Earthworm density and biomass in relation to plant diversity and soil properties in a Palouse prairie remnant

Earthworms are important soil animals in grassland ecosystems and are considered to be important to soil quality. The overall impact of earthworms on soil properties and plant diversity, however, depends on earthworm species, functional group and the type of ecosystem. The primary purpose of this study was to document the relationship among earthworms, key soil properties and native and exotic plant diversity in the little studied, Palouse prairie grassland (Idaho, USA). A secondary objective was to determine the effectiveness of three methods commonly used to sample earthworms. A hillslope characterized by Palouse prairie vegetation, well-expressed, hummocky (mounded) topography and known to support both exotic and native earthworm species was selected for study. The hillslope was divided into three zones [annual-dominated (AD), mixed (MX) and perennial-dominated (PD)] based on characteristics of the inter-mound plant communities described in previous research. Total earthworm biomass in the MX zone (53.5 g m⁻²) was significantly greater than in the PD zone (14.7 g m⁻²) (P = 0.0384), but did not differ from the AD zone. Earthworm density ranged from 52 to 81.1 individuals m⁻² but was not significantly different across zones. Total C and N at 0 to 10 and 30 to 50 cm depths were significantly greater in the AD and PD zones as compared to the same depths in the MX zone. Soil textural class was silt loam within all zones and the soil silt fraction was positively correlated with total exotic earthworm density (R = 0.783, P = 0.0125) and biomass (R = 0.816, P = 0.0072). Native earthworms were only found in the zone with the greatest total and native plant diversity (PD). Total soil C and N were not correlated to earthworm density, but soil total C and N were significantly negatively correlated with exotic plant density, which indicates that invasive plants may be decreasing soil total C (R = −0.800) and N (R = −0.800). Calculated earthworm densities using data from the electroshocker were generally lower than those based on the hand-sorting method. Electroshocking, however, created lower disturbance and was the only method that resulted in the collection of the deep-burrowing, native species Driloleirus americanus.     

Can commercial mulches be reservoirs of invasive earthworms? Promotion of ligninolytic enzyme activity and survival of Amynthas agrestis (Goto and Hatai, 1899)

Amynthas agrestis is an exotic, invasive earthworm in North America that has been associated with horticulture settings as well as damage to forest soil. An experiment was conducted to find out whether A. agrestis, an earthworm commonly found in mulches in Vermont, stimulates ligninolytic enzymes in the presence of commercial wood mulches. Mesocosms filled with a sandy loam soil were topped with either spruce, cedar or pine mulch. Half of the mesocosms received juvenile A. agrestis, the other half did not. After 7 weeks soils were analyzed for phenoloxidase and peroxidase activity. Most A. agrestis survived and developed into adults during the incubation period. Significantly greater phenoloxidase activity was detected in soils with A. agrestis than without earthworms. Mean (standard deviation) phenoloxidase activities in the presence of A. agrestis were 0.15 (±0.10), 1.14 (±0.46), 2.71 (±0.98) μmol g⁻¹ h⁻¹ for pine, spruce and cedar respectively, and 0.012 (±0.023), 0.25 (±0.25), 0.78 (±0.45) μmol g⁻¹ h⁻¹ in the absence of A. agrestis. There was significantly greater peroxidase activity for the pine and spruce treatment when earthworms were present. Mean peroxidase activities were 0.47 (±0.21), 0.94 (±0.29), 1.20 (±0.77) μmol h⁻¹ g⁻¹ soil for pine, spruce and cedar, respectively for soils with A. agrestis and 0.15 (±0.10), 0.37 (±0.10), 0.63 (±0.30) μmol h⁻¹ g⁻¹ soil in the absence of earthworms. The increased ligninolytic activity in combination with successful maturation of juveniles into adult A. agrestis suggests that mulch can be habitat for these invasive earthworms. This finding is supported by a survey of master gardeners in Vermont and New Hampshire 20% of whom reported to have seen these earthworms mainly in their gardens and mulched beds.     

