Associations between soil texture,soil water characteristics and earthworm populations in grassland

Abstract

The aim of the present study was to investigate the relationships between soil physical characteristics and earthworms in a regional-scale field study in Denmark. The earthworm populations along within-field gradients in soil texture were quantified at five field sites, representing dominant soil types of Denmark. Eleven earthworm species were found, but populations were mainly dominated by Aporrectodea tuberculata and A. longa. Despite considerable variation in soil parameters across the five study sites the results suggest that the biomass of anecic worms (or A. longa as a species) was not causally associated with the soil parameters studied. This indicates that there must be other causal factors associated with the abundance (and composition) of anecic worms that are not among the soil texture and structure parameters studied. On the other hand, soil texture (Coarse sand) was associated with the abundance of the dominant endogeic species, A. tuberculata, but not endogeic worms in general. It was hypothesized that anecic and endogeic earthworms might respond to local soil water characteristics rather than soil texture, but this hypothesis could not be confirmed with the present data.     

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.     

Earthworm burrowing behaviour in 2D terraria with single- and multi-species assemblages

Experiments were performed in 2D terraria to investigate the burrowing behaviour of different earthworm species from various ecological categories in single- and multi-species assemblages. The burrowing behaviour was quantified using image analysis software during a 2-week period. Terraria were found to reveal realistic impressions of the burrowing behaviour of various species according to the ecological classification of Bouché into epigeic, endogeic and anecic species. Results of the study also permit the recommending of classifying various earthworms as intermediate species, e.g. Aporrectodea longa as endo-anecic and Lumbricus rubellus as epi-endogeic. Burrowing activity of endogeic species was significantly reduced in multi-species compositions compared to single-species treatments. Moreover, burrowing activity of Octolasion tyrtaeum was significantly reduced in the presence of Lumbricus terrestris compared to the specific single treatment. This endogeic species profited from the burrowing behaviour, e.g. bioturbation, of L. terrestris, and as such this is circumstantial evidence for commensalism (species interaction, in which one partner benefits while the other is unaffected) between anecic and endogeic earthworm species. Simultaneous burrowing activity of a combined assemblage of both endogeic species, Aporrectodea caliginosa and O. tyrtaeum, was also significantly reduced compared with the particular single treatments. Thus, this seems likely to be a response of interspecific competition and trophic niche separation between endogeic species.     

Spatio-temporal variation of stable isotope ratios in earthworms under grassland and maize cropping systems

We investigated the specific diet and habitat of earthworms in relation to land use changes by integrating spatial and temporal scales and by using stable isotope (¹³C and ¹⁵N) techniques. The study involved two sites: Santiago (Northwest Spain) and North Wyke (Southwest England), both consisting of long term grassland which was partly converted to a maize crop in 1997. In 1998, the maize crop in Spain was divided into two, and one half was re-planted with maize (2 years maize) and the other half reconverted to grassland (1 year grassland); the same procedure was followed for the grassland resulting in two treatments, 2 years grassland and 1 year maize. At the English site only 2 years maize and 2 years grassland were under investigation. Within each of the four treatments in Spain and the two in England, three replicate plots were established.Random soil samples from three different depths (0–10, 10–20 and 20–30 cm), and earthworm specimens belonging to four different ecological categories (epigeic, anecic, epi/anecic and endogeic), were taken from each plot, treatment and site at the peak of the maize growth and after harvesting.Spanish soils were ¹³C-enriched and ¹⁵N-depleted when compared to the English ones, which was also reflected in the earthworm tissue, and allowed a direct relationship between the delta values of the animals and the cropping treatments.The enrichment in the ¹³C values of the worms feeding on the maize (C₄) plots, when compared to those found under the grassland (C₃) plots, was greater than the difference detected in the C₄ vs C₃ soils. This result clearly indicates selective feeding by earthworms with a preference for fresh C₄ residue over older native C₃.Different ecological and age groups appeared to consume organic material of differing quality, with endogeic species and mature worms showing the highest N isotope values as a result of preferential feeding in deeper soil profiles. This information proves that combined C and N isotope analysis constitutes a powerful tool in studying feeding ecology and emphasises the need for long-term studies which incorporate spatial and temporal scales to the experimental set-up.     

