Quantitative estimates of burrow construction and destruction,by anecic and endogeic earthworms in repacked soil cores

Although the role of earthworms in soil functioning is often emphasised, many important aspects of earthworm behaviour are still poorly understood. In this study we propose a simple and cost-effective method for estimating burrow system area and continuity, as well as a new and often neglected parameter, the percentage of burrow refilling by the earthworms own casts. This novel parameter is likely to have a huge influence on the transfer properties of the burrow system. The method uses standard repacked soil cores in PVC cylinders and takes advantages of clay shrinkage and the fact that earthworms were previously shown to prefer to burrow at the PVC/soil interface. In this way, after removing the PVC cylinders off dry cores, the external section of the burrow system made by earthworms along the soil walls could be easily described. We applied this method to characterise the burrow systems of four earthworms species: two anecics (Aporrectodea caliginosa nocturna and Aporrectodea caliginosa meridionalis) and two endogeics (Aporrectodea caliginosa icaliginosa and Allolobophora chlorotica). After one month the burrow's area generated by both anecic species were much larger (about 40 cm²) than the endogeic burrow's area (about 15 cm²). A. nocturna burrow system continuity was higher than that of A. meridionalis and both anecic burrow systems were more continuous than those made by the endogeic earthworms. This was partly explained by the far larger proportion of the burrow area that was refilled with casts: approximately 40% and 50% for Al. chlorotica and A. caliginosa, respectively compared with approximately 20% for the anecic burrows. We discuss whether these estimates could be used in future models simulating the dynamics of earthworm burrow systems by taking into account both burrow creation and destruction by earthworms.     

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.     

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.     

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.     

Relationships between earthworm communities and burrow numbers under different land use systems

This study addresses the influence of three different land use systems (continuous maize, pasture/maize rotation, permanent pasture) on the relationships between earthworm populations and the number of earthworm burrows quantified in a soil profile. Quantified burrows were limited to those observable by the naked eye (i.e. >2 mm in diameter) and enumerated earthworms were limited to those which could have created the observable burrows (i.e. >0.3 g).The results were combined with data from the literature coming from different geographical regions. This study showed that earthworm abundance decreased with the increasing land management intensity (maize crop vs. pasture), while the number of burrows could be higher or similar under maize compared to pasture. Under maize, despite lower earthworm abundance and the annual destruction of the burrows by tillage, the number of burrows was almost as high as under pasture. This absence of a relationship between burrow numbers and earthworm abundance was observed in the soil profile and for each layer of the profile for each land use system. Furthermore, the burrow number/m^?2 per earthworm strongly varied depending on land use and was far higher under maize when compared to pasture (74 vs. 7). Therefore, a power-law type relationship was clearly established between burrow number/m^?2 per earthworm and earthworm abundance. This power type relationship was also observed when including data from the literature although it followed a different mathematical model. These results were explained by (i) increased earthworm burrowing activity (i.e. an increase in the number of burrows produced by each earthworm) under maize, and (ii) the dynamics of burrow number under pasture (i.e. decreased burrowing activity and burrow destruction process); both results of food accessibility combined with inter-individual competition. The results of the study suggest that farmers should not use the number of pores as an indicator of earthworm abundance, but as an indicator of earthworm activity, which could be integrated in an indicator of soil quality.     

Burrow systems of endogeic earthworms: Effects of earthworm abundance and consequences for soil water infiltration