What are the limits of the drilosphere? An incubation experiment using Metaphire posthuma

The near infrared reflectance spectroscopy (NIRS) method was used in the present study to compare earthworm-made soil aggregates to aggregates found in the surrounding bulk soil. After initially assessing the daily cast production of Metaphire posthuma, boxes with soil incubated with M. posthuma and control soils were subjected to wetting in order to reorganize the soil structure. After two months of incubation, soil aggregates produced by earthworms (casts and burrows), soil aggregates that were appeared to be unaffected by earthworms (bulk soil without visible trace of earthworm bioturbation from the earthworm treatment) and soil aggregates that were entirely unaffected by earthworms (control – no earthworm – treatment) were sampled and their chemical signatures analyzed by NIRS. The production of below-ground and surface casts reached 14.9 g soil g worm^?1 d^?1 and 1.4 g soil g worm^?1 d^?1, respectively. Soil aggregates from the control soils had a significantly different NIRS signature from those sampled from boxes with earthworms. However, within the earthworm incubation boxes the NIRS signature was similar between cast and burrow aggregates and soil aggregates from the surrounding bulk soil. We conclude that the high cast production by M. posthuma and the regular reorganization of the soil structure by water flow in and through the soil lead to a relatively homogenous soil structure. Given these results, we question the relevance of considering the bulk soil that has no visible activity of earthworm activity as a control to determine the effect of earthworms on soil functioning.     

Endogeic and anecic earthworm abundance in six Midwestern cropping systems

Endogeic and juvenile anecic earthworm abundance was measured in soil samples and anecic populations were studied by counting midden numbers at the sites of two long-term cropping systems trials in South-central Wisconsin. The three grain and three forage systems at each site were designed to reflect a range of Midwestern USA production strategies. The primary objectives of this work were to determine if the abundance of endogeic or anecic earthworms varied among cropping systems or crop phases within a cropping system and were there specific management practices that impacted endogeic or anecic earthworm numbers. The earthworms present in the surface soil were: Aporrectodea tuberculata (Eisen), A. caliginosa (Savigny), A. trapezoides (Dugés); and juvenile Lumbricus terrestris (L.). True endogeic abundance was greatest in rotationally grazed pasture [188 m^?2 at Arlington (ARL) and 299 m^?2 at Elkhorn (ELK)], and smallest in conventional continuous corn (27 m^?2 at ARL and 32 m^?2 at ELK). The only type of anecic earthworm found was L. terrestris L. There was an average of 1.2 middens per adult anecic earthworm and the population of anecics was greatest in the no-till cash grain system (28 middens m^?2 at ARL, 18 m^?2 at ELK) and smallest in the conventional continuous corn system (3 middens m^?2 at ARL, 1 m^?2 at ELK). Earthworm numbers in individual crop phases within a cropping system were too variable from year-to-year to recommend using a single phase to characterize a whole cropping system. Indices for five management factors (tillage, manure inputs, solid stand, pesticide use, and crop diversity) were examined, and manure use and tillage were the most important impacting earthworm numbers across the range of cropping systems. Manure use was the most important management factor affecting endogeic earthworm numbers; but no-tillage was the most important for the juvenile and adult anecic groups and had a significantly positive influence on endogeic earthworm counts as well. The pesticides used, which were among the most commonly applied pesticides in the Midwestern USA, and increasing crop diversity did not have a significant effect on either the endogeic or anecic earthworm groups in this study. Consequently, designing cropping systems that reduce tillage and include manure with less regard to omitting pesticides or increasing crop diversity should enhance earthworm populations and probably improve sustainability.     

Identifying earthworm's organic matter signatures by near infrared spectroscopy in different land-use systems in Tabasco,Mexico

In the state of Tabasco, South-eastern, Mexico, land-use changes such as the conversion of natural into agricultural systems, modify soil quality and the abundance of soil macrofauna, including earthworms. The aim of this study was to characterize by near-infrared spectroscopy (NIRS) the earthworms’ fingerprint in soil, in six sites including natural and agricultural ecosystems with low and high earthworm biomass and low and high earthworm diversity, in order to identify specific wavelengths that discriminate the presence/abundance of earthworm species and functional groups. The spectral region of 1860–1870 nm was significantly correlated with total earthworm density, particularly at one of the sites (Cedar polyculture; r = 0.8, p < 0.05). Earthworm biomass had a specific NIRS wavelength according to the earthworm species and feeding category: 1820 and1860–1870 nm wavelengths were significantly correlated with Polypheretima elongata (r² = 0.7, p < 0.05; mesohumic species) biomass and 2090 nm for biomass of all Lavellodrilus species (polyhumics). Two species had a much wider spectral range: L. bonampakensis and Dichogaster saliens (an epigeic worm; 1690–2300 nm, r² = 0.7, p < 0.05). Biomasses of Periscolex brachysistis and Diplotrema murchiei were not significantly correlated with any near infrared wavelength spectra analyzed. Combining a maximum of 4 species per wavelength, mesohumic earthworms had a wider wavelength spectrum than polyhumics. Therefore, earthworm species diversity, biomass and abundance are associated with soil quality (as measured by NIR spectra) and this relationship varies with species and ecological category. Sites with lower and higher earthworm diversity have lower and higher soil organic matter quality, respectively, as observed by the wider or narrower spectral range with which earthworm biomasses are correlated.     