Does the deep-burrowing earthworm,Aporrectodea longa,compete with resident earthworm communities when introduced to pastures in south-eastern Australia?

Pastures in southern Australia are dominated by endogeic earthworms such as Aporrectodea caliginosa (Sav.). Introductions of the anecic earthworm, A. longa (Ude), which is mostly restricted to Tasmania at present, are likely to increase the functional diversity of local communities and thereby enhance plant production and agricultural sustainability. However, the potential impact of A. longa on resident earthworm communities first needs assessing. Glasshouse and field experiments reported here suggested that A. longa can reduce the abundance and biomass of A. caliginosa, but that these effects are likely to be offset by overall increases in worm abundance. There was no evidence to suggest that species richness was reduced by A. longa introduction.     

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.     

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.     

The effect of feeding behavior on Hg accumulation in the ecophysiologically different earthworms Lumbricus terrestris and Octolaseon cyaneum: A microcosm experiment

 A soil microcosm experiment was performed to assess the uptake of Hg from various Hg-spiked food sources (soil, leaf litter and root litter of Trifolium alexandrinum) by two earthworm species, Lumbricus terrestris (anecic) and Octolaseon cyaneum (endogeic). Treatments were applied in which one of the three food sources was Hg spiked and the other two were not. Additional treatments in which all or none of the food sources were Hg spiked were used as controls. Uptake of Hg from soil into tissues of both earthworm species was significantly higher than uptake of Hg from leaf litter or root litter, indicating that soil may be the most important pool for the uptake of Hg into earthworms. In addition, the anecic L. terrestris significantly accumulated Hg from all Hg-spiked food sources (leaf litter, root litter and soil), whereas the endogeic O. cyaneum took up Hg mainly from soil particles. Interestingly, there was no further increase in Hg in L. terrestris when all food sources were Hg spiked compared to the single Hg-spiked sources. This may be attributed to the relatively high Hg content in the soil, which may have influenced the feeding behavior of the earthworms, although their biomass did not significantly decline. We suggest that, in addition to the physiological differences, feeding behavior may also play a role in the contrasting uptake of Hg by the two earthworm species.     

C and N turnover of fermented residues from biogas plants in soil in the presence of three different earthworm species (Lumbricus terrestris,Aporrectodea longa,Aporrectodea caliginosa)

A soil microcosm experiment was performed to assess (1) the C- and N- turnover of residues from biogas plants in soils in the presence of three earthworm species (Lumbricus terrestris, Aporrectodea longa and Aporrectodea caliginosa) and (2) the resulting changes in soil chemical and microbiological properties when using these residues as fertilizer in comparison to conventional slurry. Earthworms were exposed in soils, fertilized with an equivalent amount of 120 kg of NH₄-N ha^?1 from: (1) conventional cattle slurry and (2) a fermented residue derived from cattle slurry, grass (silage) and maize. Additional treatments without slurry and earthworms were used as controls.There was considerable evidence that soils fertilized by fermented slurry comprised fewer amounts of readily available nutrients for microbial C and N turnover. We observed significant stimulation of microbial biomass, basal respiration and nitrification in treatments with conventional slurry, especially in the presence of earthworms. However, the stimulation of microbial activity by manure and earthworms were significantly lower in treatments with fermented slurry. Moreover, the results showed clear interactions between different earthworm species and manures. While the biomass of the anecic species (L. terrestris and A. longa) increased in both slurry treatments, the biomass of A. caliginosa (endogeic) decreased, with a significantly stronger biomass decline in treatments with fermented slurry. The metabolic quotients revealed microbial stress metabolism in fermented slurry treatments, predominantly in treatments with A. caliginosa. We conclude that particularly A. caliginosa and soil microorganisms competed for labile C sources in treatments with fermented slurry. An application of these residues as fertilizer might result in a reduction of microbial activity in agricultural soils and in a decline of endogeic earthworms.     

