaminated with a single hexa-chlorinated congener (PCB 155), no formation of polar metabolites in earthworms could be detected. This allowed to plan the following two experiments, using a soil from a PCB contaminated site and rich in relatively high levels (10–130 μg kg−1) of hydroxy-, sulfonated-, and hydroxy-sulfonated-PCBs. Bioaccumulation factors (BAFs) and bioconcentration factors (BCFs) were then obtained in the second and third experiments, to compare the accumulation behavior of these chemicals in laboratory and natural conditions. Regressions between BAF/BCF and Log Kow/Log D, produced a variety of results, being generally significant between BCF and PCBs and not significant in the other cases. In general, the metabolites accumulated in earthworms with detectable concentrations in their tissues (8–115 μg kg−1), although sulfonated and hydroxy-sulfonated PCBs showed BAF and BCF values lower (up to two orders of magnitude) than those calculated for the parent PCBs, given their lower lipophilicity.

Interactions of earthworm species with their environment and in particular the soil they inhabit is poorly understood. In this study, we aimed to use mid infrared absorption (MIR) spectroscopy of earthworm bodies in combination with discriminant analysis of principal components (DAPC), in order to investigate if their biogeochemical composition is determined by their species belonging or the soil they inhabit. Our conceptual approach included the comparison of MIR spectra of four earthworm species (Lumbricus terrestris, Allolobophora chlorotica, Aporrectodea caliginosa and Aporrectodea icterica) sampled from four different agricultural fields in France. Moreover, we tested the influence of the habitat on the earthworm MIR spectra by exchanging individuals and soils in an incubation experiment. MIR spectra varied according to species and sampling site. The MIR spectra of earthworm bodies also changed during the incubation experiment, and for L. terrestris, A. chlorotica and A. icterica depended on the properties of the soil ingested. This suggested that the food source in different habitats impacted the biogeochemical composition of earthworm bodies. DAPC allowed for the identification of earthworm species ranging from 64 to 100% of accuracy (mean 78%). We thus conclude that MIR spectroscopy may be a suitable approach to identify earthworm species, but that their signature may also allow to investigate their ecology as determined by the soil they inhabit.

It is generally accepted that populations of most species experience some level of density dependence; however, this has rarely been shown for soil-dwelling invertebrate species, in particular for earthworms. Experimental data, mostly obtained at high laboratory densities, suggest intense intraspecific competition for resources and/or living space and density-dependent dynamics in earthworm populations, irrespective of their ecological group affiliation. In the present study performed in large field microcosms, I investigated whether density-dependent responses occur within the earthworm density gradients more realistic for the natural sites. Five lumbricid species from epigeic ( Dendrobaena octaedra , Lumbricus rubellus ), endogeic ( Allolobophora chlorotica , Aporrectodea caliginosa ) and anecic ( Lumbricus terrestris ) ecological groups were tested. Soil systems populated with earthworm monocultures in large (20-L) microcosms were exposed in a beech-oak forest for 4.5 months; each species was represented by two ( L. terrestris ) or three treatments forming gradients of increasing density. In two endogeic species, manifold and generally similar density-dependent responses (a retardation of growth, maturation and reproduction rates, but higher mortality rates with density increase) were revealed, likely explained by spatial competition rather than by direct food competition. In each of epigeic species density-dependent responses were less variable and more species-specific. In contrast to endogeics, direct food competition was presumably a more important cause of density-dependence. In L. terrestris no significant density-dependent responses in adult earthworms were revealed; however, they need to be further investigated in relation to the age and territorial behaviour of individual earthworms. Importantly, in any earthworm species the density variations in the reproducing generation had significant consequences for the advancing generation, affecting either the numbers or/and the size (individual weight) of the cocoons produced. It is suggested that an underestimation of density-dependent processes may cause inaccurate estimates of the activities of local lumbricid populations.

Phosphorus (P) is an important nutrient for plant growth. However, P is often poorly available for uptake by roots because it strongly adsorbs to the soil mineral phase. Recent research shows that earthworms can temporally and locally increase P availability to plants. However, the pathways through which they do so are not fully understood, and it remains unclear to what extent this capacity varies among earthworm species. Here we study the variation among earthworm species with respect to readily-available P in casts as well as other physico-chemical cast properties, in a greenhouse pot experiment using a soil with a low P status. The earthworms belong to eight commonly occurring earthworm species in the Netherlands: two epigeic species (a mixture of the compost earthworms Dendrobaena veneta/Eisenia fetida; Lumbricus rubellus); four endogeic species (Allolobophora chlorotica, Aporrectodea caliginosa, Aporrectodea rosea, Octolasion lacteum); and two anecic species (Aporrectodea longa; Lumbricus terrestris). For all species, the pH in water extracts of earthworm cast (pH = 7.4–8.2) was significantly higher (p < 0.001) than for the control bulk soil (pH = 6.6) and differed significantly (p = 0.003) among earthworm species. Similarly, the dissolved organic carbon (DOC) concentration in the same water extracts was an order of magnitude higher in earthworm cast compared to the control bulk soil and varied among species (p < 0.001). The size of the total pool of reversibly adsorbed P in earthworm cast was greater than in the control bulk soil, but no significant differences were found among earthworm species. Differences among species were present for the readily-available P pools extracted from casts, including P-Olsen and water-extractable ortho-P. Water-extractable ortho-P concentrations were much higher in the casts of all species as compared to the control bulk soil (0.9–6.8 vs 0.06 mg l−1 or to 9.0–68 vs 0.6 mg kg−1). Highest ortho-P levels were measured in L. rubellus casts and the lowest in casts of A. chlorotica. A positive correlation between the concentrations of DOC and ortho-P was observed (R2 = 0.72, p < 0.001). The observed variation in all measured physico-chemical cast properties could not be explained by conventional ecological earthworm classifications. Our results show that the nature and magnitude of earthworm-induced increased P availability differs dramatically among earthworm species. This strongly suggests that, apart from its size, species composition of the earthworm community is key to optimizing P availability to plants.