Deep-burrowing earthworm additions changed the distribution of soil organic carbon in a chisel-tilled soil

We investigated the influence of earthworms on the three-dimensional distribution of soil organic carbon (SOC) in a chisel-tilled soil. By burrowing, foraging, and casting at the surface and throughout the soil, anecic earthworms such as Lumbricus terrestris L. may play a major role in regulating the spatial distribution of organic matter resources both at the surface and within the soil. In the fall of 1994, we manipulated ambient earthworm communities, which were without deep burrowing species, by adding 100 earthworm individuals m⁻² in spring and fall for 3 years. Overall, the biomass of L. terrestris was increased with earthworm additions and total earthworm biomass declined compared with ambient control treatments. To investigate the spatial variability in soil organic carbon due to this shift in earthworm community structure, we sampled soil on a 28×24 cm grid from the surface to 40 cm in four layers, 10 cm deep. Samples were analyzed for total carbon. We found that additions of anecic earthworms significantly increased average soil organic carbon content from 16.1 to 17.9 g C kg⁻¹ for the 0–10 cm soil, and from 12.4 to 14.7 g kg⁻¹ at 10–20-cm depth, and also changed the spatial distribution of soil organic carbon from uniform to patchy, compared with the ambient treatment.     

Effects of cropping and tree density on earthworm community composition and densities in central Cameroon

Earthworms can have positive effects upon crop growth in the tropics. If soils are to be managed sustainably, then more attention should be paid to the effects of cultivation and cropping practices upon earthworms. When forest vegetation is cleared, slashed, burned and land is tilled and cultivated, earthworm abundance, diversity and activity are reduced. Conversely, retaining trees in agroecosystems may maintain earthworm populations during the cropping phase.Here, we assessed the impact on earthworm species diversity and densities of crop cultivation in the understorey of timber plantations thinned to two tree densities and compared these with uncropped, undisturbed timber plantation controls. The plots were reassessed after two and a half years of fallow to see whether populations had recovered. The experiment was in central Cameroon.Seventeen earthworm species were recorded from Eudrilidae subfamilies Eudrilinae and Pareudrilinae, Ocnerodrilidae and Acanthodrilidae, most of which were endemics. This included two new species from two new genera from the sub-family Pareudrilinae, one new species from one new genus of Ocnerodrilidae, two new species of Dichogaster and one new species of Legonodrilus. Ten species were epigeic, six were endogeic and one was anecic.Generally, earthworm densities were lower in cropped plots than in the undisturbed plantation control. The most abundant species was a Legonodrilus sp. nov. with average densities of 49 individuals m⁻² in the crop phase and 80 ind. m⁻² in the fallow phase. By the fallow phase, densities in the low tree density (120 ind. m⁻²) were higher than in the high density (40 ind. m⁻²). The densities of the epigeic Acanthodrilidae were significantly reduced to 7 ind. m⁻² in the cropped plots compared with 42 ind. m⁻² in the control plots. The effects of cropping were thus species-specific and more work is required to identify which of these endemics are the ecosystem engineers in the system.     

Allyl isothiocyanate: an alternative chemical expellant for sampling earthworms

Allyl isothiocyanate (AITC), a natural breakdown product of glucosinolates in many Cruciferae and a component imparting the sharp taste to mustard, was tested for its effectiveness as a chemical expellant for sampling earthworms. Testing was performed in an arable field with earthworm populations dominated by Lumbricus terrestris Linnaeus, 1758 and Aporrectodea tuberculata (Eisen, 1874). The optimal concentration was found by comparing concentrations ranging from 5 to 250 mg l⁻¹ in water. Total biomass and numbers of earthworms collected increased hyperbolically with increasing AITC concentration, with the highest biomass and numbers collected using 100 mg l⁻¹ AITC. Biomass and numbers of earthworms collected using 250 mg l⁻¹ AITC, but not 150 or 200 mg l⁻¹ AITC, were significantly less than with 100 mg l⁻¹ AITC. Less earthworm biomass was collected by hand sorting than with chemical expulsion using 100 mg l⁻¹ AITC, but the number of earthworms collected by the two methods were not different. A comparison of hand sorting and 100 mg l⁻¹ AITC expulsion using analysis of similarities (ANOSIM) showed that the two methods produced samples differing in the distribution of both numbers and biomass of species-by-size classes. Hand sorting collected more of the smallest size class of L. terrestris and the largest size classes of A. tuberculata than AITC expulsion, whereas AITC expulsion recovered more of the largest size classes of L. terrestris than hand sorting. When 100 mg l⁻¹ AITC expulsion was compared with chemical expulsion using 200 mg l⁻¹ formalin, no differences were found in the total number, total biomass or in the species-by-size class distribution of the earthworms collected. This suggests that the AITC method may be a favorable alternative to formalin expulsion for sampling earthworms. Further studies under other environmental conditions and with other species of earthworms are warranted to establish its general utility.     