Plant influences on native and exotic earthworms during secondary succession in old tropical pastures

Land-use changes can drastically alter earthworm communities. Native species are often lost and few exotic species, such as Pontoscolex corethrurus, rapidly prevail when tropical forests are converted to pastures. However, this process can be reversed when forests recover from abandoned pastures through secondary succession. We hypothesized (1) that the formation of forest floor mass during secondary succession in pastures promotes the recovery of native, anecic earthworms and (2) that the shift from grass vegetation in pastures to woody plants in secondary forests decreases the abundance and biomass of the exotic, endogeic P. corethrurus. To test the first hypothesis, we developed a litter manipulation experiment by removing and adding plant litter in plots of mature secondary forests in the Cayey Mountains, Puerto Rico. To test the second hypothesis we performed a greenhouse experiment to examine the influence of a pasture grass species Axonopus compressus and a dominating woody species Miconia prasina of the secondary forests on the number and biomass of the earthworm P. corethrurus. We found in the litter manipulation experiment that earthworm diversity, density and fresh weight were not affected by litter input. However, in the greenhouse experiment, A. compressus increased the number and biomass of P. corethrurus, whereas M. prasina decreased the exotic, endogeic earthworm. Our results suggest that the quantity of litter does not promote rapid changes (<1 year) in native, anecic earthworm diversity, and that the exotic, endogeic P. corethrurus is favored by grass A. compressus compared to the woody plant M. prasina. The shift in vegetation from grass to woody plants promotes the decrease in the density and biomass of the exotic, endogeic P. corethrurus during secondary succession in old tropical pastures.     

Earthworm community development in soils of a reclaimed steelworks

Long-term studies are essential to learn earthworm community development and soil formation post reclamation. Investigations were undertaken at a former steelworks site at Hallside, near Glasgow, UK, reclaimed in the 1990s using a mixture of colliery spoil and sewage sludge. The site was largely planted for production of short rotation coppice willow (Salix spp.). Earthworm inoculation formed a part of the restoration process. Minimal monitoring occurred in the interim, but some records of earthworm sampling existed in 2000 and 2005. This study focused on monitoring earthworms and soil properties across the site, drawing comparisons with adjacent unspoiled soil. Results showed that after 22 years, a species-rich community of earthworms (n=16) colonized the site, with endogeic Aporrectodea caliginosa being dominant by number and anecic A. longa by mass. Across the site, earthworm community density and biomass were 208 individuals m^-2 and 71 g m^-2, respectively. The Shannon diversity index for earthworms was 1.89, with an evenness of 0.68. The sewage sludge increased the soil organic matter, but the stone content of the colliery spoil prevented digging in some locations. Soil chemistry had no negative effect on earthworms, but the compacted substrate did hinder water infiltration. Earthworms colonized the reclaimed site from adjacent areas, and community structure and density below well-drained, scrub-free willow, birch, and grassland were not significantly different (P>0.05) from those of the adjacent unspoiled areas. The results show that the historical earthworm inoculation was unnecessary and badly timed. Future reclamations of similar sites can learn from this investigation.     

Interactions between residue placement and earthworm ecological strategy affect aggregate turnover and N2O dynamics in agricultural soil

