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.     

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.     

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.     

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.     

Characteristics of earthworms (Eisenia fetida) in PAHs contaminated soil amended with sewage sludge or vermicompost

Earthworms can be used to remove polycyclic aromatic hydrocarbons (PAHs) from soil, but this might affect their survival and they might accumulate the contaminants. Sterilized and unsterilized soil was contaminated with phenanthrene (Phen), anthracene (Anth) and benzo(a)pyrene (BaP), added with or without Eisenia fetida, sewage sludge or vermicompost. Survival, growth, cocoon formation and concentrations of PAHs in the earthworms were monitored for 70 days. Addition of sewage sludge to sterilized or unsterilized soil maintained the number of earthworms and their survival was 94%. The addition of sludge significantly increased the weight of earthworms 1.3 times compared to those kept in the unamended soil or in soil amended with vermicompost. The weight of earthworms was significantly lower in sterilized than in unsterilized soil. Cocoons were only detected when sewage sludge was added to unsterilized soil. A maximum concentration of 62.3 μg Phen kg⁻¹ was found in the earthworms kept in sterilized soil amended with vermicompost after 7 days and 22.3 μg Phen kg⁻¹ when kept in the unamended unsterilized soil after 14 days. Concentrations of Phen in the earthworms decreased thereafter and ≤2 μg kg⁻¹ after 28 days. A maximum Anth concentration of 82.5 μg kg⁻¹ was found in the earthworms kept in sterilized soil amended with vermicompost and 45.8 μg Anth kg⁻¹ when kept in the unamended unsterilized soil after 14 days. A maximum concentration of 316 μg BaP kg⁻¹ was found in the earthworms kept in sterilized soil amended with vermicompost after 56 days and 311 μg BaP kg⁻¹ when kept in the unsterilized soil amended with vermicompost after 28 days. The amount of BaP in the earthworm was generally largest after 28 days, but after 70 days still 60 μg kg⁻¹ was found in E. fetida when kept in the sterilized soil amended with sewage sludge. It was found that E. fetida survived in PAHs contaminated soil and accumulated only small amounts of the contaminants, but sewage sludge was required as food for its survival and cocoon production.     

Earthworm communities and soil properties in shaded coffee plantations with and without application of glyphosate

In central Veracruz, Mexico, many coffee plantations are managed using agrochemicals for weed control, with glyphosate-based herbicides (GBH) the most commonly used. To date, however, no studies in this region have characterized the soil biological and physicochemical properties in coffee plantations under such glyphosate application. In this study, earthworms were used as bioindicator organisms by measuring differences in the earthworm community in plots within shaded coffee plantations, with and without repeated applications of glyphosate. Differences in earthworm-induced soil processes, such as water infiltration rates, potential net carbon mineralization rates and soil physicochemical properties were also evaluated. Eight plots were selected in shaded coffee plantations; four had received regular applications of GBH over the preceding 22 years, while the other four had received no herbicides over the preceding 7 years. The earthworm species found in plots with no GBH treatment were Pontoscolex corethrurus (99%) and Amynthas corticis (1%), while A. corticis was absent in plots that had been treated with GBH. Significant differences (P < 0.01) in earthworm density (168 ± 16 and 353 ± 37 ind m⁻²) and biomass (22.7 ± 1.1 and 45.4 ± 6.9 g m⁻²) were observed in soils with and without GBH, respectively. No significant difference (P = 0.08) was observed in the water infiltration rate (2 × 10⁻⁴ ± 4 × 10⁻⁵ and 4 × 10⁻⁴ ± 1 × 10⁻⁴ cm s⁻¹ with and without GBH, respectively). Soil carbon flow was greater in plots with GBH (76 ± 7 μg dry soil⁻¹ d⁻¹) than in those without GBH (62 ± 1 μg dry soil⁻¹ d⁻¹, P < 0.005). Significant differences (P < 0.05) were found in pH and in the clay, silt and Ca content of the soil. Our findings indicated reduced species number, density and biomass of earthworms, and increased net carbon mineralization rate in plots with GBH. The plots managed with glyphosate presented a negative effect on the earthworm parameters measured, and we conclude that the earthworms therefore acted as indicators of perturbation. It is also possible that this effect could be due to factors unrelated to the glyphosate that were not considered in this study, such as chemical fertilization or legume litter spatial variability, among others.     

