Influence of clay content and acidity of soil on development of the earthworm <Emphasis Type="Italic">Lumbricus rubellus</Emphasis> and its population level consequences

In this paper we report on the influence of clay content and acidity of soil on growth and reproduction of the epigeic earthworm species Lumbricus rubellus (Hoffm.), which is common in most temperate soils and abundant in grasslands. Growth, cocoon production and survival of L. rubellus were tested in 12 Dutch soils which differed in soil properties. A matrix model was used to assess the population-level consequences of changes in growth and reproduction. Soil acidity had a strong negative effect on earthworm survival, and the maturation weight decreased with clay content. Individual weight gain in L. rubellus decreased with both acidity and clay content. The acidity of soils had a larger influence on population growth rate than the clay content. The acidity of the soil also changed the population composition towards younger age classes, whereas in soils rich in clay, the population composition did not change. The average individual weight of L. rubellus in clayey soils, however, was lower compared with that in soils low in clay, a result that agrees with literature data.     

Nitrogen transformations in copper-contaminated soils and effects of lime and compost application on soil resiliency

 A neutral and an acidic soil were treated with different doses (0–3,000 mg Cu kg^–1 soil as CuSO₄) of copper. The percentages of inhibition of nitrification in both soils varied from 5 to 97%, but for the N mineralization these percentages varied from 8 to 65%. The toxic effect of Cu for basal nitrification and N mineralization was assessed as critical. Nitrification was more sensitive than ammonification to copper toxicity. It appears that an ecological dose of inhibition for nitrification and N mineralization higher than 10% is suitable as an indicator for Cu contamination. Soil resiliency assessed by N mineralization in the lime treatments varied from 11 to 154% in the sandy soil and from 70 to 168% in the sandy loam soil. A combined application of lime and compost significantly increased soil resiliency. The percentage increase varied from 904 to 1,390% in the sandy soil and from 767 to 2,230% in the sandy loam soil. It appears that compost was a powerful agent for recovering the soil fertility of Cu-contaminated soils as assessed by N transformation. The acidic sandy soil showed a lower capacity for recovery after Cu toxicity stress. Received: 27 February 1999     

Effect of earthworms on decomposition processes in raw humus forest soil: A microcosm study

Summary The earthworms Lumbricus rubellus (Hoffmeister) and Dendrobaena octaedra (Savigny) were studied in the laboratory to determine their effects on decomposition and nutrient cycling in coniferous forest soil. CO₂ evolution was monitored, and pH, PO ₄ ^3– –P, NH ₄ ⁺ –N, NO ₃ ^– –N, total N, and total C in the leaching waters were measured. After three destructive samplings, numbers of animals, mass loss, pH, and KCl-extractable nutrients were analysed.The earthworms clearly enhanced the mass loss of the substrate, especially that of litter. L. rubellus stimulated microbial respiration by 15–18%, whereas D. octaedra stimulated it only slightly. The worms significantly raised the pH of the leaching waters and the humus; L. rubellus raised the value by 0.2–0.6 pH units and D. octaedra by 0.1–0.4 units. Both worms increased N mineralization. Although the biomass of both worms decreased during the experiment, the N released from decomposing tissues did not explain the increase in N leached in the presence of earthworms. The worms influenced the level of PO ₄ ^3– –P only slightly.     

The effect of earthworms (Lumbricidae) on nitrogenase activity in soil

Summary Nitrogenase activity associated with earthworms, their faeces and activity in soil was measured by the acetylene reduction technique. A clear increase in nitrogenase activity was found in field-deposited casts of Aporrectodea caliginosa in comparison with surrounding soil, although potential nitrogenase activity was significantly higher in soil than in casts. Nitrogenase activity associated directly with earthworms (Lumbricus rubellus ) was detected, indicating the presence of active N₂-fixing bacteria on the body surface and/or in the gut. Laboratory experiments showed that nitrogenase activity in the casts of L. rubellus was higher than in unmodified soil, and that nitrogenase activity in soil was significantly increased by the burrowing and feeding activity of these worms. This paper discusses the possible causes of these earthworm effects on soil nitrogenase activity and some methodological problems of determining the nitrogenase activity.     