While many studies have examined the effects of flooding on earthworm population distributions, few studies have investigated physiological and behavioural responses of earthworms to the low oxygen conditions caused by flooding. An earthworm's skin is its oxygen exchange organ, allowing earthworms to survive in flooded environments provided that the water contains sufficient dissolved oxygen. Individuals of three species of earthworm, the anecic Lumbricus terrestris (Linneaus, 1758), the green morph of the endogeic Allolobophora chlorotica (Savigny, 1826) and the epigeic Lumbricus castaneus (Savigny, 1826) were placed in reconstituted groundwater that was either kept aerated or kept in a sealed container so that dissolved oxygen was gradually consumed as the earthworm respired. Oxygen saturation of the water was measured over time in sacrificial triplicate replicates from each treatment at discrete time points; earthworm mortality and mass were recorded. Before treatments, oxygen levels in all treatment tubes were 9.53 (±0.64) mg O2 L−1. L. terrestris, a large species which emerges at night to forage at the soil surface died when oxygen levels reached 0.82 (±0.46) mg O2 L−1 after approximately 36 h. L. castaneus, a smaller species which lives on the soil surface, died when oxygen levels reached 3.60 (±2.01) mg O2 L−1 after approximately 168 h. A. chlorotica, which is similar in size to L. castaneus, lives in the upper 20 cm of soil and is known to aestivate during the summer, did not die, even when oxygen levels reached 1.49 (±0.40) mg O2 L−1 after 280 h. The results suggest that earthworm respiration is closely linked to both body size and to behavioural ecotype. These findings suggest that if flooding increases in frequency resulting in episodic reductions in soil oxygen levels, the species composition of earthworm communities may change, with an increased presence of endogeic earthworms which show a responsive plasticity to flooding events.

We performed a greenhouse experiment to test how the functional diversity of earthworms, the dominant group of soil macro-invertebrates in many terrestrial ecosystems, affects nitrogen cycling and plant growth. Three species were chosen to represent a range of functional traits: Lumbricus terrestris (large, mainly detritivorous, makes vertical permanent burrows open at the surface), Aporrectodea longa (medium-large, feeds on both detritus and soil, makes burrows more branched than L. terrestris), and Allolobophora chlorotica (small, geophagous, makes ephemeral burrows below the soil surface). Mesocosms with ryegrass (Lolium perenne) were inoculated with none to all three species (similar total biomass), using an experimental design (Simplex) suited to partition single species and diversity effects. Two contrasting N sources, urea or mammalian dung, were labelled with 15N so that the acquisition by plants and earthworms and recovery of applied 15N could be estimated. Over 3 months, plant production was higher with urea applications, but there were also species-specific earthworm effects: A. chlorotica and, to a lesser extent, A. longa increased shoot biomass, whereas L. terrestris increased root biomass. Earthworms did not affect soil N concentrations or leaching losses, whereas more N was leached under urea. A. chlorotica tended to increase dung-15N recovery in grass shoots, but in interaction with A. longa had the opposite effect, possibly through increased N immobilization in the microbial biomass. Earthworms assimilated negligible amounts of urea-15N but a substantial proportion (17% on average) of the dung-15N, with no clear-cut differences between species. Our findings show that earthworm species may have similar trophic response to N sources and yet different effects on plant N uptake and primary production, and that inter-specific earthworm interactions can result in non-additive diversity effects.

By means of a literature survey, earthworm species of significant relevance for soil functions in different biogeographical regions of Europe (Atlantic, Boreal, Mediterranean) were identified. These focal earthworm species, defined here according to the EFSA Guidance Document on the environmental risk assessment (ERA) of genetically modified plants, are typical for arable soils under crop rotations with maize and/or potatoes within the three regions represented by Ireland, Sweden and Spain, respectively. Focal earthworm species were selected following a matrix of four steps: Identification of functional groups, categorization of non-target species, ranking species on ecological criteria, and final selection of focal species. They are recommended as appropriate non-target organisms to assess environmental risks of genetically modified (GM) crops; in this case maize and potatoes. In total, 44 literature sources on earthworms in arable cropping systems including maize or potato from Ireland, Sweden and Spain were collected, which present information on species diversity, individual density and specific relevance for soil functions. By means of condensed literature data, those species were identified which (i) play an important functional role in respective soil systems, (ii) are well adapted to the biogeographical regions, (iii) are expected to occur in high abundances under cultivation of maize or potato and (iv) fulfill the requirements for an ERA test system based on life-history traits. First, primary and secondary decomposers were identified as functional groups being exposed to the GM crops. In a second step, anecic and endogeic species were categorized as potential species. In step three, eight anecic and endogeic earthworm species belonging to the family Lumbricidae were ranked as relevant species: Aporrectodea caliginosa, Aporrectodea rosea, Aporrectodea longa, Allolobophora chlorotica, Lumbricus terrestris, Lumbricus friendi, Octodrilus complanatus and Octolasion cyaneum. Five out of these eight species are relevant for each biogeographical region with an overlap in the species. Finally, the earthworm species Ap. caliginosa (endogeic, secondary decomposer) and L. terrestris (anecic, primary decomposer) were selected as focal species. In the Mediterranean region L. terrestris may be substituted by the more relevant anecic species L. friendi. The selected focal species are recommended to be included in a standardized laboratory ERA test system based on life-history traits. © 2016 Elsevier B.V.

International audience; Recent EFSA (European Food Safety Authority) reports highlighted that the ecological risk assessment of pesticides needed to go further by taking more into account the impacts of chemicals on biodiversity under field conditions. We assessed the effects of two commercial formulations of fungicides separately and in mixture, i.e., Cuprafor Micro® (containing 500 g kg−1 copper oxychloride) at 4 (C1, corresponding to 3.1 mg kg−1 dry soil of copper) and 40 kg ha−1 (C10), and Swing® Gold (50 g L−1 epoxiconazole EPX and 133 g L−1 dimoxystrobin DMX) at one (D1, 5.81 10−2 and 1.55 10−1 mg kg−1 dry soil of EPX and DMX, respectively) and ten times (D10) the recommended field rate, on earthworms at 1, 6, 12, 18 and 24 months after the application following the international ISO standard no. 11268-3 to determine the effects on earthworms in field situations. The D10 treatment significantly reduced the species diversity (Shannon diversity index, 54% of the control), anecic abundance (29% of the control), and total biomass (49% of the control) over the first 18 months of experiment. The Shannon diversity index also decreased in the mixture treatment (both fungicides at the recommended dose) at 1 and 6 months after the first application (68% of the control at both sampling dates), and in C10 (78% of the control) at 18 months compared with the control. Lumbricus terrestris, Aporrectodea caliginosa, Aporrectodea giardi, Aporrectodea longa, and Allolobophora chlorotica were (in decreasing order) the most sensitive species to the tested fungicides. This study not only addressed field ecotoxicological effects of fungicides at the community level and ecological recovery, but it also pinpointed some methodological weaknesses (e.g., regarding fungicide concentrations in soil and statistics) of the guideline to determine the effects on earthworms in field situations.