Effects of azoxystrobin,chlorothalonil, and ethoprophos on the reproduction of three terrestrial invertebrates using a natural Mediterranean soil

The potential terrestrial toxicity of three pesticides, azoxystrobin, chlorothalonil, and ethoprophos was evaluated using reproduction ecotoxicological tests with different non-target species: the collembolan Folsomia candida, the earthworm Eisenia andrei, and the enchytraeid Enchytraeus crypticus. All reproduction tests were performed with natural soil from a Mediterranean agricultural area (with no pesticide residues) in order to improve the relevance of laboratory data to field conditions. Controls were performed with natural and standard artificial soil (OECD 10% OM). The fungicide azoxystrobin showed the highest toxicity to earthworms (EC₅₀ = 42.0 mg a.i. kg⁻¹ dw soil). Collembolans were the most sensitive taxa in terms of sublethal effects of chlorothalonil with an EC₅₀ of 31.1 mg a.i. kg⁻¹ dw soil followed by the earthworms with an EC₅₀ of 40.9 mg a.i. kg⁻¹ dw soil. The insecticide ethoprophos was the most toxic to collembolans affecting their reproduction with an EC₅₀ of 0.027 mg a.i. kg⁻¹ dw soil. Enchytraeids were generally the least sensitive of the three species tested for long-term effects. Earthworms were not always the most sensitive species, emphasizing the need to increase the number of mandatory assays with key non-target organisms in the environmental risk assessment of pesticides.     

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.     

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.     

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.     

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.     

Compacted soil affects soil macrofauna populations in a semi-arid environment in central Queensland

Population densities of soil macrofauna were assessed in a field experiment with annually compacted treatments (applied to whole plots) and management treatments to repair initially compacted soil. Earthworms accounted for about half the macrofauna recovered during the experiment. Compaction of wet surface soil (water content>plastic limit) by agricultural machinery generally reduced numbers of macrofauna and earthworms. Annual compaction with a 10 Mg axle load on wet soil reduced mean macrofauna numbers from 70 to 15 m⁻² and mean earthworm numbers from 41 to 2 m⁻². Annual compaction with 6 Mg on soil drier than the plastic limit to a depth of 0.08 m had no adverse effect on the soil macrofauna. A 3-year pasture ley had more macrofauna (211 m⁻²) than a control treatment under cropping (29 m⁻²) but numbers declined when cropping was resumed.     

Darwin's earthworms revisited

Down House was Charles Darwin's home from 1842 until his death in 1882 and where he wrote “The Formation of Vegetable Mould through the Action of Worms”. The work described here is based upon passages from this book and from further observations on earthworms in this area. General observations were made in addition to systematic sampling in areas selected either from signs of earthworm activity, habitat type or in direct relation to Darwin's documented work.Greatest species richness (n = 9) was found in Middle Field. Greatest earthworm density was present in Darwin's Kitchen Garden (715 m⁻²) with the largest associated biomass (261 g m⁻²). Aporrectodea longa was the most abundant species. Lumbricus terrestris, described by Darwin in terms of its behaviour, but not directly named, was located in relatively low numbers, but its diagnostic middens and associated burrows were easily detected. Earthworms associated with Darwin's cinder and chalk application experiments were also examined. In total, 19 of Britain's 28 earthworm species were located within the nominated World Heritage Site.     

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.     

