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.     

Earthworms and collembola relationships: effects of predatory centipedes and humus forms

Relationships between anecic earthworms (Lumbricus terrestris and Aporrectodea giardi) and the collembolan species Heteromurus nitidus (Templeton, 1835), which is known to be attracted to earthworms, were investigated in an 8-week laboratory experiment. Our aims were (1) to assess whether earthworms influence the population dynamics of H. nitidus, and (2) to study pathways of influence and how earthworm effects are modified by humus forms and predators. Using microcosms with three defaunated humus forms, then provided with earthworms and predators, we intended to demonstrate that, amongst possible favourable effects of earthworms on springtail populations, earthworm activity may provide greater access and more pathways for springtails to explore soil and avoid predation. We expected that the effects of predators (centipedes) on the abundance of H. nitidus would increase from less (calcic mull) to more (moder) compact soil, and we hypothesized that earthworms would reduce predation pressure on H. nitidus by providing escape routes through increased macroporosity. Humus forms and earthworms only affected the population size of H. nitidus under high predation pressure, when collembolan numbers were higher in calcic mull than in moder, and were increased by the presence of earthworms. These results corroborate the hypothesis that earthworms, by increasing soil macroporosity, improve the escape routes for Collembola and thus evade predation. In moder humus earthworms increased the density of H. nitidus whether predators were present or not, so we cannot exclude that earthworms were also directly beneficial to H. nitidus. However, the hypothesis of a functional relationship mediated by soil macroporosity seems relevant since it was supported by differences observed when considering body size. When two size classes were distinguished within populations of H. nitidus (1) the positive effect of earthworms in moder was observed only on larger Collembola (>1 mm), (2) the density of the larger Collembola was decreased by predation only in moder and not in mull, (3) the effects of predators on the smaller individuals were not influenced by the presence of earthworms whatever the humus form, and was not decreased by the presence of earthworms. Nevertheless, factors other than macroporosity may operate as the presence of earthworms in acidic mull led to an unexplained decrease in the abundance of small-sized H. nitidus.     

The impact of earthworms on the abundance of Collembola: improvement of food resources or of habitat?

I assessed the direct influence of earthworm excretions, and the impact of earthworms through their action on the soil structure (increased macroporosity), on the population dynamics of the collembolan species Heteromurus nitidus. The intestinal content of Collembola arising from cultures on different soil types was observed, and two experimental cultures of H. nitidus were run: (1) a culture performed on an inert substrate supplied either with earthworm casts or with soil as food resource, (2) an experiment using microcosms with cores of two humus forms (moder and calcic mull), in the presence or absence of earthworms. The observation of gut contents revealed that H. nitidus feeds on excrements, the composition of which (ratio organic matter/mineral matter) varies according to the humus form where it lived. Slightly aged (10–15 days) organo-mineral casts of earthworms appeared to be a better food than calcic mull aggregates or organic material from moder. Densities of H. nitidus cultured in cores of calcic mull were higher than in moder, except when cores of moder were inhabited by an anecic earthworm for 2 months. The humus form strongly influenced populations of H. nitidus, firstly because densities of predators were higher in moder than in calcic mull, and probably also because of soil macroporosity. It was concluded that earthworms would affect predation on H. nitidus by creating a network of interconnected macropores in which Collembola can move and find shelter.     

Stimulating effect of earthworm excreta on the mineralization of nitrogen compounds in soil

The effect of excreta of earthworm species Aporrectodea caliginosa and Eisenia fetida on the mineralization of nitrogen compounds in soils has been studied. A single application of excreta obtained from three earthworms in one day increased the formation of nitrate nitrogen compounds in the soil by 10–50%. The application of ammonium nitrogen (in the form of NH₄Cl) in amounts equivalent to the ammonium nitrogen content in the daily excreta of three earthworms had the same effect on the mineralization of nitrogen compounds. The effect of earthworm excreta, as well as the effect of ammonium nitrogen, on the nitrification process was an order of magnitude higher than their contribution to the formation of nitrates due to the oxidation of the introduced ammonium. Hence, ammonium—an important component of the earthworm excreta—can exert a stimulating effect on nitrification processes in the soil and produce long-term cumulative effects that are much more significant than the direct effect of this nitrogen compound.     

