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.     

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     

Characterisation of the three-dimensional structure of earthworm burrow systems using image analysis and mathematical morphology

The aim of this work was to exemplify the specific contribution of both two- and three-dimensional (3D) X-ray computed tomography to characterise earthworm burrow systems. To achieve this purpose we used 3D mathematical morphology operators to characterise burrow systems resulting from the activity of an anecic ( Aporrectodea nocturna), and an endogeic species ( Allolobophora chlorotica), when both species were introduced either separately or together into artificial soil cores. Images of these soil cores were obtained using a medical X-ray tomography scanner. Three-dimensional reconstructions of burrow systems were obtained using a specifically developed segmentation algorithm. To study the differences between burrow systems, a set of classical tools of mathematical morphology (granulometries) were used. So-called granulometries based on different structuring elements clearly separated the different burrow systems. They enabled us to show that burrows made by the anecic species were fatter, longer, more vertical, more continuous but less sinuous than burrows of the endogeic species. The granulometry transform of the soil matrix showed that burrows made by A. nocturna were more evenly distributed than those of A. chlorotica. Although a good discrimination was possible when only one species was introduced into the soil cores, it was not possible to separate burrows of the two species from each other in cases where species were introduced into the same soil core. This limitation, partly due to the insufficient spatial resolution of the medical scanner, precluded the use of the morphological operators to study putative interactions between the two species     

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.     

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.     

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.     

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.     

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.     

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.     

Earthworm excreta (mucus and urine) affect the distribution of springtails in forest soils

The collembolan species Heteromurus nitidus, exclusively located in soils at pH>5, can be cultured in acidic humus. As this species is attracted to the excreta of earthworms from calcic mull, its distribution is supposed to be determined only indirectly by soil pH through the distribution of earthworms. Higher densities and biomasses of Lumbricidae were observed in a calcic mull (pH 7.8) than in an acidic mull (pH 4.8) and a moder humus (pH 4.2). Choice experiments were performed to compare the attraction of H. nitidus to the mucus-urine mix of five earthworm species from the calcic mull and the acidic mull. H. nitidus was attracted to the excreta of the five species, whatever their ecological category and the humus form from which they originated. The collembolan Heteromurus major, which was indifferent to soil pH, was not attracted to earthworm excreta, which emphasizes the significance of this phenomenon for the distribution of H. nitidus over a pH range. The attraction of H. nitidus to earthworm excreta tended to be weaker and more variable when earthworms originated from acidic mull compared to calcic mull, particularly in the case of Lumbricus terrestris. Increased earthworm density reinforced by better mucus quality and quantity could determine the distribution of H. nitidus according to soil acidity. The only urine compound capable of attracting H. nitidus was NH₃ at a low concentration (0.03 g l^-1). The NH₃ content of the mucus-urine mix which attracted H. nitidus was 0.037 g l^-1, and was therefore responsible, at least partly, for the attraction.     

Evolution of burrow systems after the accidental introduction of a new earthworm species into a Swiss pre-alpine meadow

 The unintentional introduction of a new earthworm species (Aporrectodea nocturna) into a Swiss pre-alpine meadow resulted in a great increase in earthworm density in the newly colonized area (386 m^–2) compared with the density observed in the natural area (273 m^–2) where an earthworm community was already present. To investigate the impact of this introduction on the burrow systems, eight soil cores (length 25 cm, diameter 16 cm) were taken (four in the colonized area and four in the natural area) and analysed with computer-assisted X-ray tomography. The resulting images were processed to obtain the 3D-skeleton reconstructions of the earthworm burrow systems. Due to high variability in these burrow systems, only slight differences were observed between the two areas. The total burrow length and the mean burrow lengths tended to be greater in the colonized area. Moreover, the distribution of pore numbers with depth showed different patterns with a maximum for depths between 10 cm and 15 cm in the colonized area and a maximum for depths between 20 cm and 25 cm in the natural area. These differences may have been related to: (1) the particular behaviour of A. nocturna, which was observed to cast at the surface in this site, and (2) the predominance of juvenile earthworms around the colonization front. These differences were sufficient to create significant effects on the continuity of the burrow systems (assessed by the number of different pathways between virtual horizontal planes) for the two areas. The colonized area was characterized by a greater pore continuity, which could have resulted in enhanced transfer properties. Received: 2 July 1999     