Earthworms as soil quality indicators in Brazilian no-tillage systems

It is well known that earthworm populations tend to increase under no-tillage (NT) practices, but abundances tend to be highly variable. In the present study, data from the literature together with those on earthworm populations sampled in six watersheds in SW Paraná State, Brazil, were used to build a classification of the biological soil quality of NT systems based on earthworm density and species richness. Earthworms were collected in 34 farms with NT aging from 3 to 27 yr, in February 2010, using an adaptation of the TSBF (Tropical Soil Biology and Fertility) Program method (hand sorting of five 20 cm × 20 cm holes to 20 cm depth). Six forest sites were also sampled in order to compare abundances and species richness with the NT systems. Species richness in the 34 NT sites and in the 6 forests ranged from 1 to 6 species. Most earthworms encountered were exotics belonging to the genus Dichogaster (D. saliens, D. gracilis, D. bolaui and D. affinis) and native Ocnerodrilidae (mainly Belladrilus sp.), all of small individual size. In a few sites, individuals of the Glossoscolecidae (P. corethrurus, Glossoscolex sp., Fimoscolex sp.) and Megascolecidae (Amynthas gracilis) families were also encountered, in low densities. Urobenus brasiliensis (Glossoscolecidae) were found only in the forest fragments. In the NT farms, earthworm abundance ranged from 5 to 605 ind m⁻² and in the forest sites, from 10 to 285 ind m⁻². The ranking of the NT soil biological quality, based on earthworm abundance and species richness was: poor, with <25 individuals per m⁻² and 1 sp.; moderate, with ≥25–100 individuals per m⁻² and 2–3 sp.; good, with >100–200 individuals per m⁻² and 4–5 sp.; excellent, with >200 individuals per m⁻² and >6 sp. About 60% of the 34 farms fell into the poor to moderate categories based on this classification, so further improvements to the NT farm's management system are needed to enhance earthworm populations. Nevertheless, further validation of this ranking system is necessary to allow for its wider-spread use.     

Earthworms in New Zealand sheep- and dairy-grazed pastures with focus on anecic Aporrectodea longa

Earthworms play an important role as primary decomposers in the incorporation and initial mixing of plant litter. This study explored the response of earthworms to increasing fertiliser inputs, pasture production and livestock numbers (and their influence on food availability and soil physical condition) on six different managements in sheep-grazed and fifteen different managements in dairy-grazed pastures in a variety of New Zealand soils.Native earthworms were only found in some low-fertility pastures. Accidentally introduced peregrine earthworms, when present, dominate pasture soils. Of these, endogeic earthworms dominated the earthworm community and were positively associated with soil types with higher bulk densities. Peregrine anecic earthworms were absent from most hill-country sheep-grazed pastures, however in more fertile and productive dairy-grazed pastures they reached a biomass of up to 2370 kg ha^?1. Only anecic earthworms showed a positive response to the increasing pressures associated with higher potential dry matter inputs and liveweight loadings of grazing livestock on soil, while epigeic earthworms declined. The positive response of anecic earthworms probably reflects the combined effect of the increase in food resources, including dung and plant litter, available on the soil surface, and their lower susceptibility to livestock treading pressure. Anecic species may be a suitable substitute for incorporation of surface litter in those soils where livestock treading limits epigeic earthworm populations.This study confirmed previous observations of limited distribution of the introduced Aporrectodea longa in pastoral hill-country soils in the North Island, and their near absolute absence from the South Island of New Zealand. This would suggest that large areas of New Zealand pastoral farmed soils could benefit from the introduction of anecic species from other parts of New Zealand which already contain A. longa.     

