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.     

Organic carbon sequestration in earthworm burrows

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

3D skeleton reconstructions of natural earthworm burrow systems using CAT scan images of soil cores

 Four soil cores (length, 20 cm; diameter, 16 cm) were sampled in a Swiss pre-Alpine meadow with high earthworm abundance (>400 individuals/m²); two cores were taken in October 1993 and the other two cores in April 1994. The cores were described using computer assisted tomography which gives a series of section images every 3 mm. A method for reconstructing the three-dimensional (3D) skeleton of the earthworm burrow system is presented and discussed. This method provides an image of the structural organisation of the burrow system and was found to be adequately sensitive for use in ecological and functional studies. The seasonal variation of these 3D skeletons was investigated using two approaches, i.e. the analysis of: (1) global burrow system characteristics, and (2) individual burrow characteristics. At the scale of the global burrow system no difference was found between seasons (same number of burrows and same total burrow length) except for the vertical segment distribution, which was homogeneous in spring and decreased with depth in the fall. The study of individual burrow characteristics revealed that burrows tended to be more vertical in spring and that their branching intensity was higher in this season. Received: 10 June 1997     

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.     

Impact of soil compaction on earthworm burrow systems using X-ray computed tomography: preliminary study

This study is a first approach of the impact of soil compaction due to trafficking by machinery on earthworm burrow systems. To this end, two experiments were established. In the first one, microcosms were incubated in the laboratory for 70 d with Lumbricus terrestris or Aporrectodea giardi. In the second experiment, soil cores were excavated from a sugar beet field mainly colonised by L. terrestris and Aporrectodea caliginosa. The cores were then artificially compacted at 0.12 MPa or 0.25 MPa (which corresponds to the compaction due to trafficking by machinery in the field) or remained non-compacted. The whole cores were submitted to an X-ray computed tomography scan. This method allowed to compare the characteristics of the entire burrow system (total and mean burrowed length, mean length and number of burrows) and of the burrows themselves (number, area and roundness of pores constituting the burrows) in the compacted and non-compacted cores. The results showed that soil compaction contributes to close numerous pores, reduce mean length of burrows and increase the number of fragmented burrows. We concluded that soil compaction affects to a large extent the functionality of burrow systems by fragmenting them and affecting their continuity. This impact increases with the intensity of compaction. The applied soil compaction mainly affected the burrows located in the upper part of the soil. From a general point of view and especially for the upper part of the cores, horizontal burrows seemed to be more affected by compaction than vertical or oblique ones. Globally, the effects of soil compaction were especially observed at the greatest applied pressure (0.25 MPa). At 0.12 MPa, the damage to the burrows was less obvious.     

Earthworm burrowing activity of two non-Lumbricidae earthworm species incubated in soils with contrasting organic carbon content (Vertisol vs. Ultisol)

The aim of this study was to investigate the burrowing activity of two earthworm species: the endogeic Drawida sinica and one undescribed Amynthas species incubated in Vertisol and Ultisol presenting different soil organic C content. Because of their contrasting feeding behaviours, we hypothesised that soil type would have a bigger influence on the burrowing activity of the endogeic than the anecic species. Repacked soil columns inoculated with earthworms for 30 days were scanned using X-ray tomography and the compiled images used to characterise the burrow systems. After scanning, the saturated hydraulic conductivity (K _sat) was also measured. The Amynthas species burrows were less numerous (30 vs. 180), more vertically oriented (57 vs. 37°), more connected from the surface to the bottom of the columns (73 vs. 5 cm³) and had a higher global connectivity index (83 vs. 28%) than those of D. sinica. The K _sat was threefold faster in columns incubated with Amynthas and was linked to the volume of percolating burrows (R ² = 0.81). The soil type did not influence Amynthas burrow characteristics. In contrast, there were 30% more D. sinica burrows in the Vertisol than in the Ultisol while other burrow characteristics were not affected. This result suggests that these burrows were more refilled with casts leading to shorter and discontinuous burrows. The K _sat was negatively related to the number of burrows (R ² = 0.44) but was not statistically different between the Vertisol and the Ultisol, suggesting a constant impact of this species on the K _sat. We found that a decrease in the amount of soil organic C by 50% had only a small influence on earthworm burrowing activity and no effect on the K _sat.     