By creating burrows, earthworms influence the transfer properties of soils. The effects of endogeic species on soil transfer properties, however, are not yet well understood because these earthworms generally create burrows that are refilled by casts and have no preferential vertical orientation. Thirty soil cores were incubated for various periods (1–3 or 4 weeks) at different earthworm densities (70, 210, 345 or 480 individuals m⁻²). The cores were then scanned using X-ray tomography and the burrow systems were characterised by measuring the total burrow volume, bioturbation volume (refilled burrows and lateral compaction around the burrows), the number of branches, tortuosity and continuity (assessed by computing the number of burrows with a vertical extension greater than 15, 20 and 25% of the core). We also computed the mean geodesic distance, i.e. the mean distance from the bottom to the top of the core assuming that distances inside burrows are null. Rainfall simulations were carried out on 17 cores chosen to encompass the variations observed in the burrow systems. The water transfer efficiency of each core was estimated by measuring two parameters: breakthrough volume and the percentage of water transmitted after 1 h of rain. Burrow and bioturbation volume increased significantly and steadily with time and earthworm density. We estimated that on average Allolobophora chlorotica burrowed 22 cm per week. All other burrow system characteristics also increased with time and earthworm density except the mean geodesic distance, which decreased significantly. This suggests that intraspecific interactions had no significant effect on burrow system geometry. Univariate PLS regressions were used to understand which burrow system characteristics had the strongest influence on water transfer. These regressions showed that the mean geodesic distance was the most important parameter. This means that in addition to individual burrow characteristics, the spatial arrangement of the whole burrow system also had a major effect on transfer properties.     

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.     

The drilosphere concept: Fine-scale incorporation of surface residue-derived N and C around natural Lumbricus terrestris burrows

Anecic (deep-burrowing) earthworms are important for soil biogeochemical functioning, but the fine-scale spatial range at which they incorporate C and N around their burrows (the drilosphere sensu stricto) needs to be investigated under realistic conditions. We conducted a field experiment to delimit spatially the extent to which soil around natural Lumbricus terrestris burrows is influenced biochemically. We placed plant litter dual-labelled with ¹³C and ¹⁵N stable isotope tracers on L. terrestris burrow openings and we measured residue-derived ¹³C and ¹⁵N in thin concentric layers (0–2, 2–4, 4–8 mm) around burrows with or without a resident earthworm. After 45 days, earthworms were significantly enriched in ¹³C and ¹⁵N as a result of feeding on the plant litter. At 0–5 cm soil depth, soil ¹⁵N concentrations were significantly higher around occupied than unoccupied burrows, and they were significantly higher in all burrow layers (including 4–8 mm) than in bulk soil (50–75 mm from burrow). This suggests that biochemical drilosphere effects of anecic earthworms, at least in the uppermost portion of the burrow, extend farther than the 2 mm layer assumed traditionally.     

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.     

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.     

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.     

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.     

Earthworms,soil mineral nitrogen and forage production in grass-based hayfields

This study was designed to address how earthworm activity influences soil mineral nitrogen (N), plant N uptake and forage yield in grass-based hayfields. Earthworm populations were reduced by applying carbaryl pesticide to the experimental field plots every 2-weeks, effectively eliminating the earthworms for up to 12-weeks from May to August. Grass yields and tissue N concentrations were measured every 2 weeks, and the soil mineral N concentration determined at the final harvest. Reducing earthworm populations for up to 12-weeks did not affect grass yield or N uptake. However, regression analysis showed that plots with undisturbed earthworm populations had higher soil N by 0.8 kg N ha^?1 per week, representing mineralization of about 10 kg N ha^?1 during the 12-week study. This was a fraction of the fertilizer N recommendation (75 kg N ha^?1) for grass-based hayfields in this region. Therefore, the increase in soil mineral N from earthworm activity was small, relative to the N requirements of the hayfield.     

Earthworm collection from agricultural fields: Comparisons of selected expellants in presence/absence of hand-sorting