Effects of seasonal and diurnal temperature fluctuations on population dynamics of two epigeic earthworm species in forest soil

Temperature fluctuations are a fundamental entity of the soil environment in the temperate zone and show fast (diurnal) and slow (seasonal) dynamics. However, responses of soil ecosystem engineers, such as earthworms, to annual temperature dynamics are virtually unknown. We studied growth, mortality and cocoon production of epigeic earthworm species (Lumbricus rubellus and Dendrobaena octaedra) exposed to temperature fluctuations in root-free soil of a mid-European beech-oak forest. Both earthworm species (3 + 3 individuals of each species) were kept in microcosms containing soil stratified into L, F + H and A_h horizons. In the field, earthworm responses to smoothing of diurnal temperature fluctuations were studied, simulating possible global change. In the laboratory, earthworm responses to seasonal (±5 °C of the annual mean) and diurnal temperature fluctuations (±5 °C of the seasonal levels) were analyzed in a two-factorial design. Both experiments lasted 12 months to differentiate between seasonal and diurnal responses. In the third experiment overwintering success of both earthworm species was investigated by comparing effects of constant temperature regime (+2 °C), and daily or weekly temperature fluctuations (2 °C ± 5 °C).Temperature regime strongly affected population performance of the earthworms studied. In the field, smoothed temperature fluctuations beneficially affected population development of both earthworm species (higher biomass, faster maturity and reproduction, lower mortality). Consequently, density of both species increased faster at smoothed than at ambient temperature conditions. In the laboratory, responses of L. rubellus and D. octaedra to temperature treatments differed; however, in general, earthworms benefited from the absence of diurnal fluctuations. Total earthworm numbers were at a maximum at constant temperature and lowest in the treatment with both diurnal and seasonal temperature fluctuations. However, after one year L. rubellus tended to dominate irrespective of the temperature regime. In the overwintering experiment L. rubellus sensitively responded to even short-term winter frost and went extinct after one week of frost whereas D. octaedra much better tolerated frost conditions. Earthworms of both species which survived frosts were characterized by a significant body weight decrease during the period of frosts and fast recovery in spring suggesting a different pattern of individual resource expenditure as compared with constant +2 °C winter regime. Contrasting trends in the population dynamics of L. rubellus and D. octaedra during the frost-free period and during winter suggest that in the long-term temperature fluctuations contribute to the coexistence of decomposer species of similar trophic position in the forest litter. The results are discussed in context of consequences of climate change for the functioning of soil systems.     

Effects of metal pollution on soil macroinvertebrate burrow systems

A reduction in the numbers of macroinvertebrates present in soil may have a negative effect on soil structure, infiltration rates, and gas exchanges. Soil pollution by metal is known to have a detrimental effect on soil macrofauna. The aim of the present study was to evaluate (1) direct and indirect effects of soil pollution on soil macroinvertebrate bioturbation and (2) effects of the two macroinvertebrate communities found in a polluted and a nonpolluted area (one supposed sensitive, the other tolerant to metals) on burrow systems parameters. Macroinvertebrate porosity was studied using X-ray tomography. Three-dimensional reconstructions and characterisation of the burrow system were obtained using image analysis. Results showed that metal pollution principally affected the spatial distribution of macropores (more macropores were found near the soil surface) and the shape of the burrow system (branching rate was higher in the polluted soil), whereas soil macroinvertebrate composition principally affects burrow density parameters (the number of burrows was higher for the sensitive macroinvertebrate community).     

Influence of earthworms and leaf litter on edaphic variables in sewage-sludge-treated soil microcosms

Summary A series of 48 greenhouse soil microcosms were established and treated with combinations of sewage sludge, Acer saccharum leaves, and the earthworms Eisenia fetida and Lumbricus terrestris. The microcosms were incubated at constant moisture for 110 days. Samples were then taken and analysed for sludge depth, organic-matter content, and waterstable soil aggregates. Weights of surface leaves and weights and numbers of surviving earthworms were determined for each microcosm. L. terrestris significantly reduced sludge depth and the surface organic-matter content of microcosm soil and significantly increased percentages of 4-mm diameter water-stable aggregates. Leaf litter also significantly reduced sludge depth and increased 4-mm water-stable aggregates. E. fetida inhibited surface feeding by L. terrestris, reduced its 110-day survival rate, and inhibited the production of 4-mm water-stable aggregates in L. terrestris treatments. Numbers of E. fetida increased in L. terrestris treatments. Sludge depth, organic-matter content and water-stable aggregates were not significantly different from controls in E. fetida treatments.     