Earthworm burrowing in laboratory microcosms as influenced by soil temperature and moisture

Earthworm burrows contribute to soil macroporosity and support diverse microbial communities. It is not well known how fluctuations in soil temperature and moisture affect the burrowing activities of earthworms. The objective of this experiment was to evaluate the maximum depth and length of burrows created by the endogeic earthworm Aporrectodea caliginosa (Savigny) and the anecic earthworm Lumbricus terrestris L. for a range of temperatures (5–20 °C) and soil water potentials (−5 and −11 kPa). The laboratory microcosm was a plexiglass chamber (45 cm high, 45 cm wide) containing 0.14 m² of pre-moistened soil and litter, designed to house a single earthworm for 7 days. Earthworm mass, surface casting and burrowing activities were affected significantly by soil temperature, moisture and the temperature×moisture interaction. Burrow length and maximum burrow depth increased with increasing temperature, but there was less burrowing in wetter soil (−5 kPa) than drier soil (−11 kPa). Weight gain and surface casting, however, were greater in soil at −5 kPa than −11 kPa. Our results suggest more intensive feeding and limited burrowing in wetter soil than drier soil. Earthworms inhabiting the non-compacted, drier soil may have pushed aside particles without ingesting them to create burrows. The result was that earthworms explored a larger volume of soil, deeper in the chamber, when the soil was drier. How these burrowing activities may affect the community structure and activity of soil microorganisms and microfauna in the drilosphere remains to be determined.     

Sublethal effects of imidacloprid on the burrowing behaviour of two earthworm species: Modifications of the 3D burrow systems in artificial cores and consequences on gas diffusion in soil

Earthworms have been termed ‘ecosystem engineers’ (sensu [Jones, C.G., Lawton, J.H., Shachak, M., 1994. Organisms as ecosysem engineers. Oikos 69, 373-386.]) because of the important roles they play in the soil. As a consequence, it is assumed that if earthworms change their behaviour following exposure to pesticides or pollutants this could have a drastic impact on soil functioning. To test this assumption under laboratory conditions, we studied the burrow systems made by two earthworm species (the anecic Aporrectodea nocturna and the endogeic Allolobophora icterica) in artificial soil cores containing imidacloprid, a widely used neonicotinoid insecticide. After 1-month incubation period, the macropores created in the soil core were analyzed by tomography. In order to further characterize transfer properties associated with burrow systems gas diffusion measurements were also carried out. The burrow systems made by the two earthworm species were very different: A. nocturna made more continuous, less branched, more vertical and wider burrows than A. icterica. Some changes to A. nocturna burrow systems were observed after exposure to imidacloprid (they made a smaller burrow system and burrows were more narrow), but only at the highest concentration of imidacloprid used (0.5 mg kg⁻¹). A. icterica worms were more sensitive to imidacloprid and many differences in their burrow systems (length, sinuosity, branching rate and number of burrows) were observed at both concentrations tested (0.1 and 0. 5 mg kg⁻¹). As a consequence, the continuity of the burrow systems made by both species was altered following imidacloprid treatment. Gas diffusion through the A. nocturna soil cores was reduced but no difference in gas diffusion was observed in the A. icterica soil cores.     

Assessment of earthworm contribution to soil hydrology: a laboratory method to measure water diffusion through burrow walls

The capacity for water diffusion in burrow walls (i.e. the coefficient of sorptivity) either burrowed by Lumbricus terrestris (T-Worm) or artificially created (T-Artificial) was studied through an experimental design in a 2D terrarium. In addition, the soil density of earthworm casts, burrow walls (0–3 mm around the burrow) and the surrounding soil (>3 mm) were measured using the method of petroleum immersion. This study demonstrated that the quantity of water which transits through burrows of L. terrestris in the soil matrix was lower than that transited through soil fractures, due to a reduction of soil porosity in burrow walls (compaction: cast > worms burrow walls > surrounding soil > artificial burrow walls). Earthworm behaviour, in particular burrow reuse with associated cast pressing on walls, could explain the larger burrow wall compaction in earthworm burrows. If water diffusion was lower through the compacted burrows, burrow reuse by the worms makes them more stable (worms would maintain the structure over years) than unused burrows. The present experimental design could be used to test and measure the specific differences between earthworm species in their contributions to water diffusion. Probably, these contributions depend on the presumed related-species behaviours which would determine the degree of burrow wall compaction.     