International audience; Aims : The passage of seeds through the earthworm gut potentially damages seeds, altering seed and seedling performances depending on seed traits. This work was conducted to study to what extent chemical and morphological seed traits determine the seed attractiveness for earthworms. Methods : We tested seed selection via the ingestion and digestion of 23 grassland plant species spanning a range of 14 morphological and chemical traits by two common earthworm species: the anecic Lumbricus terrestris and the endogeic Allolobophora chlorotica. Results : Both earthworm species ingested seeds from all plant species. A. chlorotica digested almost all ingested seeds (out of the 15 % ingested), whereas L. terrestris excreted them in varying quantities (out of the 86 % ingested), depending on plant species identity. Seed ingestion rate by L. terrestris was driven by seed oil content and earthworm initial weight. The apparent effect of seed length was explained via seed oil content. Seed digestion rate by L. terrestris was negatively impacted by seed size. Seed ingestion rate by A. chlorotica tended to be impacted by seed protein content and seed length. Conclusion : Earthworms–seed interactions depend on a variety of seed traits and earthworm identity. Thus, earthworms, via their specific feeding behavior, might facilitate or impede the regeneration of certain plant species and drive plant communities.

Through their manipulation of soil physical and chemical properties, earthworms have the potential to increase nutrient mineralization and benefit crop production. However, this effect is dependent on numerous factors including earthworm densities and ecological strategies. In addition to impacting density, soil environmental conditions may limit earthworm activity to specific times during the growing season, moderating their effects on nutrient availability and crop growth. The purpose of this study was to determine the active period of earthworms to better assess their potential impact on crop productivity in dryland wheat production systems in the Palouse region of northern Idaho and eastern Washington, U.S.A. Earthworm community density, age structure, diversity, and soil moisture and soil temperature were monitored over a 14-month (April 2014–June 2015) period in six dryland wheat fields. All earthworms collected were exotic species, with the endogeic species, Aporrectodea trapezoides being predominant at all sites (87% of all adults identified). Allolobophora chlorotica, Lumbricus terrestris and Aporrectodea tuberculata were collected at lower frequencies (1.8–6.4% of adults). Mean densities ranged from 14 to 75 individuals m−2, and populations transitioned to aestivation beginning in mid-June of both years. All earthworms were active (no aestivating individuals) on 56.9% of the sampling dates in 2014 and 71% of the days in 2015. A short active period and average densities at most sites of less than 100 individuals per m2 suggest that the impact of earthworms on crop growth may be limited in conventional, dryland agricultural fields of this region.

Short Rotation Forestry (SRF) practice which includes use of rapidly growing native and non-native tree species has been introduced to the UK as a method to increase woody biomass production. A largely unknown aspect of SRF is the quality of leaf litter, and its palatability to the soil decomposer community of which earthworms are a major component. The aims of the present study were to investigate and compare the preference of selected native British earthworms for selected SRF species litter. These were addressed through a series of controlled laboratory experiments. Choice chambers were used to quantify litter removal by Allolobophora chlorotica, Aporrectodea caliginosa, Aporrectodea longa and Lumbricus terrestris over a period up to five weeks. In addition, an infrared webcam recording technique was used to directly observe litter selection behaviour of L. terrestris under cover of darkness. Choice chamber experiments revealed that earthworms significantly preferred (p

Within the regulatory framework for authorisation of plant protection products (PPPs) (EU Directive 91/414/1991/EEC replaced by Regulation (EC) 1107/2009), higher tier risk assessments and earthworm field tests are conducted in different countries across Europe. This paper describes dominant earthworm species for regulatory and biogeographical regions in agricultural landscapes across Europe and examines regional differences in earthworm communities and densities and their respective response to a toxic reference. For the assessment of earthworm abundance and species distribution, data of untreated control plots from 30 earthworm field studies were analysed; each conducted according to the ISO 11268–3 (1999) guideline by European Crop Protection Association member companies in the context of registration of PPPs. For the evaluation of the response to PPPs under different regional and climatic conditions, the effect on earthworm abundance was assessed by comparing plots treated with toxic references with untreated control plots. Additionally, a comparative literature review was included providing an overview of earthworm species composition and densities in agricultural crops from 14 European countries. The assessment of earthworm field studies from six different European countries revealed that common earthworm species of anecic and endogeic ecological groups are present at most field sites. Dominant species groups of endogeic and anecic earthworms can be defined that are abundant in all assessed countries. These are the endogeic species Aporrectodea caliginosa, Aporrectodea rosea and Allolobophora chlorotica, and the anecic species Lumbricus terrestris (Northern and Central Europe) and Lumbricus friendi (Southern Europe). Taking into account the high variability in total earthworm abundances, it can be concluded that the variability within regions was larger than the variability between regions. Analysis of the earthworm community and data of toxic references lead to the conclusion that testing in different zones is not considered necessary.

We have shown previously that riboflavin is stored in chloragosomes of chloragocyte-derived earthworm eleocytes what may have an adaptive value for worms vulnerable to soil-derived pathogen invasion. Thus, it is important to determine whether riboflavin stores are also present in earthworm species with low eleocyte counts, especially in the chloragocytes of their chloragogen tissue that surrounds the coelomic site of the intestine, and find out its putative roles in earthworm immunity. Experiments were performed on adult worms from 6 species. Freely floating coelomocytes (consisting of amoebocytes plus species-specific numbers of eleocytes) were extruded by mild electric shock and then chloragocytes from the same anaesthetized worm were mechanically detached into buffer. Both cell suspensions were analyzed by spectrofluorometry for riboflavin content. It turned out that riboflavin is stored in species-specific quantities in all investigated species. Riboflavin storage predominates in free coelomocytes of eleocyte-rich species (Eisenia andrei, Dendrobaena veneta, Allolobophora chlorotica) while in chloragocytes of species with few eleocytes (Lumbricus terrestris, L. rubellus, and Aporrectodea caliginosa). Upon a massive microbial impact, the coelomocytes (both amoebocytes and eleocytes) are involved in the formation of multicellular bodies encapsulating soil-derived pathogens, what is connected with enhanced coelomocyte mobility. Thus the second aim of the present investigation was to check if riboflavin can participate in coelomocyte accumulations. Tests performed in a 48-well chemotaxis chamber revealed that riboflavin behave as chemoattractant for coelomocytes of all investigated earthworm species, which may be one of mechanisms underpinning its immunomodulatory functions.

The introduction of earthworms into soils contaminated with metals and metalloids has been suggested\ud to aid restoration practices. Eisenia veneta (epigeic), Lumbricus terrestris (anecic) and Allolobophora\ud chlorotica (endogeic) earthworms were cultivated in columns containing 900 g soil with 1130, 345, 113\ud and 131 mg kg1 of As, Cu, Pb and Zn, respectively, for up to 112 days, in parallel with earthworm-free\ud columns. Leachate was produced by pouring water on the soil surface to saturate the soil and generate\ud downflow. Ryegrass was grown on the top of columns to assess metal uptake into biota. Different\ud ecological groups affected metals in the same way by increasing concentrations and free ion activities in\ud leachate, but anecic L. terrestris had the greatest effect by increasing leachate concentrations of As by\ud 267%, Cu by 393%, Pb by 190%, and Zn by 429% compared to earthworm-free columns. Ryegrass\ud grown in earthworm-bearing soil accumulated more metal and the soil microbial community exhibited\ud greater stress. Results are consistent with earthworm enhanced degradation of organic matter leading\ud to release of organically bound elements. The degradation of organic matter also releases organic acids\ud which decrease the soil pH. The earthworms do not appear to carry out a unique process, but increase\ud the rate of a process that is already occurring. The impact of earthworms on metal mobility and\ud availability should therefore be considered when inoculating earthworms into contaminated soils as\ud new pathways to receptors may be created or the flow of metals and metalloids to receptors may be\ud elevated.