Previous laboratory studies using epigeic and anecic earthworms have shown that earthworm activity can considerably increase nitrous oxide (N₂O) emissions from crop residues in soils. However, the universality of this effect across earthworm functional groups and its underlying mechanisms remain unclear. The aims of this study were (i) to determine whether earthworms with an endogeic strategy also affect N₂O emissions; (ii) to quantify possible interactions with epigeic earthworms; and (iii) to link these effects to earthworm-induced differences in selected soil properties. We initiated a 90-day ¹⁵N-tracer mesocosm study with the endogeic earthworm species Aporrectodea caliginosa (Savigny) and the epigeic species Lumbricus rubellus (Hoffmeister). ¹⁵N-labeled radish (Raphanus sativus cv. Adagio L.) residue was placed on top or incorporated into the loamy (Fluvaquent) soil. When residue was incorporated, only A. caliginosa significantly (p < 0.01) increased cumulative N₂O emissions from 1350 to 2223 μg N₂O-N kg⁻¹ soil, with a corresponding increase in the turnover rate of macroaggregates. When residue was applied on top, L. rubellus significantly (p < 0.001) increased emissions from 524 to 929 μg N₂O-N kg⁻¹, and a significant (p < 0.05) interaction between the two earthworm species increased emissions to 1397 μg N₂O-N kg⁻¹. These effects coincided with an 84% increase in incorporation of residue ¹⁵N into the microaggregate fraction by A. caliginosa (p = 0.003) and an 85% increase in incorporation into the macroaggregate fraction by L. rubellus (p = 0.018). Cumulative CO₂ fluxes were only significantly increased by earthworm activity (from 473.9 to 593.6 mg CO₂-C kg⁻¹ soil; p = 0.037) in the presence of L. rubellus when residue was applied on top. We conclude that earthworm-induced N₂O emissions reflect earthworm feeding strategies: epigeic earthworms can increase N₂O emissions when residue is applied on top; endogeic earthworms when residue is incorporated into the soil by humans (tillage) or by other earthworm species. The effects of residue placement and earthworm addition are accompanied by changes in aggregate and SOM turnover, possibly controlling carbon, nitrogen and oxygen availability and therefore denitrification. Our results contribute to understanding the important but intricate relations between (functional) soil biodiversity and the soil greenhouse gas balance. Further research should focus on elucidating the links between the observed changes in soil aggregation and controls on denitrification, including the microbial community.     

Vertical distribution of earthworms in grassland soils of the Colombian Llanos

 The vertical distribution of native earthworm species from natural and disturbed savannas in the Oxisols of the Colombian Llanos was assessed in a native savanna and in a 17-year-old grazed grass-legume pasture during a period of 17 months. Different patterns of vertical stratification were observed for all species with a strong migration of populations to deeper layers in the dry season. The correlation between the size of the earthworms and the average depth at which they were found was not significant (P>0.05), despite the fact that bigger species are located deeper in the soil. The living habits and adaptive strategies of the smallest species, Ocnerodrilidae n. sp., found in both ecosystems studied are responsible for this pattern. This endogeic species is associated with organic pools generated by an anecic species and further studies should assess the role of this species in ecosystem functioning. Mature worms of one anecic species were located deeper than immature ones in the soil (P<0.01). Soil moisture had an important effect on the vertical distribution of earthworms, although differences between immature and mature worms of the anecic Martiodrilus carimaguensis are likely to be of biotic origin. New data on the biology and ecology of these Neotropical species are shown. Received: 24 November 1999     

Different earthworm ecological groups interactively impact seedling establishment

There is increasing evidence that direct interactions between earthworms and seeds impact the assembly of plant communities. However, effects of earthworms of different ecological groups and their interactions on plant germination and establishment are little known. We set up a full-factorial greenhouse experiment in order to explore impacts of different ecological groups of earthworms (epigeic, endogeic and anecic) on the establishment of seedlings. The three ecological groups of earthworms affected seedling establishment in an interactive way with the effects varying in time. While anecic earthworms detrimentally affected the number of established seedlings, impacts of epigeic and endogeic species depended on the presence of earthworms belonging to other ecological groups. Impacts of anecic earthworms were more significant and consistent than those of epigeic and endogeic ones pointing to the dominance of the effect of anecic earthworms for seedling establishment. The initial positive effect of the combined presence of epigeic and endogeic earthworms compared to treatments with only endogeic and only anecic earthworms was likely due to the acceleration of seed germination and elevated nutrient availability for seedlings in earthworm casts. By contrast, reduced numbers of seedlings in presence of both epigeic and endogeic earthworms compared to the control treatment might have been due to seedling predation after the litter layer had disappeared. The results extend current knowledge on interactions between earthworms and seeds by showing that, in addition to anecic species, also epigeic and endogeic species influence seedling establishment with their effect depending on the presence of anecic species. Moreover, the results indicate that impacts of earthworms vary with the developmental stage of seedlings highlighting the importance of interactions between earthworms and seeds.     