Mucus excretion and carbon turnover of endogeic earthworms

Summary Mucus excretion of endogeic earthworms, by the body surface to burrow walls and by the intestine to casts, was investigated using uniformly ¹⁴C-labelled adult or subadult specimens of Octolasion lacteum (Örley) in laboratory incubations in soil from a beechwood on limestone. The daily loss of C due to mucus excretion from the body surface and in casts was calculated as 0.2 and 0.5% of total animal C, respectively. The C loss due to mucus excretion by subadult or adult individuals of O. lacteum is assumed to account for 63% of total C losses (including mucus excretion and respiration) of the earthworms. In a second experiment we studied the incorporation of ¹⁴C from labelled soil, again from a beechwood on limestone, into the tissue of the endogeic earthworm species Aporrectodea caliginosa (Savigny). The results of this experiment indicate the existence of two C pools, one more labile and one more stable, in earthworms. It is assumed that the C investment for respiration and mucus excretion is derived from the labile C pool of endogeic earthworms.     

Fat but slim: Criteria of seed attractiveness for earthworms

Earthworms were shown to significantly affect seeds and seedlings survival via their ingestion and digestion for nutritive purposes. Such selective feeding of earthworms on plant seeds is likely to favour certain plant species and to affect seed bank composition, plant recruitment and plant community structure. Relationships between earthworms and seeds, particularly seed traits that determine attractiveness of seeds for earthworms, are yet to be determined. In this study, the influence of six seed traits was tested on the ingestion, digestion and germination of seeds by two earthworm species (Lumbricus terrestris, anecic and Satchellius mammalis, epigeic). The seed traits tested were their length, width, weight, shape, oil content and the presence of trichomes on their surface. Each earthworm species was introduced into a microcosm with eleven seed species from a chalk grassland that represented those different traits. Ingested, digested and germinated seeds were counted after voiding the guts of the earthworms. Univariate and multivariate analyses showed that seed length, width, weight and seed oil content could significantly affect the ingestion of seeds for both earthworm species. Seed width and seed oil content were the two traits that influenced the digestion of seeds the most, but only for L. terrestris. We also found that seed ingestion was earthworm species-specific but we found no correlation between earthworm traits and number of ingested or digested seeds. Few seeds germinated from L. terrestris casts and no seeds germinated from S. mammalis casts. Implications in terms of plant evolution strategies are further discussed.     

Subsurface earthworm casts can be important soil microsites specifically influencing the growth of grassland plants

Earthworms (Annelida: Oligochaeta) deposit several tons per hectare of casts enriched in nutrients and/or arbuscular mycorrhizal fungi (AMF) and create a spatial and temporal soil heterogeneity that can play a role in structuring plant communities. However, while we begin to understand the role of surface casts, it is still unclear to what extent plants utilize subsurface casts. We conducted a greenhouse experiment using large mesocosms (volume 45 l) to test whether (1) soil microsites consisting of earthworm casts with or without AMF (four Glomus taxa) affect the biomass production of 11 grassland plant species comprising the three functional groups grasses, forbs, and legumes, (2) different ecological groups of earthworms (soil dwellers—Aporrectodea caliginosa vs. vertical burrowers—Lumbricus terrestris) alter potential influences of soil microsites (i.e., four earthworms × two subsurface microsites × two AMF treatments). Soil microsites were artificially inserted in a 25-cm depth, and afterwards, plant species were sown in a regular pattern; the experiment ran for 6 months. Our results show that minute amounts of subsurface casts (0.89 g kg^?1 soil) decreased the shoot and root production of forbs and legumes, but not that of grasses. The presence of earthworms reduced root biomass of grasses only. Our data also suggest that subsurface casts provide microsites from which root AMF colonization can start. Ecological groups of earthworms did not differ in their effects on plant production or AMF distribution. Taken together, these findings suggest that subsurface earthworm casts might play a role in structuring plant communities by specifically affecting the growth of certain functional groups of plants.     