Summary The soil seed bank functions to escape unfavourable environmental conditions and seed predation. Anecic earthworms are increasingly recognised as important dispersers and predators of plant seeds. In contrast, the role of the usually more abundant endogeic earthworms which live and feed in the soil on plant seeds is largely unknown. We tested whether (A) endogeic earthworms ( Aporrectodea rosea , Allolobophora chlorotica , Octolasion tyrtaeum , Aporrectodea caliginosa ) ingest and digest grassland plant seeds ( Phleum pratense , Bellis perennis , Trifolium repens , Poa trivialis , Plantago lanceolata , Medicago varia ), (B) the passage of seeds through the gut of endogeic earthworms modifies seed germination, and (C) excreta (mucus and casts) of endogeic earthworms modify seed germination. As a reference effects of the well-studied anecic species Lumbricus terrestris were determined. Endogeic earthworms ingested and digested all of the studied plant seeds; however, both ingestion and digestion were earthworm and plant species specific. Moreover, passage through the gut of endogeic earthworms and their excreta modified plant seed germination (gut passage: B. perennis ; excreta: Ph. pratense and Pl. lanceolata ). The results indicate that endogeic earthworms may strongly impact the composition of the soil seed bank and, consequently, plant community assembly via direct and indirect effects on plant seeds. Since post-dispersal seed predation is a key factor for the structure of plant communities, with the effect on seed survival potentially exceeding that of pre-dispersal predation, seed predation and changes in germination of seeds by endogeic earthworm species is likely to drive plant community composition.

An investigation of earthworms across a sand dune system in NW England examined species distribution and abundance with respect to soil physico-chemical conditions and management. Replicated 0.1 m2 quadrats were examined every 50 m along 700 m transects across areas of vegetation succession and samples were then taken every 10 m where earthworms were first encountered. A translocation of Aporrectodea longa assessed the ability of this species to live in soils with a high sand content and laboratory work examined growth, maturation and survival of this species in soils with increasing proportions of sand (0–100%). Nine earthworm species were found on the dunes, but none in yellow dunes where organic matter (OM) content was

Earthworm (Lumbricidae) populations were assessed in three replicated, field experiments with different management intensities conducted on grassland research farms on medium-textured mineral soils, located at Solohead (Co. Tipperary), Grange (Co. Meath) and Johnstown Castle (Co. Wexford), Ireland. The experiment at Solohead involved four levels of mineral fertilizer application (80, 175, 225 and 350 kg N ha−1), and the plots were rotationally grazed by dairy cows. Three levels of mineral fertilizer application were compared at Johnstown Castle (0, 225 and 390 kg N ha−1) and two at Grange (100 and 225 kg N ha−1), and the plots at these sites were rotationally grazed by suckler cows and followers or dry cattle. Between 10 and 15 earthworm species were recorded per site, with Allolobophora chlorotica and Aporrectodea caliginosa being dominant. Other abundant species included Aporrectodea rosea, Aporrectodea longa, Lumbricus terrestris, Lumbricus friendi, Lumbricus festivus, Lumbricus rubellus and Satchellius mammalis. Mean earthworm population densities per treatment ranged from 203 to 324 individuals m−2 and biomass from 60 to 176 g m−2. ANCOVA revealed significant overall positive relationships between management intensity (as indicated by N application rate) and earthworm abundance (F = 4.47, p

Specific composition of earthworm community has indicative value for evaluating the impact of agricultural practice on soil. The occurrence of species only like Aporrectodea caliginosa, Aporrectodea rosea, Lumbricus rubellus tolerant to disturbance is the result of intensive tillage and agricultural practice or the influence of strong limiting ecological factor. A community including more sensitive species Lumbricus terrestris and Aporrectodea longa, or the most sensitive species Allolobophora chlorotica and Lumbricus castaneus, indicates more favourable conditions of habitat.

International audience; Earthworm burrow systems are generally described based on postulated behaviours associated with the three ecological types. In this study, we used X-ray tomography to obtain 3D information on the burrowing behaviour of six very common anecic (Aporrectodea nocturna and Lumbricus terrestris) and endogeic (Aporrectodea rosea, Allolobophora chlorotica, Aporrectodea caliginosa, Aporrectodea icterica) earthworm species, introduced into repacked soil cores for 6 weeks. A simple water infiltration test, the Beerkan method, was also used to assess some functional properties of these burrow systems. Endogeic worms make larger burrow systems, which are more highly branched, less continuous and of smaller diameter, than those of anecic worms. Among the anecic species, L. terrestris burrow systems are shorter (9.2 vs 21.2 m) with a higher number (14.5 vs 23.5) of less branched burrows (12.2 vs 20.2 branches m(-1)), which are also wider (7.78 vs 5.16 mm) than those of A. nocturna. In comparison, the burrow systems made by endogeic species appeared similar to each other. However, A. rosea burrows were short and narrow, whereas A. icterica had a longer burrow system (15.7 m), more intense bioturbation intensity (refilled macropores or soil lateral compaction around them) and thus a greater number of burrows. Regarding water infiltration, anecic burrow systems were far more efficient due to open burrows linking the top and bottom of the cores. For endogeic species, we observed a linear relationship between burrow length and the water infiltration rate (R (2) = 0.49, p < 0.01). Overall, the three main characteristics significantly influencing water infiltration were burrow length, burrow number and bioturbation volume. This last characteristic highlighted the effect of burrow refilling by casts

Summary Developments in ecotoxicology and soil restoration have led to an increase in earthworm research over the past 20 years. Rearing of earthworms under laboratory conditions has therefore become desirable, but perceived difficulties have beset this practice. A review of recent research on the development of earthworm culture techniques for temperate, soil dwelling (anecic and endogeic) species is presented. Critical abiotic and biotic factors including soil moisture, organic matter, temperature, pH, earthworm density and species composition are considered, and optimal values provided for four species ( Allolobophora chlorotica, Aporrectodea caliginosa, Aporrectodea longa and Lumbricus terrestris ). Published figures relating to growth to maturity, cocoon production and cocoon viability, for these frequently studied species are collated and advice offered for culture of other soil dwelling species.