The fate of catechol in soil as affected by earthworms and clay

The effect of endogeic earthworms (Octolasion tyrtaeum) and the availability of clay (Montmorillonite) on the mobilization and stabilization of uniformly ¹⁴C-labelled catechol mixed into arable and forest soil was investigated in a short- and a long-term microcosm experiment. By using arable and forest soil the effect of earthworms and clay in soils differing in the saturation of the mineral matrix with organic matter was investigated. In the short-term experiment microcosms were destructively sampled when the soil had been transformed into casts. In the long-term experiment earthworm casts produced during 7 days and non-processed soil were incubated for three further months. Production of CO₂ and ¹⁴CO₂ were measured at regular intervals. Accumulation of ¹⁴C in humic fractions (DOM, fulvic acids, humic acids and humin) of the casts and the non-processed soil and incorporation of ¹⁴C into earthworm tissue were determined.Incorporation of ¹⁴C into earthworm tissue was low, with 0.1 and 0.44% recovered in the short- and long-term experiment, respectively, suggesting that endogeic earthworms preferentially assimilate non-phenolic soil carbon. Cumulative production of CO₂-C was significantly increased in casts produced from the arable soil, but lower in casts produced from the forest soil; generally, the production of CO₂-C was higher in forest than in arable soil. Both soils differed in the pattern of ¹⁴CO₂-C production; initially it was higher in the forest soil than in the arable soil, whereas later the opposite was true. Octolasion tyrtaeum did not affect ¹⁴CO₂-C production in the forest soil, but increased it in the arable soil early in the experiment; clay counteracted this effect. Clay and O. tyrtaeum did not affect integration of ¹⁴C into humic fractions of the forest soil. In contrast, in the arable soil O. tyrtaeum increased the amount of ¹⁴C in the labile fractions, whereas clay increased it in the humin fraction.The results indicate that endogeic earthworms increase microbial activity and thus mineralization of phenolic compounds, whereas clay decreases it presumably by binding phenolic compounds to clay particles when passing through the earthworm gut. Endogeic earthworms and clay are only of minor importance for the fate of catechol in soils with high organic matter, clay and microbial biomass concentrations, but in contrast affect the fate of phenolic compounds in low clay soils.     

Spatial distribution of earthworms [Lumbricidae] in recultivated soils of the Rhenish lignite‐mining area,Germany

The spatial distribution of earthworms was studied by means of combined formalin expulsion and hand sorting in three arable fields of the Rhenish lignite‐mining area that differed in their recultivation age (6, 12, 24 yr). In addition, pH and the spatial distribution of penetration resistances were measured to see if they are corresponding with the distribution of earthworms. Already the 6 yr old field had a rich population of endogeic, anecic, and epigeic earthworms (119 ind. m^–2, 48 g m^–2, 6 species). This quantity was similar to the 24 yr old site. The 12 yr old field was only sparsely populated by earthworms (5 ind. m^–2, 5 g m^–2, 3 species). In the 6 yr old field, the spatial distribution pattern showed a center of maximal earthworm abundances, corresponding to the distributional pattern of penetration resistances. In the old field (24 yr), the species varied in their spatial distribution, and there was no correspondence with the distribution of penetration resistance. In general, the penetration resistance at the youngest site was clearly lower than at the two older sites. The earthworm population in the 6 yr old field can be explained by cocoons contained in the dumped material. A calculation using literature data on earthworm‐population dynamics shows that a founding population of 400–600 reproductive individuals per hectare and a continuity of favorable growth conditions during the time of soil management is necessary for the development of the situation found at the 6 yr old site in this study.     

Hierarchical classification of environmental factors and agricultural practices affecting soil fauna under cropping systems using Bt maize