Effects of biomass and size of Millsonia anomala (Oligochaeta: Acanthodrilidae) on particle aggregation in a tropical soil in the presence of Panicum maximum (Gramineae)

Summary The present study assessed the effect of the tropical geophagous earthworm, Millsonia anomala, on the aggregate-size distribution of a sieved (2 mm), tropical ferruginous soil in the presence and absence of the perennial tropical grass Panicum maximum. The effect of two size groupings and graded biomass densities of M. anomala on soil aggregation was analysed in time and with depth in the containers within which the plants were grown. In the absence of earthworms, aggregation was rapid although limited (13.5% of soil as aggregates >2.0 mm), and probably arose from a combination of microbial activity and physical processes (interparticle bonding due to clays and other colloids). The roots of the test species contributed little to aggregation. In contrast, the effect of earthworm activity on soil aggregation was rapid and important. The effects of both biomass and, to a lesser extent, size were significant. After 79 days, aggregation reached a maximum with 3 g per container of small earthworms (ca. 59% of soil as aggregates >2.0 mm) and a minimum with 1 g per container of large earthworms (ca. 35% of aggregates >2.0 mm). Aggregation was considered to occur through three different mechanisms: (1) A rapid aggregation due to the interactions of colloids; (2) an intermediate aggregation due to a combination of unquantified processes related to earthworm activity (mucus secretion, development of fungus hyphae); (3) egestion of soil as earthworm casts which are stable macroaggregates.     

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 Collembola community of a Central European forest: Influence of tree species composition

The present study investigates the response of the Collembola community to replacement of beech by spruce or by mixed stands of beech and spruce in the Solling mountains (Germany). The study was carried out in three beech (Fagus sylvatica), spruce (Picea abies) and mixed stands of beech and spruce arranged in three blocks. The density, diversity and community structure of Collembola as well as microbial and abiotic parameters in the organic layers and mineral soil of the three spruce, three beech and three mixed stands were investigated. Major results are: (i) Collembola communities did not differ strongly between stand types and were dominated by Folsomia quadrioculata and Mesaphorura species, (ii) neither total abundance of Collembola nor densities of the hemiedaphic species F. quadrioculata, Parisotoma notabilis and Isotomiella minor significantly responded to stand type, (iii) in the mixed stands the fungal biomass was increased leading to high densities of fungal feeding Collembola (e.g. Mesaphorura sp.) and high species numbers of Collembola, (iv) the density of the epedaphic and partly herbivorous group Entomobryidae/Tomoceridae in the spruce stands exceeded that in the mixed and beech stands; presumably this was due to the higher diversity of the ground vegetation in the spruce stands. Canonical correspondence analysis (CCA) of the collembolan communities of L/F and H/Ah horizons also indicated that most of the epedaphic species were associated with the spruce stands. Moreover, results of the CCA indicated that soil pH is an important structuring force for collembolan communities. Overall, results suggest that stand type impact collembolan communities, presumably via changes in the amount and quality of food resources, such as fungal biomass and living plant material. However, differences in collembolan community structure between the investigated stand types were moderate supporting earlier findings that Collembola generally respond little to changes in the vegetation structure.     

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.     

Dynamic study of the burrowing behaviour of Aporrectodea nocturna and Allolobophora chlorotica: interactions between earthworms and spatial avoidance of burrows

The behaviour of earthworms belonging to two different species and ecological types (Aporrectodea nocturna and Allolobophora chlorotica) was studied using two-dimensional (2D) terraria. Two experiments were set up to gain insight into the nature of interactions between these earthworms. Firstly, the evolution of the burrow systems was analysed with the density of the earthworms varying from one to five individuals. Secondly, a burrow system was first established by using one earthworm which was then removed before the introduction of a second earthworm. This second earthworm therefore encountered a burrow system created either by a conspecific earthworm or by an earthworm of the other species. These experiments showed that: (1) intra- and interspecific interactions occur between earthworms, (2) these interactions are dependent on the physical presence of the earthworms, and (3) spatial avoidance can occur (A. chlorotica avoiding burrows created by A. nocturna). The results suggest that earthworm burrow systems are "individual structures", rarely used by other earthworms when inhabited. When abandoned, the burrows may be recolonised depending on the ecological type of the earthworm under consideration.     

Changes in nutrient pools,microbial biomass and microbial activity in soils after transit through the gut of three endogeic earthworm species of the genus Postandrilus Qui and Bouché, 1998

Purpose

Endogeic earthworms play a significant role in biogeochemical cycles due to the large amount of soil they ingest, and because after transit through their guts, casts usually show differences in nutrient contents and microbial populations with bulk soil. Here, we studied how three endogeic earthworm species, Postandrilus majorcanus, Postandrilus sapkarevi and Postandrilus palmensis, inhabiting soils in Majorca island (Balearic Islands, W Mediterranean), modify nutrient pools and microbial communities of soil.