Earthworms dominate the animal biomass in moist floodplain soils. They are known to survive long periods in aerated water, but little is known about earthworm population dynamics in floodplain systems with changing inundation frequencies. This study determined earthworm population dynamics in a floodplain system, in relation to frequency and duration of flooding events. From October 2000 to May 2003 earthworms were hand sorted in the 'Afferdensche en Deestsche Waarden', a floodplain on the south bank of the river Rhine, near Druten, The Netherlands. Earthworm numbers and biomasses per age class (adult, subadult, juvenile) were recorded. Numbers and biomasses tend to decrease during flooding. Lumbricus terrestris was found in high numbers (>10/m2) only at the end of a flooding period. Allolobophora chlorotica was hardly affected by flooding; their biomass remained stable during the year. Aporrectodea caliginosa showed fluctuating numbers and biomasses during the sampling period that did not correlate with flooding frequency. Numbers and biomasses of Lumbricus rubellus were strongly reduced at the end of each flooding event, but their population densities fully recovered until next flooding event. Earthworm populations in floodplains fluctuate in time, depending on the season and on the time, duration and frequency of flooding. Different earthworm species react differently towards these flooding dynamics. © 2004 Elsevier GmbH. All rights reserved.

Summary Coelomocytes (extruded via dorsal pores of earthworms subjected to 5V electric shock) of adult Allolobophora chlorotica, Lumbricus terrestris, Dendrobaena veneta and Eisenia fetida consisted of species-specific proportions of hyaline amoebocytes, granular amoebocytes, and eleocytes. In all species a significant increase of granular amoebocytes with a parallel decrease of hyaline amoebocytes was recorded by the end of Spring and beginning of Summer, both in animals from the natural environment and in those kept in the laboratory at the room temperature. It indicates that coelomocyte proliferation/maturation is dependent mainly on the annual endogenous rhythm, which is relatively stable even at atypical ambient temperatures. During four successive seasons viability and functions (plastic adherence or neutral red pinocytosis) were recorded in coelomocytes of A. chlorotica assayed at various in vitro temperatures. In coelomocytes retrieved from animals freshly collected from their natural environment, coelomocyte activity was better at low temperatures (0 °C and 10 °C) during the Winter, while during other seasons (especially in the Summer) it was best at 22 °C. Such seasonal effects were abolished in coelomocytes from animals tested after one-month acclimation to the room temperature, as in all seasons cell functions were best at 22 °C. These observations indicate that earthworm coelomocytes can thermally adapt. This phenomenon was also recorded in L. terrestris, D. veneta and E. fetida . In conclusion, earthworm coelomocyte proliferation/maturation seems to be endogenously controlled while coelomocyte activity seems to be more plastic and subjected to thermal adaptation.

The inverse relationship between the particle size of food resources and earthworm growth rate is widely recognised. The aim was to determine the effects of the particle size of food on interactions between combinations of adult and hatchling A. chlorotica and L. terrestris under laboratory conditions. Treatments were set up in 1 litre mesocosms, maintained in darkness at 15 ′ 1 °C. Separated cattle solids (SCS) were used as food either unmilled or milled to < 1 mm. The latter led to a 185 and 54 % increase in the mass of A. chlorotica and L. terrestris respectively, compared with unmilled SCS. Intra-specific competition in A. chlorotica was detected in adult and hatchling pairings. A positive inter-specific interaction was recorded with hatchling A. chlorotica and mature L. terrestris fed unmilled SCS with growth of the former significantly greater (P

Hatchling Lumbricus terrestris, Aporrectodea longa, Allolobophora chlorotica and L. rubellus were cultured in 1-l pots at 15°C in darkness. Each species was grown in isolation and also in the presence of selected adult species representing a range of earthworm ecological groupings. The experiments were continued for 24–36 weeks (until animals matured). The effects of adult presence were determined by measuring survival, growth rate, maturation and cocoon production of original hatchlings. In all treatments greater than 90% of animals survived. Both inter- and intra-specific interactions negatively influenced growth, maturation and fecundity of experimental earthworms. The intensity of interactions was directly related to the degree of niche overlap between pairings. Growth of both anecic and endogeic species was significantly (P

Earthworms are recognized to increase soil porosity, reorganize soil structure, and stimulate soil microflora and nutrient mineralization. The properties of earthworm casts should depend both on earthworm species or ecological group and on soil properties. Interactions between earthworm species and soil types have been suggested, but only poorly demonstrated. In order to better understand those interactions, two hypotheses led our study: (1) Soil type has a greater influence on cast properties than earthworm; (2) Earthworms from different species influence cast properties differently; (3) The intensity and direction of the impact of each earthworm species on cast properties vary with soil properties. Fifteen physical and chemical variables (N–NH 4 + , N–NO 3 − , total organic C and N, C/N ratio, CaCO 3 , pH, P, K + , Mg 2+ , Mn 2+ , Na + , CEC, moisture, wettability) were measured in casts of three earthworm species ( Lumbricus terrestris , Allolobophora chlorotica and Aporrectodea rosea ) produced in three temperate soils. Univariate and multivariate analyses showed that earthworm species and soil types significantly impacted cast properties. pH, N t , K and Mg contents were interactively altered by both factors. Multivariate analysis showed that a difference of soil type had a major impact on casts properties (62%) compared to the impact of a difference of earthworm species (10%). Cast properties were most impacted by L . terrestris , then by A . chlorotica and last by A . rosea . The response ratio (ratio of the properties of the casts to the properties of the bulk soil) was used to quantify the effect of earthworm species compared to the control soil. It showed a higher response of variables in casts in nutrient-rich soils, especially in casts of L . terrestris . The interactions between earthworm species and soil types on cast properties were discussed with regards to earthworm ecology, properties of the soil, and earthworm modifications of cast microflora.

To study the effects of earthworm casting on organic matter dynamics, control soil and casts were added as a surface layer (Horizon I) to perspex cylinders containing a ‘base’ soil depleted in organic C (Horizon II). Three treatments with different Horizon I were used; a control containing uningested soil and oak litter (Quercus petraea (Mattuschka) Lieblein), cast derived from the same substrates, and a third (Ew+cast) where cast as well as endogeic (Aporrectodea caliginosa Savigny, Allolobophora chlorotica Savigny) and anecic (Lumbricus terrestris L.) earthworms were included. These were monitored over a 2 1 2 year period. Moisture fluctuations were reduced and higher amounts of organic matter and C were present in Horizon I with the cast treatment after 2 years. In addition, the proportion of clay C (and to a lesser extent light fraction) in Horizon I decreased significantly in the control and cast treatments while there was a corresponding increase in the proportions of silt C. Overall, earthworm activity had a limited effect on C distribution in the particle size fractions studied, though the dynamics of organic matter/particle associations may require more than 2 1 2 years before clear patterns emerge.

