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.     

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).     

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.     

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.     

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.     

Earthworm colonisation of abandoned arable soil polluted by copper

The distribution, density and biomass of earthworms were investigated at the copper polluted site, Hygum (Denmark). In 1994, shortly after farming of the area was abandoned, only four earthworm species were present and their distribution was restricted to areas where copper concentration did not exceed 200 mg kg^?1 dry soil. Sixteen years later (in 2010), without any agricultural activity, ten species of earthworms were found, in particular, epigeic species were present where soil copper concentrations reached >1000 mg kg^?1 dry 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.     

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.     

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.     

Earthworms in a 15 years agricultural trial

Alternative cropping systems have been proposed to enhance sustainability of agriculture, but their mid and long-term effects on soil biodiversity should be studied more carefully. Earthworms, having important agro-ecological functions, are regarded as indicators of soil biological health. Species composition, abundance, and biomasses of earthworms were measured in autumn 2005–2007 (period 1) and 2011–2013 (period 2) in a trial initiated in 1997 near Paris, France. A conventional, an organic and a direct seeded living mulch-based cropping systems were compared. Earthworms were sampled in a wheat crop by combining the application of a chemical expellant and hand-sorting.In period 1, earthworm abundance did not usually differ in the three cropping systems, but sometimes it was higher in the conventional system. Mean total abundance was 122, 121 and 149 individuals m⁻² in period 1 and 408, 386 and 216 in period 2 in the organic, living mulch and conventional systems respectively. While earthworm abundance and biomass increased slightly in the conventional system between the two periods, they at least tripled in the other two systems. This was mainly due to the species Aporrectodea caliginosa and Aporrectodea longa in the living mulch cropping system, and to A. caliginosa, Lumbricus castaneus, Lumbricus terrestris and A. longa in the organic system.After at least 14 years, organic and living mulch cropping systems contained between 1.5 and 2.3 times more earthworms than the conventional system. Considering the inter-annual variations in earthworm communities due to climatic conditions and cultural practices, earthworm communities should be assessed over several years before conclusions can be drawn. Moreover, since changes in cultural practices may take a long time to affect earthworm communities, mid and long-term trials are needed to assess the effects of cropping systems on soil biodiversity.     

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.     

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.     

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.     

Impact of ecologically different earthworm species on soil water characteristics

A laboratory experiment was performed to assess the impact of ecologically different earthworm species on soil water characteristics, such as soil tension, water content, and water infiltration rate. Three earthworm species (Lumbricus rubellus, Aporrectodea caliginosa, Lumbricus terrestris) were exposed in soil columns (diameter 30 cm, height 50 cm) for 100 days with a total fresh earthworm biomass of 22.7 ± 0.4 g per column, each in duplicate. Each column was equipped with tensiometers at 10 and 40 cm and FD-probes at 10 cm depth, to continuously measure the temporal development of soil tension and soil moisture. Additionally, 30 g of sieved and rewetted horse manure was placed on the soil surface as a food source. Precipitation events (10 mm) were simulated at day 28 and day 64. At the end of the experiment the water infiltration rate and the runoff at 55 cm depth were determined.The results showed considerable evidence, that ecologically different earthworms modify soil water characteristics in different ways. The anecic L. terrestris and the endogeic A. caliginosa showed the tendency to enhance the drying of the topsoil and subsoil. Their intensive and deep burrowing activity might enhance the exchange of water vapor due to a better aeration in soil. In contrast, the epigeic L. rubellus tended to enhance the storage of soil moisture in the topsoil, which might be linked to lower rates of litter loss from soil surface and thus a thicker litter layer remaining. A. caliginosa led to considerable higher water infiltration rates and faster water discharges in the subsoil, relative to the other species, probably due to a high soil dwelling activity.     

Effects of bioenergy crop cultivation on earthworm communities—A comparative study of perennial (Miscanthus) and annual crops with consideration of graded land-use intensity

Energy crops are of growing importance in agriculture worldwide. This field study aimed to investigate earthworm communities of different intensively cultivated soils during a 2-year period, with special emphasis on annual and perennial energy crops like rapeseed, maize, and Miscanthus. These were compared with cereals, grassland, and fallow sites. Distribution patterns of earthworm abundance, species, and ecological categories were analysed by constrained ordination procedures (redundancy analysis; CANOCO) using a set of environmental variables as predictors, such as CN value of harvest residues, SOC and N_t content, soil pH, soil texture, and land-use intensity. The latter was determined by principal component analysis using average soil coverage and intensity of tillage, weed control, and fertilisation as input variables. It was clearly found that land-use intensity was the dominant regressor for earthworm abundance and total number of species. The diversity of earthworm communities was especially enhanced and showed a more balanced species composition in extensively managed soils under grassland, fallow, and Miscanthus. For the total number of species, Miscanthus (5.1 ± 0.9) took a medium position and neither differed significantly from intensively managed rapeseed (4.0 ± 0.9), cereals (3.7 ± 1.1), and maize sites (3.0 ± 1.4), nor from grassland (6.8 ± 1.5) and fallow (6.4 ± 1.0) sites. Total earthworm abundance ranged between 355 (±132) and 62 (±49) individuals m⁻² in fallow and maize sites, respectively.Interestingly, Miscanthus had quite positive effects on earthworm communities although the CN value of harvest residues was very high. It is recommended that Miscanthus may facilitate a diverse earthworm community even in intensive agricultural landscapes.     