Experimental evidence for the role of earthworms in compacted soil regeneration based on field observations and results from a semi-field experiment

In laboratory experiments, earthworms are often observed to burrow through compacted soil layers, leading to the general assumption that these animals play a significant role in regenerating compacted soils in agricultural plots. To demonstrate this role under field conditions, the abundance of earthworm macropores inside compacted zones was estimated on plots under reduced (RT) or conventional tillage (CT). Then, different types of compacted zones typically found in CT (plough pan and compacted clods) and RT plots (compacted volume under wheel tracks) were experimentally simulated in wooden boxes, buried in the field and inoculated with different earthworm species. After 6 weeks of incubation, the number of macropores inside the compacted zones was examined. Field observations showed that approximately 10% and 30% of the compacted zones were colonised by at least one macropore in CT and RT plots, respectively. A significantly greater number of anecics was found in RT plots, but we could not conclude that this ecological type of earthworm plays a more major role in the regeneration process in these plots since there were fewer compacted zones and these covered a smaller area in CT. The semi-field experiment provided evidence that earthworm-mediated regeneration of compacted zones is possible and its nature varies between ecological types of earthworm. Lumbricus terrestris, which makes individual burrows that are vertical and deep, was the main species to cross through the plough pan. The other three earthworm species (Aporrectodea giardi, A. caliginosa and A. rosea) did burrow inside the other types of compacted zones (“wheel tracks” and “compacted clods”). In every case, however, macropore density was far greater in non-compacted zones, illustrating that avoidance of compacted soil by earthworms is important and should be taken into account when extrapolating results from laboratory studies.     

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.     

Earthworm composition,diversity and biomass under three land use systems in south-eastern Australia

In south-eastern Australia, strips of planted native trees and shrubs (shelterbelts) are frequently established to restore ecosystem services altered by agriculture. Despite their wide use, little is known about the effects of establishing shelterbelts on soil macro invertebrates, especially earthworms, which are of major importance in soil processes. We assessed earthworm composition, diversity and biomass in three land use systems: native shelterbelts dominated by Acacia and Eucalyptus species, agricultural pastures and native remnant woodland fragments dominated by Eucalyptus blakelyi and/or Eucalyptus melliodora. Earthworm communities differed significantly among systems, with abundance, biomass and diversity greatest under pasture. Within shelterbelts we saw a shift from high earthworm biomass and density to low with increasing time after establishment. Soil edaphic variables did not correlate strongly with earthworm biomass or density, but were correlated with earthworm community composition. Overall the introduction of native woody vegetation was associated with a decline in density and biomass of earthworms, including a decrease in the relative abundance of exotic species. As such shelterbelts can be used to promote native earthworm relative abundance, which may be important for local diversity, soil function and landscape connectivity.     

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.     

Burrow systems made by Aporrectodea nocturna and Allolobophora chlorotica in artificial cores: morphological differences and effects of interspecific interactions

Burrow systems of earthworms contained in artificial cores were analysed through X-ray computed tomography and 3D skeleton reconstructions. Gas diffusion experiments were carried out on these cores to characterize soil transfer properties associated with the different burrow systems. Three types of cores were studied: cores in which Aporrectodea nocturna, an anecic earthworm was introduced (treatment 1), cores in which Allolobophora chlorotica, an endogeic earthworm was introduced (treatment 2) and cores that contained both species (treatment 3). Comparisons of the characteristics of the burrow systems of treatment 1 and 2 show important differences: the burrow system of A. nocturna comprises fewer burrows, which are longer, less branched, more vertical and have a lower sinuosity. The burrow system of A. chlorotica is characterized by lower continuity, which however did not result in a lower soil diffusivity. To study the burrow systems made by the two species in the same core (treatment 3), a separation that was based on differences in pore diameter between the two species and that takes into account the burrow orientation was designed. This separation was proven to be efficient since it resulted in low percentages of errors (around 10%) when applied to the burrow systems of treatments 1 and 2. Comparison of the burrow systems from treatments 1 and 3 demonstrated that the burrow system of A. nocturna was influenced by the presence of A. chlorotica: in treatment 3, A. nocturna made burrows that were smaller, more vertical and less branched. However, these interactions have to be confirmed under natural conditions.     