Materials and methods

To do this, we analysed C, N and P pools, microbial biomass (phospholipid fatty acids, PLFA) and microbial activity (fluorescein diacetate hydrolysis, FDA) in paired samples of bulk soil and fresh casts.

Results and discussion

The mineral and organic N contents were generally enhanced in casts produced by all three earthworm species. However, inorganic P and organic C contents were only higher in P. sapkarevi (32 %, only P) and P. majorcanus casts (100 % for both soil nutrient pools) than in bulk soil. Bacterial and fungal biomass were only higher than in bulk soil in P. majorcanus casts (65 and 100 %, respectively), but without effects on microbial activity, that was lower in P. palmensis casts (26 %). Earthworm gut transit strongly influenced the soil microbial community structure, resulting in differences between casts and soils.

Conclusions

The increased nutrient mineralization (6-, 1.3- and 1.4-fold for N, C and P, respectively) in casts produced by these earthworm species is of particular importance because of the amount of casts released and the seasonal variations in earthworm activity, which may favour plant growth.     

Understory vegetation,soil structure and soil invertebrates in Congolese eucalypt plantations,with special reference to the invasive plant Chromolaena odorata and earthworm populations

Earthworm relationships with vegetation have received extensive attention, and earthworm density has been shown to be related to vegetation types or plant species. However, the factors involved are rarely known. In Congo, we studied the effect of Chromolaena odorata (L) R.M. King & H. Robinson, which invades eucalypt plantations, on soil invertebrates, especially earthworms. In order to investigate relationships between vegetation cover and soil invertebrates, four understory species, including C. odorata, were studied. Also, comparisons were made between plots invaded by C. odorata and plots free from it. The addition of leaf litter on experimental plots was made in order to check its influence. Plant remains were observed in the digestive tract of earthworms. An increased earthworm density was observed under C. odorata. The leaf litter and roots of this species had low lignin/N ratio. The size of leaf fragments found in the digestive tract of the earthworms, and the lack of short-term effect of experimentally added leaf litter, suggested that litter quality could influence earthworm through their feeding on fine particulate top soil organic matter. The amount of soil aggregates, in the size classes that fitted the size range of earthworm casts, was increased under C. odorata. More field experiments are needed to establish a causal effect in the relationships observed between earthworm density and C. odorata. If so, the major drawbacks, such as water and nutrient competition, resulting from C. odorata overrunning the plantations, could be somewhat offset by its positive effects through soil improvement.     

Earthworm (Lumbricidae) diversity in the Central Balkans: An evaluation of their conservation status

In this paper we summarize the current knowledge on earthworm diversity in the central part of the Balkans, in the State of Serbia. Up to now, 71 species and 8 subspecies, belonging to 18 genera, are known from investigated territory. Our aim was to produce a single priority list for earthworm conservation, particularly as the Balkan countries did not have their own Red Lists for earthworms and none of the species had a legally protected status. The list underlines the diversity of earthworms, zoogeographical position as well as category of threat. The analysis based on the IUCN (2001) Red List Categories shows that 17 of 79 taxa are Critically Endangered (5 species are serious Critically Endangered, but 12 species are only suspect for the Critically Endangered category), 9 species are Endangered and 14 species are Vulnerable. Serious candidates for their inclusion in CR category within the IUCN (2001) Red List are: Cernosvitovia biserialis, Dendrobaena kozuvensis, Lumbricus improvisus, Serbiona kosowensis montenegrina and Serbiona serbica. Of all registered earthworms, 34 taxa (43.1%) were identified only as endemic species. Unfortunately, most of the high-priority species are endemic (23 taxa). The position of the Central Balkans contributes to the great biodiversity of earthworms on its territory, but the fact that nearly 36% of its lumbricids are threatened is a strong signal that action is required.     

The dispersal of the earthworm Aporrectodea giardi responds faster to habitat quality than to cumulative use of habitat in experimental conditions

The study of the spatial distribution of soil organisms is one of the key research areas for understanding soil functioning. However, we still know little about the role of dispersal in the distribution of soil organisms such as earthworms. Critically, the relative strength of the external factors that trigger dispersal movements has not been documented. In this work, we test the relative importance of habitat quality and the cumulative use of habitat as drivers of dispersal of an anecic earthworm (Aporrectodea giardi) by observing their dispersal rates over 1, 2, 4, 8, 16 and 32 days. The results suggest that cumulative dispersal rates were higher and reached a maximum value more rapidly when individuals were introduced into unsuitable rather than suitable soil. This suggests that earthworm dispersal responded more rapidly and this response was more pronounced with respect to the cumulative use of the habitat. It seemed that there were two types of dispersal: one triggered quickly to escape unsuitable conditions and another in response to a cumulative use of the habitat.     