The role earthworms play in soil fertility is under increasing scientific scrutiny, especially in light of the fact that farmers are seeking to reduce soil tillage. However, there are many discrepancies in earthworm sampling methods. The aim of this study is to compare the efficiency of three chemical expellants (formaldehyde, commercial ‘hot’ mustard and allyl isothiocyanate, abbreviated AITC), with two sampling methods: (i) a simple method that consisted in spreading an expellant solution on the soil and retrieving earthworms that emerged at the soil surface, and (ii) a combined method that consisted in applying a chemical to expel earthworms and then hand-sorting the remaining earthworms from the block of soil. Sampling efficiency was measured in terms of earthworm density and biomass, for endogeic and anecic earthworms.With the simple method, a higher density of earthworms was sampled using formaldehyde and AITC than with mustard. Formaldehyde, AITC and mustard expelled not significantly different biomasses of 47.7, 31.9 and 20.5 g m^?2, respectively, on average over the three plots. The combined method did not yield a significantly different density or biomass with the different chemicals.Formaldehyde is toxic and commercial ‘hot’ mustard is difficult to standardise and inefficient when used without hand-sorting. Accounting for the accuracy of the sampling methods as well as the toxicity of the chemicals to users and soil organisms, AITC appears to be a reliable and promising chemical expellant whether or not in combination with hand-sorting. Its use would be a step towards standardizing earthworm sampling methods.     

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.     

Impact of soil engineering by two contrasting species of earthworms on their dispersal rates

By burrowing galleries and producing casts, earthworms are constantly changing the structure and properties of the soils in which they are living. These changes modify the costs and benefits for earthworms to stay in the environment they modify. In this paper, we measured experimentally how dispersal behaviour of endogeic and anecic earthworms responds to the cumulative changes they made in soil characteristics. The influence of earthworm activities on dispersal was studied in standardised mesocosms by comparing the influence of soils modified or not modified by earthworm activities on earthworm dispersal rates.The cumulative use of the soil by the earthworms strongly modified soil physical properties. The height of the soil decreased over time and the amount of aggregates smaller than 2 mm decreased in contrast to aggregates larger than 5 mm that increased. We found that: (i) earthworm activities significantly modified soil physical properties (such as bulk density, soil strength and soil aggregation) and decreased significantly the dispersal rates of the endogeic species, whatever the species that modified the soil; (ii) the decreasing in the dispersal proportion of the endogeic species suggests that the cost of engineering activities may be higher than the one of dispersal; (iii) the dispersal of the anecic species appeared to be not influenced by its own activities (intra-specific influences) or by the activities of the endogeic species (inter-specific influences). Overall these results suggest that the endogeic species is involved in a process of niche construction, which evolved jointly with its dispersal strategy.     

Emission of nitrous oxide from hydrocarbon contaminated soil amended with waste water sludge and earthworms

Soils in Mexico are often contaminated with hydrocarbons and addition of waste water sludge and earthworms accelerates their removal. However, little is known how contamination and subsequent bioremediation affects emissions of N₂O and CO₂. A laboratory study was done to investigate the effect of waste water sludge and the earthworm Eisenia fetida on emission of N₂O and CO₂ in a sandy loam soil contaminated with the polycyclic aromatic hydrocarbons (PAHs): phenanthrene, anthracene and benzo(a)pyrene. Emissions of N₂O and CO₂, and concentrations of inorganic N (ammonium (NH₄⁺), nitrite (NO₂^?) nitrate (NO₃^?)) were monitored after 0, 5, 24, 72 and 168 h. Adding E. fetida to the PAHs contaminated soil increased CO₂ production rate significantly 2.0 times independent of the addition of sludge. The N₂O emission rate from unamended soil expressed on a daily base was 5 μg N kg^?1 d^?1 for the first 2 h and increased to a maximum of 325 μg N kg^?1 d^?1 after 48 h and then decreased to 10 μg N kg^?1 d^?1 after 168 h. Addition of PAHs, E. fetida or PAHs + E. fetida had no significant effect on the N₂O emission rate. Adding sludge to the soil sharply increased the N₂O emission rate to >400 μg N kg^?1 d^?1 for the entire incubation with a maximum of 1134 μg N kg^?1 d^?1 after 48 h. Addition of E. fetida, PAHs or PAHs + E. fetida to the sludge-amended soil reduced the N₂O emission rate significantly compared to soil amended with sludge after 24 h. It was found that contaminating soil with PAHs and adding earthworms had no effect on emissions of N₂O. Emission of N₂O, however, increased in sludge-amended soil, but addition of earthworms to this soil and contamination reduced it.     

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.     

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.