Short rotation forestry – Earthworm interactions: A field based mesocosm experiment

Short rotation forestry (SRF) which consists of planting rapidly growing native and non-native tree species has been introduced to the UK to increase woody biomass production. A largely unknown aspect of SRF species is their interaction with soil fauna, of which the earthworm community is a major component. Earthworms have a pronounced impact on litter decomposition, nutrient cycling and tree growth. Conversely, tree litter and root chemistry can impact on the associated earthworm community development. The aim of this study was to determine direct interactions between SRF species and earthworms. A field-based mesocosm experiment was conducted using Betula pendula (birch) and Eucalyptus nitens (eucalyptus) with two earthworm species Lumbricus terrestris and Allolobophora chlorotica. The one year experiment revealed that native birch and non-native eucalyptus had a similar influence on L. terrestris population development. However, birch had a positive impact on A. chlorotica population establishment compared with eucalyptus. In the presence of earthworms, total tree biomass and leaf nitrogen concentration of eucalyptus were increased respectively by 25% and 27% compared with an earthworm-free control. In the presence of earthworms, surface litter incorporation was greater for both tree species (almost 5 times for birch and 3 times for eucalyptus) compared with controls. This work showed direct SRF-earthworm interactions which differed for tree species.     

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.     

Dehydration does not affect the radial pressures produced by the earthworm <Emphasis Type="Italic">Aporrectodea caliginosa</Emphasis>

As a soil dries, the earthworms in that soil dehydrate and become less active. Moisture stress may weaken an earthworm, lowering the radial pressure that the animal can produce. This possibility was investigated for the earthworm Aporrectodea caliginosa (Savigny). Pressures were compared for saturated earthworms (worms taken from saturated soil) and stressed earthworms (worms that had been partially dehydrated by leaving them in dry soil). A load cell was used to record the forces that earthworms produced as they moved through artificial burrows (holes that had been drilled through blocks of aluminium or Perspex). The radial pressure was calculated using the forces exerted and the dimensions of the artificial burrows. There was a negative correlation between burrow diameter and radial pressure, although radial pressure was independent of the length of the block through which the earthworms had burrowed. The highest radial pressures were produced by the anterior segments of the animal. Partial dehydration caused the earthworms to become quiescent, but did not decrease the radial pressure that the earthworms produced. It is suggested that coelomic fluid is retained in the anterior segments while the rest of the animal dehydrates. Dehydrated earthworms became lethargic, and we suggest that lethargy is due to the loss of coelomic fluid from the posterior segments. Coelomic fluid is known to be lost through dorsal pores. In burrowing species of earthworm such as Aporrectodea caliginosa, these pores are only present on the posterior segments.     

Earthworms influenced by reduced tillage,conventional tillage and energy forest in Swedish agricultural field experiments

Abstract

We compared earthworm density, depth distribution and species composition in three soil cultivation experiments including the treatments ploughless tillage and mouldboard ploughing. Sampling was done in September 2005 and for one experiment also in 1994. By yearly sampling 1995–2005, earthworms in an energy forest of Salix viminalis were compared with those in an adjacent arable field. Sampling method was digging of soil blocks and hand sorting and formalin sampling in one cultivation experiment. Both methods were used in the energy forest and arable land comparison.