Nightcrawler (Lumbricus terrestris L. [Oligochaeta: Lumbricidae]) is an anecic earthworm typically lacking from Persian walnut (Juglans regia L.) orchards in northern California, yet this earthworm may be established by inoculative release. Our purpose of was to determine the effect of L. terrestris inoculation on green manure incorporation, in this case woolypod vetch (Vicia villosa Roth ssp. varia [Host] Cabriere), and on the decomposer community. An experiment using controlled-environment microcosms showed that disappearance of surface-applied vetch residue was increased by the addition of L. terrestris compared to a resident earthworm complex consisting of the endogeic Lumbricidae Aporrectodea sp. prob. trapezoides (Duges) and Allolobophora chlorotica (Savigny) as well as the epigeic Eisenia foetida (Savigny). An experiment conducted in an organic Persian walnut orchard during the spring of 1996 also showed that disappearance of surface-applied vetch residue was increased in plots that had been inoculated with L. terrestris. However, an additional study conducted in the same orchard during the summer of 1998 showed similar rates of vetch biomass disappearance in plots with or without inoculated L. terrestris. Colonization by L. terrestris did not lead to significant differences in densities or live biomass for resident earthworms in this orchard. However, night observations showed significantly greater densities of isopod Crustacea (Isopoda: Oniscidae) in plots without L. terrestris, suggesting possible interference between L. terrestris and the isopods.

The aim of this study was to measure the pressures exerted by earthworms during burrowing. For this purpose we developed two methods with which to quantify the axial and radial pressure. The data were recorded with an electronic balance that was connected to a PC. Artificial earthworm burrows were used to standardize the measurements. Plexiglas tubes with diameters ranging from 2 to 6.3 mm which corresponded to the diameter of the earthworms were used. A pin was placed inside the tubes, on which the earthworms exerted a pressure by peristaltic locomotion. Only the maximum values of the pressure measurement were taken into account for evaluation, and the arithmetic mean was calculated. The measurements were conducted with Aporrectodea longa, Lumbricus terrestris, Aporrectodea caliginosa, Octolasion cyaneum, Allolobophora chlorotica, Aporrectodea rosea, Lumbricus rubellus and Dendrobaena octaedra. The species examined were classified into ecological groups. The mean axial pressures exerted by each group were in the order: epigeic (14–25 kPa), endogeic (27–39 kPa) and anecic (46–65 kPa). For the mean radial pressure the order was: epigeic (39–63 kPa), anecic (72–93 kPa) and endogeic (59–195 kPa). It was apparent from the results that radial pressure is the most important pressure with respect to the burrowing activity of earthworms.

Studies of earthworm species and their activity, expressed as channels, on cultivated loamy (Humic Cryaquept) and clayey soils (Typic Endoaqualf and Typic Cryaquept) were conducted in southern and central Norway before conversion from conventional to organic cropping systems. At all the three study areas: Landvik (Grimstad), Voll (As) and Kvithamar (Stjordal), the earthworm species Aporrectodea caliginosa, Aporrectodea rosea, Allolobophora chlorotica and Lumbricus rubellus were found. At Landvik, where the deep-burrowing species Aporrectodea longa, A. caliginosa and Lumbricus terrestris dominated, medium (4–6 mm) and coarse (>6 mm) earthworm channels were most numerous below the plough layer. Almost no coarse pores were found at Voll and Kvithamar. The volume of earthworm channels below the plough layer was 0.6–0.8% of total soil volume at Landvik and Voll and 0.3–0.4% at Kvithamar. Earthworm channels >6 mm below the plough layer were useful to identify present and previous activity of deep-burrowing earthworms such as L. terrestris. Below the plough layer, roots were almost entirely restricted to earthworm channels and interaggregate spaces.

Predation on three species of earthworms by the terrestrial planarian, Artioposthia triangulata, was compared in columns of artificial soil (10 cm dia, 1 m depth) at four different temperatures (5, 10, 12 and 16°C). The planarian significantly reduced the number of earthworms and affected the vertical distribution of all species by reducing prey numbers in the top 20 cm. Greater earthworm mortality occurred at higher temperatures but there was no difference in the numbers of each species that were killed.

The effects of metal contaminants on the population density and species composition of earthworms were studied at 22 sites around a primary smelting works situated at Avonmouth, southwest England. All worms were absent from six sites within 1 km of the factory and numbers were also reduced significantly at an additional four sites 2 km from the plant. Total earthworm density was found to be inversely related to concentrations of metals in soils. some species of earthworms were found to be more ‘sensitive’ to metals and were absent from sites where more ‘tolerant’ species persisted. For example, Lumbricus rubellus, Lumbricus castaneus and Lumbricus terrestris were all present at sites close to the smelter where Aporrectodea rosea, Aporrectodea caliginosa and Allolobophora chlorotica were absent. The first three ‘tolerant’ species have more active calcium secretion glands in their gut than the three ‘sensitive’ worms. Calcium is known to be involved in the sequestration and elimination of various metals through the chlorogogenous tissue. Toxicity tests were carried out with a ‘sensitive’ and a ‘non-sensitive’ earthworm species ( Lumbricus rubellus and Aporrectodea rosea respectively) to examine if earthworm species distribution could be related to their susceptibility to the metals. In addition, the OECD recommended species Eisenia fetida was tested, to compare the sensitivity of this worm to the two commonly occurring soil species. Zinc was the metal used for these tests, since earlier comparisons of metal toxicity between the laboratory and field have shown that of the four metals released from the smelting works (Cd, Cu, Pb, Zn), it is zinc that is limiting the distribution of worms. Test results indicated that Aporrectodea rosea is more sensitive to the effects of zinc than Lumbricus rubellus . Thus, it appears that differences in the distribution of species of earthworms around the smelter can be related to variation in sensitivity to zinc. Both species were affected by zinc at lower concentrations than Eisenia fetida .

Three species of earthworm, Lumbricus terrestris L., Aporrectodea longa (Ude) and Allolobophora chlorotica (Savigny), were offered a choice of mixtures of soil and small wheat straw fragments which had been inoculated individually with six saprotrophic fungi. All earthworm species showed preferences between the six fungal species offered. Early straw decomposers, capable of utilizing water-soluble sugars and cellulose, were preferred in most cases to the lignin-decomposing fungi characteristic of the later stages of decomposition. The removal of fungal-inoculated straw pieces from the soil surface by L. terrestris followed the same pattern. The palatability of two wheat pathogens to L. terrestris was found to be similar to that of the preferred saprotroph. The implications of these findings for fungal abundance and dispersal in wheat fields are discussed.