Assessing exotic plant species invasions and associated soil characteristics: A case study in eastern Rocky Mountain National Park,Colorado, USA,using the pixel nested plot design

Rocky Mountain National Park (RMNP), Colorado, USA, contains a diversity of plant species. However, many exotic plant species have become established, potentially impacting the structure and function of native plant communities. Our goal was to quantify patterns of exotic plant species in relation to native plant species, soil characteristics, and other abiotic factors that may indicate or predict their establishment and success. Our research approach for field data collection was based on a field plot design called the pixel nested plot. The pixel nested plot provides a link to multi-phase and multi-scale spatial modeling-mapping techniques that can be used to estimate total species richness and patterns of plant diversity at finer landscape scales. Within the eastern region of RMNP, in an area of approximately 35,000 ha, we established a total of 60 pixel nested plots in 9 vegetation types. We used canonical correspondence analysis (CCA) and multiple linear regressions to quantify relationships between soil characteristics and native and exotic plant species richness and cover. We also used linear correlation, spatial autocorrelation and cross correlation statistics to test for the spatial patterns of variables of interest. CCA showed that exotic species were significantly (P < 0.05) associated with photosynthetically active radiation (r = 0.55), soil nitrogen (r = 0.58) and bare ground (r = −0.66). Pearson's correlation statistic showed significant linear relationships between exotic species, organic carbon, soil nitrogen, and bare ground. While spatial autocorrelations indicated that our 60 pixel nested plots were spatially independent, the cross correlation statistics indicated that exotic plant species were spatially associated with bare ground, in general, exotic plant species were most abundant in areas of high native species richness. This indicates that resource managers should focus on the protection of relatively rare native rich sites with little canopy cover, and fertile soils. Using the pixel nested plot approach for data collection can facilitate the ecological monitoring of these vulnerable areas at the landscape scale in a time- and cost-effective manner.     

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.     

The role of biological activity (roots,earthworms) in medium-term C dynamics in vertisol under a Digitaria decumbens (Gramineae) pasture

The natural abundance of ¹³C was used to estimate the turnover of the soil organic matter in a vertisol re-grassed with Digitaria decumbens (C4 plant) following intensive market gardening (C3 plants). In addition, the experimental design allowed us to determine the respective roles of roots and earthworms (Polypheretima elongata) in soil C stock restoration in D. decumbens pasture.The C stock increased from 31 to 37 Mg C ha⁻¹ in 5 years and the δ¹³C increased from −18.1‰ in market gardening soil to −15.5‰ in the 5-year-old pasture soil in the upper 20 cm. Below the 20 cm soil layer, the C stock and the δ¹³C did not change significantly in 5 years. The net gain of 6 Mg C ha⁻¹ was the balance of a loss of 5 Mg C ha⁻¹ derived from market gardening and a gain of 11 Mg C ha⁻¹ derived from D. decumbens. Effects of earthworms on the C dynamics were not discernible.     

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.     

Removal of benzo (a) pyrene from soil using an endogeic earthworm Pontoscolex corethrurus ()

The endogeic earthworm Pontoscolex corethrurus (Müller, 1857) was the most abundant species (75%) in soil contaminated with hydrocarbons, mostly benzo(a)pyrene (BaP), in the state of Tabasco (Mexico). The earthworm P. corethrurus was tested for its capacity to remove 100 mg BaP kg⁻¹ from an Anthrosol soil (sterilized or not) and amended with legume Mucuna pruriens (L.) DC. var. utilis (Wall. ex Wight) Baker ex Burck (3%) or the grass Brachiaria humidicola (L.) DC (3%) (recently renamed as Urochloa humidicola (Rendle) Morrone & Zuloaga) in an aerobic incubation experiment. P. corethrurus removed 26.6 mg BaP kg⁻¹ from the sterilized soil and application of B. humidicola as feed increased this to 35.7 mg BaP kg⁻¹ and M. pruriens to 34.2 mg BaP kg⁻¹ after 112 days. The autochthonous microorganisms removed 9.1 mg BaP kg⁻¹ from the unsterilized soil and application of B. humidicola increased this to 18.0 mg BaP kg⁻¹ and M. pruriens to 11.2 mg BaP kg⁻¹. Adding P. corethrurus to the unsterilized soil accelerated the removal of BaP and 36.1 mg kg⁻¹ was dissipated from soil. It was found that the autochthonous microorganisms removed BaP from soil, but addition of P. corethrurus increased the dissipation 4-fold. The endogeic earthworm P. corethrurus can thus be used to remediate hydrocarbon-contaminated soils in tropical regions.