In two soil cultivation experiments, highest abundances or biomass were found in ploughless tillage. Earthworm density was higher in the upper 10 cm, especially in the ploughless tillage. Earthworm density was significantly higher in the energy forest than in the arable field. Formalin sampling revealed c. 36% of the earthworm numbers found by digging in the energy forest and gave almost no earthworms in the arable field. In all treatments with soil cultivation, species living and feeding in the rhizosphere and soil dominated. One such species, Allolobophora chlorotica, was more abundant under mouldboard ploughing than ploughless tillage. Lumbricus terrestris, browsing on the surface and producing deep vertical burrows, was more common in the ploughless tillage. Species living and feeding close to the soil surface were almost only found in the energy forest, which had not been soil cultivated since 1984.

The findings support earlier studies pointing out possibilities to encourage earthworms by reduced soil cultivation. This is one of the first published studies that followed earthworm populations in an energy forest plantation during several years. Explanation of earthworm reactions to management and environmental impacts should be done with consideration of the ecology of species or species groups. Earthworm sampling by formalin must always be interpreted with caution and calibrated by digging and hand-sorting sampling.     

Flooding responses of three earthworm species,Allolobophora chlorotica,Aporrectodea caliginosa and Lumbricus rubellus,in a laboratory-controlled environment

To get a better understanding of earthworm’ responses towards flooding, three laboratory experiments were performed with the species Allolobophora chlorotica, Aporrectodea caliginosa and Lumbricus rubellus.Flooding response was determined in a pot experiment, in which the earthworms were incubated for 42 days in flooded or non-flooded soil, with or without heavy metal pollution. To determine moisture preference, earthworms were incubated for 9 days in aquaria with five compartments, containing soil with a moisture gradient (%, w/w), ranging from 35%, 45% (field capacity), 55%, 65% (saturated) to 65%+ (saturated and an extra water layer). Effects on earthworm health were studied by incubating earthworms of each species for 42 days in soil with the same range of moisture contents and determining the dry/wet weight ratio and dry weight gain as an indication of earthworm health.A. chlorotica was tolerant to water, although the worms tended to escape from flooded soil. Their health was significantly lower in the flooded soils (P<0.05). A. caliginosa showed little response to flooding. This species, however, was affected by the heavy metal pollution in the pot experiment. L. rubellus was sensitive towards flooding, with escape and avoidance behaviour being the main mechanism of survival. This species was able to survive when forced to stay in flooded soil for at least 42 days, but this significantly reduced its health (P<0.05).These results suggest that earthworms are able to survive in inundated soils, but there are large differences between species in response to flooding conditions.     

The interactions between soil type and earthworm species determine the properties of earthworm casts

Earthworms are recognized to increase soil porosity, reorganize soil structure, and stimulate soil microflora and nutrient mineralization. The properties of earthworm casts should depend both on earthworm species or ecological group and on soil properties. Interactions between earthworm species and soil types have been suggested, but only poorly demonstrated. In order to better understand those interactions, two hypotheses led our study: (1) Soil type has a greater influence on cast properties than earthworm; (2) Earthworms from different species influence cast properties differently; (3) The intensity and direction of the impact of each earthworm species on cast properties vary with soil properties. Fifteen physical and chemical variables (N–NH₄⁺, N–NO₃⁻, total organic C and N, C/N ratio, CaCO₃, pH, P, K⁺, Mg²⁺, Mn²⁺, Na⁺, CEC, moisture, wettability) were measured in casts of three earthworm species (Lumbricus terrestris, Allolobophora chlorotica and Aporrectodea rosea) produced in three temperate soils. Univariate and multivariate analyses showed that earthworm species and soil types significantly impacted cast properties. pH, N_t, K and Mg contents were interactively altered by both factors. Multivariate analysis showed that a difference of soil type had a major impact on casts properties (62%) compared to the impact of a difference of earthworm species (10%). Cast properties were most impacted by L. terrestris, then by A. chlorotica and last by A. rosea. The response ratio (ratio of the properties of the casts to the properties of the bulk soil) was used to quantify the effect of earthworm species compared to the control soil. It showed a higher response of variables in casts in nutrient-rich soils, especially in casts of L. terrestris. The interactions between earthworm species and soil types on cast properties were discussed with regards to earthworm ecology, properties of the soil, and earthworm modifications of cast microflora.