Earthworm populations in relation to soil organic matter dynamics and management in California tomato cropping systems

Earthworms are key regulators of soil structure and soil organic matter (SOM) dynamics in many agroecosystems. They are greatly impacted by agricultural management, yet little is known about how these factors interact to control SOM dynamics. This study sought to explore linkages between agricultural management, earthworms and aggregate associated SOM dynamics through a survey of tomato (Solanum lycopersicum L.) cropping systems in northern California. Earthworms and soil samples were collected between February and April of 2005 from 16 fields under one of three types of residue management: (1) tomato mulch – no postharvest tillage and tomato residues left on the soil surface, (2) cover crop – tomato residues tilled in and leguminous cover crop planted, and (3) bare fallow – tomato residues tilled in and soil surface left exposed throughout the winter. Earthworms were collected via hand-sorting and identified to species, while soils were wet sieved to yield four aggregate size classes: large macroaggregates (>2000 μm), small macroaggregates (250–2000 μm), microaggregates (53–250 μm) and the silt and clay fraction (<53 μm). The combined large and small macroaggregate fraction was then fractionated into coarse particulate organic matter (cPOM; 250 μm), microaggregates within macroaggregates (mM; 53–250 μm) and macroaggregate occluded silt and clay (Msc; <53 μm). The earthworms identified in this survey were composed entirely of exotic species and were dominated by Aporrectodea caliginosa. Earthworm abundance was related to residue management, with the tomato mulch systems averaging 4.5 times greater fresh earthworm biomass than bare fallow (P = 0.024). Aggregate stability and total soil C and N also appeared to be influenced by residue management, such that the tomato mulch system displayed significantly greater mean weight diameters than the bare fallow system (P = 0.049), as well as more than 50% greater total soil C and N (P = 0.049 and P = 0.036; respectively). Earthworm biomass was also found to be positively correlated with total soil C (P = 0.009, R² = 0.39) and N (P = 0.010, R² = 0.039) as well as the proportion of macroaggregate C in the cPOM fraction (P = 0.028, R² = 0.30). Our findings suggest that residue handling and the associated management practices (e.g., tillage, organic vs. conventional agriculture) are important for both earthworm populations and SOM storage. Although earthworms are known to influence SOM in many ways, other factors appear to play a more prominent role in governing aggregate associated SOM dynamics.     

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.     

Role of soil water content in the carbon and nitrogen dynamics of Lumbricus terrestris L. burrow soil

We evaluated the role of soil water content in controlling C and N dynamics within the drilosphere created by the anecic earthworm Lumbricus terrestris (L.). Mesocosms (volume = 3.1 l) were each amended with corn litter and three earthworms. Control treatments received no earthworms and no other earthworm species were present in the soil. WET and DRY treatments received a total of 9.25 cm and 3.25 cm of water, respectively. Water was added on weeks 1, 3, 7, and 10 at a rate of 2.0 cm per mesocosm for WET treatments and 0.5 cm per mesocosm for DRY treatments. Mesocosms were sampled destructively after incubation at 18–20 °C for 0, 3, 7, and 13 weeks. The water content of WET burrow soil ranged from 0.12 g g⁻¹ to 0.18 g g⁻¹ and was significantly higher than in the DRY treatment throughout the incubation period. The live weight of earthworms was significantly higher in the WET treatment only on week 13, whereas litter consumption was significantly lower in the DRY treatment for week 13. Carbon mineralization, measured as CO₂ evolved after a 24-h incubation, was consistently higher in WET than in DRY burrow soil. Effects of differences in soil water content were also apparent for biomass C and metabolic quotient. Soil water content did no affect the total C concentration of burrow soil. DRY burrow soil had consistently lower levels of nitrate than WET soil throughout the experiment. Lower levels of ammonium and inorganic N were observed for WET burrow soil on weeks 3 and 7. Water content did not have a significant effect on burrow soil total N. We concluded that the water content of the drilosphere affects both C and N dynamics and can affect the speciation of inorganic N; yet, the effects of soil water content do not appear to result from differences in the feeding activities of anecic earthworms.     

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.     

Impact of soil stockpiling and mining rehabilitation on earthworm communities

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