What happens to earthworm casts in the soil? A field study of carbon and nitrogen dynamics in Neotropical savannahs

In the Oxisols of the eastern plains of Colombia, the large native anecic earthworm Martiodrilus sp. is an abundant ecosystem engineer producing long-lasting casts and burrows. Casts deposited in the soil by this species have been estimated at several tonnes per hectare per year. The physical and chemical processes occurring in these casts have never been studied. In this study, we compared the dynamics of water content (WC), total C (C_tot), and available N (N_avail) contents, and the distribution in size of aggregates in ageing below-ground casts of this species and in the bulk soil. In a native herbaceous savannah and a sown grass/legume pasture (Brachiaria humidicola, Arachis pintoi, Desmodium ovalifolium and Stylosanthes capitata), fresh surface casts were experimentally injected into artificial burrows of 1 cm Ø and 10 cm depth and sampled at different dates during a total period of 120 days. The injection procedure used resulted in a 34% decrease in WC of the casts from the sown pasture and reduced the mean mass diameter (MMD) of the aggregates of casts from the savannah by 19%. Other properties were not significantly affected by the procedure.For injected casts in both grasslands, MMD and C_tot were stable during cast ageing while WC and N_avail were initially at levels several times higher than the bulk soil and decreased to similar bulk soil values with ageing. The C_tot was twice and one third higher in casts compared with the bulk soil in the pasture and the savannah, respectively. Overall means for cast MMD (8.3 and 7.4 mm) were twice as high as those in the bulk soil (3.8 mm) in the savannah and the pasture, respectively. However, MMD was not significantly different between the casts and the bulk soil in two occasions in the pasture. Available nitrogen (N_avail) in injected casts was initially greater than bulk soil levels, reaching maximum levels just after injection (116 and 93 mg kg⁻¹) and remained significantly greater during 1-2 weeks, in the savannah and the pasture, respectively. In conclusion, the tonnes of casts deposited in the soil profile by Martiodrilus sp. each year are likely to contribute greatly to plant nutrition and to the regulation of the soil structure. For each anecic earthworm species, the ecological impact of its below-ground casts is likely to be as important as its surface casts.     

Changes of pore morphology, infiltration and earthworm community in a loamy soil under different agricultural managements

Earthworm activity produces changes at different scales of soil porosity, including the mesoporosity (between 1.000 and 30 μm eq. dia.) where both water retention and near-saturated infiltration take place. At this scale, the structural changes are poorly described in temperate agricultural systems, so we do not yet fully understand how these changes occur. The present study was conducted to determine the relationships between the morphology of the mesopores, which is mainly affected by earthworm activity, and the hydrodynamic behaviour (near-saturated infiltration) of topsoil under different agricultural managements inducing a large range of earthworm populations.Investigations were carried out at the soil surface in three fields under different management practices giving rise to three different earthworm populations: a continuous maize field where pig slurry was applied, a rye-grass/maize rotation (3/1 year, respectively) also with pig slurry, and an old pasture sown with white clover and rye-grass.Pore space was quantified using a morphological approach and 2D image analysis. Undisturbed soil samples were impregnated with polyester resin containing fluorescent pigment. The images were taken under UV light, yielding a spatial resolution of 42 μm pixel⁻¹. Pores were classified according to their size (which is a function of their area) and their shape. Hydraulic conductivity K(h) was measured using a disc infiltrometer at four water potentials: −0.05, −0.2, −0.6, and −1.5 kPa. The abundance and ecological categories groups of earthworms were also investigated.Continuous soil tillage causes a decrease in both abundance and functional diversity (cf. maize compared with old pasture) when soil tillage every 4 years causes only a decrease in abundance (cf. rotation compared with old pasture). There were no relationships between total porosity and effective porosity at h=−0.05 kPa. Image analysis was useful in distinguishing the functional difference between the three managements. Fewer roots and anecic earthworms resulted in fewer effective tubular voids under maize. There were fewer packing voids in the old pasture due to cattle trampling. Greater hydraulic conductivity in the pasture phase of rotation may arise from a greater functional diversity than in the maize and absence of cattle trampling compared with the pasture. We point to some significant differences between the three types of agricultural management.A better understanding is required of the influence of agricultural management systems on pore morphology. This study provides a new methodology in which we consider the earthworm activity as well as community in order to assess the effects of agricultural management on soil structure and water movement.     