Effect of tillage on earthworms over short- and medium-term in conventional and organic farming

Earthworms play an important role in many soil functions and are affected by soil tillage in agricultural soils. However, effects of tillage on earthworms are often studied without considering species and their interactions with soil properties. Furthermore, many field studies are based on one-time samplings that do not allow for characterisation of temporal variation. The current study monitored the short (up to 53 days) and medium term (up to 4 years) effects of soil tillage on earthworms in conventional and organic farming. Earthworm abundances decreased one and three weeks after mouldboard ploughing in both conventional and organic farming, suggesting direct and indirect mechanisms. However, the medium-term study revealed that earthworm populations in mouldboard ploughing systems recovered by spring. The endogeic species Aporrectodea caliginosa strongly dominated the earthworm community (76%), whereas anecic species remained <1% of all earthworms in all tillage and farming systems over the entire study. In conventional farming, mean total earthworm abundance was not significantly different in reduced tillage (153 m⁻²) than mouldboard ploughing (MP; 130 m⁻²). However, reduced tillage in conventional farming significantly increased the epigeic species Lumbricus rubellus from 0.1 m⁻² in mouldboard ploughing to 9 m⁻² averaged over 4 years. Contrastingly, in organic farming mean total earthworm abundance was 45% lower in reduced tillage (297 m⁻²) than MP (430 m⁻²), across all sampling dates over the medium-term study (significant at 3 of 6 sampling dates). Reduced tillage in organic farming decreased A. caliginosa from 304 m⁻² in mouldboard ploughing to 169 m⁻² averaged over 4 years (significant at all sampling dates). Multivariate analysis revealed clear separation between farming and tillage systems. Earthworm species abundances, soil moisture, and soil organic matter were positively correlated, whereas earthworm abundances and penetration resistance where negatively correlated. Variability demonstrated between sampling dates highlights the importance of multiple samplings in time to ascertain management effects on earthworms. Findings indicate that a reduction in tillage intensity in conventional farming affects earthworms differently than in organic farming. Differing earthworm species or ecological group response to interactions between soil tillage, crop, and organic matter management in conventional and organic farming has implications for management to maximise soil ecosystem functions.     

Earthworms in Chromolaena odorata (L.) King and Robinson (Asteraceae) fallows along a chronosequence: Changes in community structure and identification of persistent and indicator species

The species Chromolaena odorata (Asteraceae) is a notorious invasive shrub spreading throughout West and Central Africa and as such, there is a need to determine its environmental impact, particularly on soil biodiversity and functioning. Indeed, soil organisms such as earthworms are known to strongly influence soil properties and biogeochemical cycles. This study, conducted in Central Côte d’Ivoire, aims to investigate the temporal dynamics of earthworm communities in C. odorata fallows of different ages and to identify associated indicators and persistent species. Three distinct classes of fallows identified by local farmers, were considered: young (1–3 years, C1), medium-aged (4–8 years, C2) and old (>9 years, C3). Each of the classes included four plot replicates where earthworms were sampled using the Tropical Soil Biology and Fertility (TSBF) 25 cm × 25 cm × 30 cm soil monolith method. The study of earthworm communities was focused on density, biomass, diversity and complementarity. Indicator values (IndVals) were used to identify indicator species of the classes of fallows. The shrub exerted a mixed influence on earthworms depending on the functional group, with litter feeders and polyhumics declining over time as a result of a reduction of the litter availability on the soil surface. The species richness was significantly greater in C1 than in the other classes although the Shannon–Weaver's index did not vary significantly. However, a cluster analysis performed on densities highlighted marked differences between C2 and the two other classes in terms of community composition. Indicator species were found for C1 and C2. The geophagous Millsonia omodeoi has emerged as a persistent species as its density and biomass steadily increased so that it became the dominant species in old fallows. The roles of litters and soil parameters in influencing earthworm communities are discussed.     

The significant contribution of mycorrhizal fungi and earthworms to maize protection and phytoremediation in Cd-polluted soils