The earthworm population in a winter cereal field in Ireland was studied over a 3-year-period and its effects on soil and N turnover were assessed. The mean annual population density was 346–471 individuals m-2 and the mean biomass was 56.9–61.2 g m-2. Twelve species were recorded, the most abundant being Allolobophora chlorotica followed by Aporrectodea caliginosa, and 242 mg at 5°C to 713 mg at 10°C in the case of juvenile Lumbricus terrestris. Gut contents (dry mass of soil) comprised 6.7–15.5% of the A. caliginosa live mass, and 9.7–14.7% of the Lumbricus terrestris mass. Annual soil egestion by the field population was estimated as 18–22 kg m-2. Tissue production ranged from 81.7 to 218.5 g m-2, while N turnover resulting from mortality was calculated as 1.5–3.9 g m-2 depending on the year and the method of calculation. Earthworms were estimated to contribute an additional 3.4–4.1 g mineral N to the soil through excretion, mucus production, and soil ingestion. Independent estimates of N output via mucus and excretion derived from 15N laboratory studies with Lumbricus terrestris were 2.9–3.6 g m-2 year-1.

The effects of wastewater sludge amendments on nitrogen forms and urease activity in soil and earthworm (Lumbricus terrestris) casts were investigated. Two different wastewater sludges (municipal and canned food industry) were added to clay loam agricultural soil samples at doses of 20, 40 and 80 g dry raw sludge kg(-1). Following sludge amendment, an incubation study was performed and mineral nitrogen (NH(4)(+)-N+NO(3)(-)-N), total-N and urease activity were determined in soil and earthworm casts. Results indicated that the addition of wastewater sludge significantly increased urease activity, mineral-N and total-N contents of soil and casts. Furthermore, urease activity, mineral-N and total-N contents of casts were found to be significantly higher than that of the surrounding soil. Application of canned food industry sludge at a dose of 80 g kg(-1)(200 t ha(-1)) was the most effective application on nitrogen and urease contents. On the other hand, 40 g kg(-1) (100 t ha(-1)) treatment was found to be the most suitable dose for municipal wastewater sludge.

Concentrations of heavy metals (Pb, Cd, Zn and Cu) were determined in five species of earthworms, representing different ecophysiological groups, sampled from soils associated with an abandoned Pb and Zn mine site. Surface soil-dwelling species (Aporrectodea caliginosa and Allolobophora chlorotica) accumulated significantly greater concentrations of Cd than deep-burrowing (Lumbricus terrestris and Aporrectodea longa) or litter-dwelling (L. rubellus) species. Aporrectodea caliginosa generally accumulated greater concentrations of Pb than other species. The greatest concentrations of Zn were generally found in the deep-burrowing species, whilst lowest concentrations were found in Allolobophora chlorotica. No consistent inter-species pattern was evident for Cu. The composition (metal and organic carbon) of the ingesta of earthworms was different to that of the soil. Concentrations of Cd in the ingesta of Aporrectodea caliginosa were significantly greater than in the ingesta of Aporrectodea longa and L. terrestris; concentrations ofPb in the ingesta of Aporrectodea caliginosa and Aporrectodea longa were greater than for L. terrestris. The metal concentrations in the faeces were similar for all species. A significant correlation exists between ingesta and tissue Cd concentrations, but was not evident for Pb or Zn. This suggests that ingested total Cd concentration is a primary factor in determining tissue Cd concentrations of earthworms. For the other metals other interactive factors appear to markedly influence accumulated tissue metal concentrations.

During the past 25 yr, research by the authors at Rothamsted Experimental Station investigated many aspects of the utilization of earthworms in land improvement and environmental management. Results of some of these investigations are summarized in this paper with the aim of illustrating the general principles of how earthworm populations can be manipulated and managed for environmental improvement. The use of earthworms in land improvement and reclamation: we investigated the effects of inoculating earthworms of the species Lumbricus terrestris L., Aporrectodea longa (Ude), Aporrectodea caliginosa (Sav.) and Allolobophora chlorotica (Sav.) into intact soil profiles in the laboratory, plots on direct-drilled, arable land in the field and newly-capped waste disposal sites that had few or no earthworms. In all these studies the earthworms increased significantly in number and rate of growth and yield of plants growing on the inoculated sites. Earthworms for inoculation were obtained by field collection after watering soil with dilute formaldehyde solution. The use of earthworms in organic waste management: the life cycles and productivity of Eisenia fetida (Sav.), Eudrilus eugeniae (Kinberg), Perionyx excavatus (Michaelsen) and Dendrobaena veneta (Rosa), and their potential in processing animal and plant wastes, from sewage, agricultural, domestic, urban and industrial sources are summarized. The preprocessing of wastes, their population ecology, optimum stocking rates, the mechanization of processing and utilization of the product are discussed. Results of experiments on the effects of temperatures of 10, 15, 20 and 25°C and a range of soil moisture contents of 70, 75, 80, 85 and 90% on the growth, cocoon production and cocoon hatching of the four species are summarized. The use of earthworms in assessment of the environmental effects of chemicals: earthworms can be used as key indicators to predict the effects of chemicals on other soil invertebrates. Methods of testing chemicals against earthworms in field and laboratory are reviewed. Two standardized laboratory test methods, one exposing earthworms to chemicals on filter paper and one to chemicals in artificial soils are described, and the median lethal concentration (LC 50 ) for chloracetamide, pentachlorophenol, chlordane, carbaryl, potassium bromide, copper sulfate and trichloracetic acid calculated, based on assays done in 34 laboratories. The relevancy of the two tests in environmental toxicity testing is reviewed.

Summary Field experiments were done at two widely separated sites in England (Tadcaster, North Yorkshire and North Wyke, Devon). In each case an old permanent sward was re-seeded after ploughing or by direct drilling, or was left intact and fertilised. The carbamate pesticide aldicarb was applied to half of each experimental plot. The effects of sward improvement and pesticide on various soil invertebrate groups were assessed. Soil-dwelling macrofauna arthropods were scarce at both sites. Stem-boring dipterous larvae, however, were numerous at both sites. At Tadcaster, Oscinella vastator was the only species present in the original sward, but 0. frit was by far the most prevalent species in re-seeded areas. These can only have arisen from direct oviposition of eggs by adults on or near the emerging seedlings and not by migration of larvae from buried turf or the desiccated old pasture. Mites and Collembola were abundant. Their numbers were reduced initially by cultivation and pesticide usage, but the effects of these practices were transient and their numbers recovered to previous levels usually within 10 wk. Earthworms were present at both sites. Deep burrowing species, e.g. Lumbricus terrestris and Allolobophora longa were less affected by the treatments imposed than species such as A. chlorotica and L. rubelius which usually live close to the soil surface. Effects on their numbers were in any case, short-lived. No improvement method consistently provided the greatest herbage dry matter yield. Pesticide application, however, greatly increased seedling stand and/or herbage yield at both sites and showed the benefits of using crop protectants, especially on direct-drilled re-seeds.