Effects of earthworms on soil properties and rice production in the rainfed paddy fields of Northeast Thailand

Earthworms play an important role in soil fertility because of their effects on soil properties, and especially the creation of casts that usually have different properties than the surrounding soil. If the influence of earthworms on soil properties and plant growth has been extensively studied in aerobic ecosystems, their effects remain largely unknown in partially flooded environments such as paddy fields. The aim of this study was to describe the potential influence of earthworms on soil properties and rice yield in Northeast Thailand. We studied the abundance of aboveground casts produced by Drawida beddardi in two paddy rice systems (transplanting: TP and direct seeding: DS) and we measured the soil physicochemical properties of casts and their influence on rice growth and yield. Cast abundance was low (∼250 casts per ha) and not different in TP and DS. Casts appeared as patches of fertility with more fine particle sizes and organic matter contents, a higher CEC, as well as a much higher concentrations of mineral elements (N, P and K). As a consequence, our results suggest that earthworm activity lead to a significant higher rice yield. These results stress the importance earthworms could have in the regulation of rice productivity in paddy fields. More research is now needed to understand the ecological mechanisms that regulate earthworm abundance and activity in these environments.     

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.     

Liming upland grassland: the effects on earthworm communities and the chemical characteristics of carbon in casts

Different earthworm species have different tolerances of acid soil conditions, and the application of lime to upland grassland to improve the grazing quality may therefore alter the size and diversity of the earthworm community. Altering soil properties may also affect the chemical characteristics of organic C in earthworm casts. We surveyed the earthworm community of an upland grassland in southern Scotland at the outset of annual lime applications, and after 3 years, and used ¹³C nuclear magnetic resonance (NMR) spectroscopy to assess the distribution of C between different functional groups in the organic matter. In addition, soil was incubated for 8 weeks with several earthworm species in the presence or absence of lime, and the earthworm casts were subsequently analysed by ¹³C NMR spectroscopy. Liming did not significantly affect earthworm abundance or species diversity, but it did affect the chemical composition of the casts. Casts from earthworms incubated in unlimed soil had greater ratios of alkyl‐C to O?alkyl‐C, indicative of more decomposed, recalcitrant C, and spectra from litter‐feeding species had the greatest intensities of O?alkyl‐C signals. In limed soil, the largest O?alkyl‐C signal intensities were not restricted to litter‐feeding species, indicating an increase in the quality of organic matter ingested by geophagous species.     

Destabilization of soil during the production of earthworm (Lumbricidae) and artificial casts

The mechanisms by which soil is destabilized in the digestive tract of endogéic earthworms were investigated with artificial casts, which were moulded with a syringe from slurries of a silty loam with or without gypsum and organic matter treatments, and compared to casts produced by Aporrectodea rosea (Lumbricidae). Both types of casts generally had the same levels of mechanical dispersion, observations of slaking, and particle size distribution when the casts were fresh, aged-moist for 30 days, or air-dried. Fresh casts were significantly more dispersive than the uningested soil despite the addition of gypsum or organic matter to the soil. However, the dispersion from aged-moist or air-dried casts was not substantially greater than that of uningested soil. Air-drying was more effective than moist-ageing in increasing the stability of casts and uningested soil. The concentration of soluble carbohydrate was greater in artificial casts produced from soil treated with sheep dung or xanthan gum, and in earthworm casts produced from soil treated with xanthan, than in the uningested soil of the same treatments. An increase in the concentration of soluble carbohydrate was related to an increase in dispersion. An attempt was made to simulate the addition of mucus to soil in the digestive tract of earthworms, by the addition of sucrose or xanthan gum to the slurry during the production of artificial casts. The addition of xanthan, but not sucrose, to the slurry increased mechanical dispersion relative to that of the uningested soil in the fresh treatment. Although the production of artificial casts destabilized soil to the same degree as earthworm casts, the artificial casts did not simulate all chemical, biochemical, and microbiological aspects of digestion. Received: 24 November 1995