Mycorrhizal fungi and earthworms can individually or interactively influence plant growth and heavy metal uptake. The influence of earthworms and arbuscular mycorrhizal (AM) fungi either alone or in combination on maize (Zea mays L.) growth and cadmium (Cd) uptake was investigated in a calcareous soil artificially spiked with Cd. Soils were contaminated with Cd (10 and 20 mg Cd kg⁻¹), inoculated or un-inoculated with the epigeic earthworm Lumbricus rubellus and two AM fungal species (Rhizophagus irregularis and Funneliformis mosseae) for two months of growth under greenhouse conditions. Generally, earthworms alone increased both shoot P uptake and biomass but decreased shoot Cd concentration and root Cd uptake. AM fungi individually often increased total maize P uptake, declined shoot Cd concentration, and consequently produced higher total biomass. However, R. irregularis enhanced shoot Cd uptake at low Cd level and root Cd uptake at high Cd level. In plants inoculated with F. mosseae species, earthworms increased shoot biomass and Cd uptake, decreased root biomass and Cd uptake at all Cd levels, and increased shoot Cd concentration at low Cd level. In plants colonized by R. irregularis species, however, earthworm addition decreased maize biomass only at high Cd level and root Cd concentration and total maize Cd uptake at both Cd levels. Earthworm activity decreased Cd transfer from the soil to maize roots at low Cd level, but this was counterbalanced in the presence of F. mosseae. Mycorrhizal symbiosis significantly reduced the transfer of Cd from roots to shoots, independence of earthworm effect. Overall, it is concluded that L. rubellus and AM fungi, in particular F. mosseae isolate, improved maize tolerance to Cd toxicity both individually and interactively by increasing plant growth and P nutrition, and restricting Cd transfer to the aboveground biomass. Consequently, the single and interactive effects of the two soil organisms might potentially be important not only in protecting maize plants against Cd toxicity, but also in Cd phytostabilization in soils polluted by this highly toxic metal.     

Soil properties and metal accumulation by earthworms in the Siena urban area (Italy)

This paper reports the results of a study focused on the metal (Cd, Co, Cr, Cu, Ni, Pb, Sb, U and Zn) distribution in soils and uptake and accumulation by earthworms Nicodrilus caliginosus (Savigny) from urban, peri-urban, green-urban and non-urban zones of Siena municipality (central Italy). The main goal was to define the influence of soil properties and metal soil contents on the uptake of these contaminants by earthworms. Data indicated that Cd, Cu, Pb, Sb and Zn soil contents increased in the following order: non-urban < green-urban < peri-urban < urban soils, suggesting that vehicular traffic affects the distribution of these metals. Pb and Sb were the main soil contaminants and their highest enrichments were found in urban sites where stop-and-go traffic occurs. Concentrations of these traffic-related metals in earthworms showed a distribution pattern similar to that in soil, suggesting that soil contamination influenced the uptake of Cd, Cu, Pb, Sb and Zn by N. caliginosus. There were significant positive correlations between Cd, Pb and Sb earthworm concentrations and their soil contents. The lack of correlation for Cu and Zn could be due to the physiological regulation of these elements by earthworms. Statistical analysis pointed out that the uptake and accumulation of Cd, Cu, Pb, Sb and Zn by earthworms were affected by some soil physicochemical properties such as the organic carbon and carbonate contents that are able to rule the bioavailability of metals in soils.     

Factors influencing earthworm communities in post-industrial areas of Krakow Soda Works

Application of earthworm in soil re-cultivation and re-creation in post-industrial ecosystems make a big challenge for temporal applied zoology. The sediments of the Krakow Soda Works “Solvay” have undergone land reclamation in different ways: older sediments traps were left without any re-cultivation practices; meanwhile the newest ones were reclaimed using standard method (new soil cover planted with combination of grass and leguminous plants). The effect of different treatments on community and population structure of earthworm was estimated during consecutive years 1999–2000. Six localities differing in time of establishment, reclamation processes, vegetation type and soil properties were chosen. Nine species were recorded, among which Aporrectodea caliginosa occurred in all localities, being also the most abundant. Two other species, Lumbricus rubellus and Dendrobaena octaedra, which are epigeic species, become most important in forest assemblages and were characteristic for communities of older succession processes. Abundance of adult forms as well as total biomass were significantly affected by soil depth (r = 0.75, P < 0.05, r = 0.917, P < 0.001, respectively). Species richness however was connected with higher amount of macroelements and average plant height. Shannon diversity index and its evenness negatively correlated only with forestation (r = –0.67, P < 0.05, r = –0.68, P < 0.05, respectively). Niche overlap (α Pianka) for all analysed species extracted two groups differing in environmental requirements. First contained epigeic earthworms, closely related to plant succession (PCA results), the other one grouped endogeic and anecique species correlated significantly with soil depth and plant density. Community structure of earthworms do not reflects succession changes in post-industrial habitats, but is strongly affected by microhabitat factors in local scale (mainly soil depth and plant density).     