The population dynamics of soil organisms under agricultural field conditions are influenced by many factors, such as pedology and climate, but also farming practices such as crop type, tillage and the use of pesticides. To assess the real effects of farming practices on soil organisms it is necessary to rank the influence of all of these parameters. Bt maize (Zea mays L.), as a crop recently introduced into farming practices, is a genetically modified maize with the Cry1Ab gene which produces a protein toxic to specific lepidopteran insect pests. To assess the effects of Bt maize on non-target soil organisms, we conducted research at a field site in Foulum (Denmark) with a loamy sand soil containing 6.4% organic matter. The study focused on populations of springtails (Collembola) and earthworms (Oligochaeta) from samples taken at the beginning and at the end of the maize crop-growing season during 2 consecutive years. Farming practices, soil parameters, the biological structure of soil communities, and the type and age of the crop at the time of sampling, were used as attributes to predict the total abundance of springtails and biomass of earthworms in general and the abundance or biomass for specific functional groups (epigeic, endogeic and anecic groups for earthworms, and eu-, eu to hemi-, hemi-, hemi to epi- and epiedaphic groups for Collembola). Predictive models were built with data mining tools, such as regression trees that predict the value of a dependent variable from a set of independent variables. Regression trees were constructed with the data mining system M5′. The models were evaluated by qualitative and quantitative measures of performance and two models were selected for further interpretation: anecic worms and hemi-epiedaphic Collembola. The anecic worms (r²=0.83) showed preferences for less clay and more silt soil with medium pH but were not influenced directly by farming practices. The biomass of earthworms was greater in early autumn than in spring or late autumn. Biomass of hemi-epiedaphic Collembola (r²=0.59) increased at the end of the maize growing season, while higher organic matter content and pH tended to increase their biomass in spring. Greater abundance of Collembola was also noted in early autumn if the crop was non-Bt maize. The models assessed by this research did not find any effects of the Bt maize cropping system on functional groups of soil fauna.     

Impact of five different tillage systems on soil organic carbon content and the density,biomass, and community composition of earthworms after a ten year period

To assess the impact of different types of soil tillage on the density, biomass, and community composition of earthworms, a long-term field study was performed in which soils were tilled in different ways for ten years. This study included five different types of tillage: (i) plough, (ii) grubber, (iii) disc harrow, (iv) mulch sowing, and (v) direct sowing. At the end of the experiment the earthworm density, biomass, and community composition, and the SOC (soil organic carbon) content were determined. The results show that density, biomass, and community composition of earthworm populations varied in relation to the type of soil tillage used. The density of anecic earthworm species decreased when soils were managed by conventional ploughing, relative to reduced tillage practices, whereas conversely the density of endogeic species increased. Additionally, the varying types of soil tillage influenced the abundance and biomass of different earthworm species in different ways. The density of Aporrectodea caliginosa was positively influenced by ploughing, whereas Aporrectodea longa, Lumbricus castaneus, and Satchellius mammalis showed a positive relationship to the grubber and Allolobophora chlorotica to direct sowing. We attribute these changes to modifications in the vertical distribution of SOC and varying potentials for mechanical damage of earthworms by tillage. A decrease in tillage intensity modified the vertical SOC distribution in the topsoil and consequently revealed positive effects on earthworm biodiversity, thus sustaining soil functioning.     

Axial and radial pressure exerted by earthworms of different ecological groups

 The aim of this study was to measure the pressures exerted by earthworms during burrowing. For this purpose we developed two methods with which to quantify the axial and radial pressure. The data were recorded with an electronic balance that was connected to a PC. Artificial earthworm burrows were used to standardize the measurements. Plexiglas tubes with diameters ranging from 2 to 6.3 mm which corresponded to the diameter of the earthworms were used. A pin was placed inside the tubes, on which the earthworms exerted a pressure by peristaltic locomotion. Only the maximum values of the pressure measurement were taken into account for evaluation, and the arithmetic mean was calculated. The measurements were conducted with Aporrectodea longa, Lumbricus terrestris, Aporrectodea caliginosa, Octolasion cyaneum, Allolobophora chlorotica, Aporrectodea rosea, Lumbricus rubellus and Dendrobaena octaedra. The species examined were classified into ecological groups. The mean axial pressures exerted by each group were in the order: epigeic (14–25 kPa), endogeic (27–39 kPa) and anecic (46–65 kPa). For the mean radial pressure the order was: epigeic (39–63 kPa), anecic (72–93 kPa) and endogeic (59–195 kPa). It was apparent from the results that radial pressure is the most important pressure with respect to the burrowing activity of earthworms. Received: 28 April 1998     

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.