To validate cholinesterase activity as a biomarker of pesticide exposure, we characterized the tissue distribution (whole body, nervous tissue and crop/gizzard), activity at two seasons of cholinesterase in six different species of earthworms collected in an unpolluted field: Lumbricus terrestris, Lumbricus castaneus, Aporrectodea nocturna, Aporrectodea caliginosa, Allolobophora chlorotica and Aporrectodea rosea. The major part of total cholinesterase activity was found in the nervous tissue while activity in crop/gizzard was weak. The level of the total cholinesterase activity was stable for each species considered throughout the year (spring and autumn). Lumbricus species exhibited three-fold higher specific activity than the others (0.086+/-0.015 U mg(-1) and 0.235+/-0.036 U mg(-1) for Allolobophora or Aporrectodea, and Lumbricus species respectively). This stability of the base level makes cholinesterase activity a useful biomarker for monitoring effects of pesticide under natural conditions. Cholinesterase activity was characterized using different substrates and inhibitors. It seems likely that the cholinesterases are acetylcholinesterases in most species investigated as they preferentially hydrolyzed acetylthiocholine and were inhibited by eserine, but not by tetraisopropyl pyrophoramide (iso-OMPA). Characterization of cholinesterase from Allolobophora chlorotica is uncertain and it cannot be classified as a true AChE.

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Oil spills are one of the most common types of soil pollution. Bioremediation has become an attractive alternative to physicochemical methods of remediation, where feasible. Earthworms have been shown to stimulate the degradation of petroleum hydrocarbons in soil, and it was hypothesized that the role of earthworms in remediation lies in the enhancement of an oil degrading microbial community. The aim of this study was to characterize microbial activity and community dynamics in oil-contaminated soil incubated with or without earthworms. Three earthworm species (Eisenia fetida, Allolobophora chlorotica and Lumbricus terrestris) were incubated in crude oil polluted soil (ca. 10,000 mg/kg total petroleum hydrocarbons (TPH)) and a reference soil for 28 d. Control treatments with manual mixing and/or cattle dung amendment were also included. In the oil-contaminated soil, respiration and concentration of microbial biomass was significantly enhanced by earthworm amendment, and TPH concentrations decreased significantly. These effects were less evident in treatments with A. chlorotica, possibly due to a difference in behavior, since individuals of this endogeic species were found in a state of inactivity (aestivation). Microbial community dynamics were described by phospholipid fatty acid (PLFA) analyses. After 28 d, similar shifts in the soil PLFA composition were observed in the oil-contaminated soil irrespective of worm species. Fungal:bacterial ratios were increased in the presence of worms, but also by addition of dung as a food source, indicating a non-specific effect of metabolizable substrates. In contrast, the fatty acids 17:1ω8 (=Δ9-heptadecenoic acid) and 20:4ω6c (arachidonic acid) were specifically stimulated by the presence of earthworms in the oil-contaminated soil. The results showed that earthworms can contribute positively to bioremediation of oil-contaminated soil, but that the effect may be species-dependent.

Earthworm response to a single cultivation sequence was assessed. Cropping and management of cultivated land was similar to that of non-cultivated areas. Effects were measured in a trial on recently replaced opencast mining land. A subsequent survey investigated responses to cultivation on both replaced and undisturbed land at periods since cultivation ranging from 1.5 to 7.5 years. On undisturbed land, adverse effects of cultivation, if any, were short-lived, with numbers of all species, but particularly deep-burrowing species, having recovered or increased after 3 years. However, on replaced land the initial adverse effect appeared to be larger, and recovery from this effect was delayed until at least 5 years after tillage. Different species present (Lumbricus terrestris/ Aporrectodea longa, Lumbricus rubellus/festivus, Aporrectodea caliginosa/Octalasion tyrtaeum and Allolobophora chlorotica) also varied in the extent of the initial reduction and in their rate of recovery. Reasons for different responses to cultivation and implications for the management of replaced land are considered.

Comparison was made of the heavy metal concentrations of lead, zinc and calcium in earthworms and soil from pasture near a large lead and zinc smelting plant and a control area 9 km away. The concentration of lead in the earthworms was less than that in the soil, whereas zinc and cadmium concentrations were several times the value in the soil. The zinc and cadmium concentration factors for all worm species were significantly lower at the contaminated site than at the control site and both were significantly greater than for lead. There was 7–9 times as much zinc and cadmium in the soil from the smelting site as in that from the control area and 1·6 times as much lead. Corresponding values in the worms were 1–3·5 times as much zinc and cadmium and 1·4–4·0 times as much lead. The zinc:cadmium ratio decreased from 81 in soil to 26 for earthworms (average of aggregation of species) at the control site and from 63 to 18 at the contaminated site. The best overall relationship between metal concentration in earthworms and concentration in soil was y = k.xb where y = concentration of metal per gramme of worm, dry weight, and x = concentration of metal per gramme of soil, dry weight. Comparison of regression slopes of the log transformed data showed that uptake was greater for lead than for the other two metals. For each metal the worms are in two species groups. For the Lumbricus terrestris, Allolobophora chlorotica, A. rosea group the rates of uptake for zinc and cadmium were not significantly different. There was no evidence that the earthworm populations near the smelting plant had been reduced by the heavy metal contamination.

The water content of Lumbricus terrestris, after keeping on moist filter-paper for 3 or 4 days, is 84-8 % of its body weight. That of Allolobophora chlorotica is 80 % of its body weight. Both species can survive a loss of 60 % of the body weight, but not much more. Earthworms of the species A. chlorotica, A. terrestris f. longa, Dendrobaena subrubicunda, L. rubellus and L. terrestris are all able to survive from 31 to 50 weeks in soil totally submerged beneath aerated water. The same species, and A. caliginosa can survive for 72-137 days in aerated tap water without food. Garden specimens of A. chlorotica make U-shaped burrows in soil beneath water. They do not irrigate either the burrows or glass tubes. Egg-cocoons of A. chlorotica, taken from culture pots of soil, will hatch under water and the young worms will feed and grow though totally immersed. D. subrubicunda, L. rubellus and L. terrestris avoid immersion in water when allowed a choice between water-saturated and moist but air-filled soil. When A. caligmosa, A. chlorotica and A. terrestris f. longa are allowed a similar choice, a small proportion of the population is always found in the water-filled soil. Enforced submergence before being allowed a choice of environment does not affect the behaviour of L. rubellus, i.e. water is still avoided. Similar treatment of A. chlorotica and L. terrestris results in a more frequent choice of water. It is unnecessary to postulate that the individuals of A. chlorotica occurring in Lake Windermere constitute a ‘biological race’. The avoiding reaction of earthworms to immersion may explain why they leave the soil after heavy rain.