Impact on soil fauna of sheep faeces containing a range of parasite control agents

To compare the impact of parasite control agents in sheep faeces, 1 kg quantities of fresh faeces were spread uniformly over 1 m² pasture plots in June 2001 (winter; a time of high earthworm activity). Faecal treatments applied to five replicate plots were C− (none), C+ (from untreated sheep), B (from sheep with an intra-ruminal bolus releasing a benzimidazole anthelmintic—‘albendazole’), ML (from sheep with a bolus releasing a macrocyclic lactone anthelmintic—‘ivermectin’), F (from sheep receiving a daily feed supplement containing chlamydospores of the nematophagous fungus, Duddingtonia flagrans). The disappearance of faeces was assessed visually over the 50 days following faecal application, then soil samples were taken to assess: (a) populations of earthworms and other soil macrofauna, (b) nematodes and other soil microfauna, and (c) the presence of D. flagrans in soil. Faecal disappearance was greatest in F and C+ plots and least in ML and B plots at 12 and 23 days (P < 0.05). Earthworm casting after 23 and 50 days was greater (P < 0.05) in plots with faeces (C+, ML, F, but not B) than in plots without faeces (C−). Greater earthworm activity in plots with faeces was reflected in greater numbers of earthworms, cocoons and greater biomass m⁻² than in C− plots. On the basis of faecal dry weight applied, F plots had most earthworms and ML plots the least. After 50 days total nematodes in 0–5 cm soil showed a treatment effect (P < 0.001), being more abundant in F, C+ and B than in C− and ML plots; enchytraeids, rotifers, tardigrades and copepods showed no treatment effects. A few nematode taxa (Acrobeles, Alaimus, Pungentus, Tylencholaimus) showed significant treatment effects. The greatest effect among nematodes was in nematode channel ratio (NCR) (P < 0.008), with a decrease in F plots; changes in NCR may reflect the impact of earthworm activity on soil processes rather than a direct effect of the fungal treatment on nematodes. D. flagrans did not become established in the soil. During the trial conditions were favourable for earthworms and their activity was high in all treatments receiving faeces, with F and ML plots being the extremes. There was an apparent shift towards fungal-mediated decomposition in F plots. At the end of the 50-day trial, in a period when earthworms were active, there was no evidence of differential effects of any of the anthelmintic treatments on environmental indicators.     

Organic carbon sequestration in earthworm burrows

Earthworms strongly affect soil organic carbon cycling. The aim of this study was to determine whether deep burrowing anecic earthworms enhance carbon storage in soils and decrease C turnover. Earthworm burrow linings were separated into thin cylindrical sections with different distances from the burrow wall to determine gradients from the burrow wall to the surrounding soil. Organic C, total N, radiocarbon (¹⁴C) concentration, stable isotope values (δ¹³C, δ¹⁵N) and extracellular enzyme activities were measured in these samples. Anecic earthworms increased C stocks by 270 and 310 g m^?2 accumulated in the vertical burrows. C-enrichment of the burrow linings was spatially highly variable within a distance of millimetres around the burrow walls. It was shown that C accumulation in burrows can be fast with C sequestration rates of about 22 g C m^?2 yr^?1 in the burrow linings, but accumulated C in the burrows may be mineralised fast with turnover times of only 3–5 years. Carbon stocks in earthworm burrows strongly depended on the earthworm activity which maintains continuous C input into the burrows. The enhanced extracellular enzyme activity of fresh casts was not persistent, but was 47% lower in inhabited burrows and 62% lower in abandoned burrows. Enzyme activities followed the C concentrations in the burrows and were not further suppressed due to earthworms. Radiocarbon concentrations and stable isotopes in the burrow linings showed an exponential gradient with the youngest and less degraded organic matter in the innermost part of the burrow wall. Carbon accumulation by anecic earthworm is restricted to distinct burrows with less influence to the surrounding soil. Contrary to the initial hypothesis, that organic C is stabilised due to earthworms, relaxation time experiments with nuclear magnetic resonance spectroscopy (NMR) did not reveal any enhanced adsorption of C on iron oxides with C stabilising effect. Our results suggest that earthworm activity does not substantially increase subsoil C stocks but burrows serve as fast ways for fresh C transport into deep soil horizons.     

The efficiency of soil hand-sorting in assessing the abundance and biomass of earthworm communities. Its usefulness in population dynamics and cohort analysis studies