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

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

Abbau von Isoproturon in Regenwurm‐Makroporen und in der Unterbodenmatrix – Eine Feldstudie

Degradation of isoproturon in earthworm macropores and subsoil matrix—a field study The objective is to compare the time scale of microbial degradation of the herbicide Isoproturon at the end of earthworm burrows with the time scale of microbial degradation in the surrounding soil matrix. To this end, we developed a method which allows the observation of microbial degradation on Isoproturon in macropores under field conditions. Study area was the well‐investigated Weiherbach catchment (Kraichgau, SW Germany). The topsoil of a 12 m² large plot parcel was removed, the parcel was covered with a tent and instrumented with TDR and temperature sensors at two depths. After preliminary investigations to optimize application and sampling techniques, the bottom of 55 earthworm burrows, located at a depth of 80–100 cm, was inoculated with Isoproturon. Within an interval of 8 d, soil material from the bottom of 5–6 earthworm burrows was taken into the laboratory and analyzed for the Isoproturon concentration for investigation of the degradation kinetics. Furthermore, the degradation of Isoproturon in the soil matrix, that surrounded the macropores at the field plot, was observed in the laboratory. Microbial degradation of Isoproturon at the bottom of the earthworm burrows was with a DT‐50‐value of 15.6 d almost as fast as in the topsoil. In the soil matrix that closely surrounded the center of the earthworm burrows, no measurable degradation was observed within 30 d. The clearly slower degradation in the soil matrix may be likely explained by a lower microbial activity that was observed in the surrounding soil matrix. The results give evidence that deterministic modeling of the fate of pesticides once transported into heterogeneous subsoils by preferential flow requires an accuracy of a few centimeters in terms of predicting spatial locations: time scales of microbial degradation in the subsoil drop almost one order of magnitude, in case the herbicides dislocates from the bottom of an earthworm burrow a few centimeter into the surrounding soil matrix. If at all, predictions of such an accuracy can only be achieved at locations at sites where the soil hydraulic properties and the macropore system are known at a very high spatial resolution.     

Differences in burrowing behaviour and spatial interaction between the two earthworm species Aporrectodea nocturna and Allolobophora chlorotica

 To study intraspecific and interspecific interactions between different ecological types of earthworm, the burrowing behaviour of two earthworm species (the anecic earthworm Aporrectodea nocturna and the endogeic earthworm Allolobophora chlorotica) was observed in a microcosm. Earthworms were either alone in the microcosm, together with a conspecific earthworm, or with an earthworm of the other species. Observations under red light, including those of the position of the animals and the burrow dug, were recorded 4 times a day for 8 days and provided the data needed to reconstruct the probable trajectories of each earthworm. Differences in movement and burrowing behaviour were observed. Comparisons between the two species confirmed the expected behaviour of each ecological type: A. nocturna reused its burrow system regularly, whereas A. chlorotica rarely did. Moreover, it was shown that A. chlorotica burrowed less and explored a smaller surface when in the presence of A. nocturna. Besides, A. nocturna burrowed less and explored a smaller surface when in the presence of another A. nocturna. If these interactions occur under natural conditions, they could affect the structure of the burrow systems of the earthworm species examined. Received: 15 January 1999