Pit digging and manually revising soil blocks is a frequently used method used for field studies of earthworm communities. The aim of this study was to compare the efficiency of hand-sorting (HS) to extract small earthworms, ca. 0.2 g, and the usefulness in studies of population dynamics and cohort analysis. Many earthworms are not recovered when revising manually the soil. Factors include soil characteristics, i.e. moisture, texture, etc. and also a human factor, which is more relevant if the study is conducted in the long-term. We used data collected in a field study of earthworm communities during 2 years in the savannas of Colombia. Small soil blocks (20 × 20 × 20 cm) were dug out in order to collect the smallest earthworms by washing-sieving (WS) and compare the results with the standard HS of large monoliths (100 × 100 × 50 cm). In fact, this methodology has rarely been addressed in earthworm population field studies. Our results showed that HS efficiency varied owing to the species and ranged from 31.4% up to 100% in the savanna and from 44% to 80% in the pasture, for two small species, i.e Aymara n. sp. (epigeic) and Ocnerodrilidae sp. (endogeic). In the case of the Glossodrilus n. sp. (endogeic) these values were similar, i.e. 51.7% and 58.1%, in the savanna and pasture, respectively. We also used frequency tables to calculate the average efficiency of HS 1 m² soil cores for each weight class in each species in order to obtain a population density correction factor. This allowed us to make corrections in earthworm density in the histograms for population dynamics analysis. We conclude that this method should be the modus operandi in long-term earthworm demography studies.     

Uptake and bioaccumulation of heavy elements by two earthworm species from a smelter contaminated area in northern Kosovo

As, Cd, Cu, Pb, Sb and Zn concentrations were determined in two earthworm species (Allolobophora rosea and Nicodrilus caliginosus) from a mining and industrial area in northern Kosovo and compared with their contents in the bulk soil and the main soil fractions. Earthworm specimens were collected at fifteen sites located at different distances from a Pb–Zn smelter along a gradient of decreasing contamination. Individuals of A. rosea and N. caliginosus showed similar tissue levels of As, Cd, Cu, Pb, Sb and Zn, suggesting that earthworm species belonging to the same eco-physiological group have a similar propensity to uptake and bioaccumulate heavy elements. Cd, Pb, Sb and Zn concentrations in both earthworm species were positively correlated with the respective total soil contents and generally decreased with distance from the smelter. The bioaccumulation factor (BAF) revealed that Cd and Zn were the only elements bioaccumulated by earthworms. The rank order of BAF values for both species was as follows: Cd > > Zn > > Cu > As = Pb = Sb. The absorption of Cd, Pb, Sb and Zn by earthworms mostly depended on the extractable, reducible and oxidable soil fractions, suggesting that the intestine is likely the most important uptake route. The extractable soil fraction constantly influenced the uptake of these heavy elements, whereas the reducible fraction was important mainly for Pb and Zn. The water soluble fraction had an important role especially for the most mobile heavy elements such as Cd and Zn, suggesting that dermal uptake is not negligible. As a whole, the analytical data indicate that soil fractionation patterns influence the uptake of heavy elements by earthworms, and the extractable fraction is a good predictor of heavy element bioavailability to these invertebrates in soil.     

The combined effects of earthworms and arbuscular mycorrhizal fungi on microbial biomass and enzyme activities in a calcareous soil spiked with cadmium

Earthworms and arbuscular mycorrhizal fungi (AMF) are known to independently affect soil microbial and biochemical properties, in particular soil microbial biomass (SMB) and enzymes. However, less information is available about their interactive effects, particularly in soils contaminated with heavy metals such as cadmium (Cd). The amount of soil microbial biomass C (MBC), the rate of soil respiration (SRR) and the activities of urease and alkaline phosphatase (ALP) were measured in a calcareous soil artificially spiked with Cd (10 and 20 mg Cd kg⁻¹), inoculated with earthworm (Lumbricus rubellus L.), and AMF (Glomus intraradices and Glomus mosseae species) under maize (Zea mays L.) crop for 60 days. Results showed that the quantity of MBC, SRR and enzyme activities decreased with increasing Cd levels as a result of the elevated exchangeable Cd concentration. Earthworm addition increased soil exchangeable Cd levels, while AMF and their interaction with earthworms had no influence on this fraction of Cd. Earthworm activity resulted in no change in soil MBC, while inoculation with both AMF species significantly enhanced soil MBC contents. However, the presence of earthworms lowered soil MBC when inoculated with G. mosseae fungi, showing an interaction between the two organisms. Soil enzyme activities and SRR values tended to increase considerably with the inoculation of both earthworms and AMF. Nevertheless, earthworm activity did not affect ALP activity when inoculated with G. mosseae fungi, while the presence of earthworm enhanced urease activity only with G. intraradices species. The increases in enzyme activities and SRR were better ascribed to changes in soil organic carbon (OC), MBC and dissolved organic carbon (DOC) contents. In summary, results demonstrated that the influence of earthworms alone on Cd availability is more important than that of AMF in Cd-polluted soils; and that the interaction effects between these organisms on soil microorganism are much more important than on Cd availability. Thus, the presence of both earthworms and AMF could alleviate Cd effects on soil microbial life.