Influence of microarthropod abundance and climatic factors on weight loss and mineral nutrient contents of Eucalyptus leaf litter during decomposition

Summary Densities of the different taxa of microarthropods per gram of litter in litter bags varied widely from the rainy to the dry season. Collembola and Acarina constituted more than 86% of the total microarthropods, and occurred in significantly greater numbers in the coarse-mesh bags than the fine-mesh bags. There were no fauna in the litter of suspended bags. The average weight loss was greater in the coarse-mesh bags (50.4%) than in the fine-mesh bags (44.5%), and the suspended bags (7.4%). Similarly, the concentrations of N and Ca were greater in the litter of coarse-mesh bags compared to that of the fine-mesh bags. In contrast, the concentrations of P and K were comparatively lower in coarse-mesh bags. The mass loss of litter showed a negative linear correlation with the total Collembola and with litter temperature. The N concentrations in the litter in both the mesh bags showed negative correlations with the abundance of total Collembola, and with that of Lepidocyrtus sp. and Sminthuridae, and rainfall. The N concentration in the litter in the coarse-mesh bags was positively correlated with the total number of arthropods but, surprisingly, was negatively correlated with the total number of Acarina. The concentration of Ca showed negative correlations with rainfall and litter moisture only. The P concentration showed positive correlations with total Collembola, with Lepidocyrtus sp. and Sminthuridae in both the mesh bags, with rainfall in the fine-mesh bags, and with total microarthropods in the coarse-mesh bags.     

The earthworm population of a winter cereal field and its effects on soil and nitrogen turnover

The earthworm population in a winter cereal field in Ireland was studied over a 3-year-period and its effects on soil and N turnover were assessed. The mean annual population density was 346–471 individuals m^-2 and the mean biomass was 56.9–61.2 g m^-2. Twelve species were recorded, the most abundant being Allolobophora chlorotica followed by Aporrectodea caliginosa, and 242 mg at 5°C to 713 mg at 10°C in the case of juvenile Lumbricus terrestris. Gut contents (dry mass of soil) comprised 6.7–15.5% of the A. caliginosa live mass, and 9.7–14.7% of the Lumbricus terrestris mass. Annual soil egestion by the field population was estimated as 18–22 kg m^-2. Tissue production ranged from 81.7 to 218.5 g m^-2, while N turnover resulting from mortality was calculated as 1.5–3.9 g m^-2 depending on the year and the method of calculation. Earthworms were estimated to contribute an additional 3.4–4.1 g mineral N to the soil through excretion, mucus production, and soil ingestion. Independent estimates of N output via mucus and excretion derived from ¹⁵N laboratory studies with Lumbricus terrestris were 2.9–3.6 g m^-2 year^-1.     

Palatability of microfungi to soil arthropods in relation to the functioning of arbuscular mycorrhizae

We investigated the feeding preferences of six species of mites and collembolans for three fungi commonly associated with roots of Acer saccharum (Glomus macrocarpum, Alternaria alternata and Trichoderma harzianum), from a maple-forest soil in southern Ontario, Canada. Experiments were also conducted in vitro to determine animal feeding responses to (1) increasing quantities of hyphal biomass, (2) the presence of root vs. litter fungal substrates, and (3) hyphae of different widths of Glomus macrocarpum. The results indicate that arthropods prefer to graze in the litter region rather than in the deeper soil layers. Under ideal moisture/temperature conditions, animals are forced to the lower regions by interspecific interactions. They prefer to graze on hyphae of conidial fungi rather than on those of arbuscular mycorrhizal fungi. When arbuscular mycorrhizal fungal hyphae are grazed, there is a clear preference for the narrower hyphae, which are those further away from the root. The thicker hyphal segments, commonly found connecting absorptive hyphal fans to roots, were less preferred. These data are not consistent with the hypothesis that microarthropods are detrimental to arbuscular mycorrhizal associations, and suggest that Glomalean fungi may have evolved mechanisms to deter grazing by microarthropods.     

Influence of soil properties on the population and activity of geophagous earthworms after five years of bare fallow

Many soil properties influence earthworm populations and activity. To determine which properties are of significance, a broad collection of soils was investigated. Samples from these different soils were kept bare at one site in large plots (3 Mg soil per plot) to liminate crop and weather interference and to isolate the dominating mechanisms of earthworm effects. Earthworm density, biomass, and tunnelling activity were assessed after 5 years of bare fallow. All earthworm parameters varied strongly. Earthworms increased soil respiration by their tunnelling activity, and in turn increased microbial activity and propagated the loss of organic C. Earthworm abundance ranged from 12 to 274 m^-2 and was about 10 times greater than on cropped soils. The range in abundance was mainly caused by variations in the numbers of juveniles. The average soil moisture content was the only soil property among the many properties investigated that was consistently correlated with earthworm abundance and biomass. Even after 5 years of bare fallow with almost no addition of fresh plant biomass and with little water loss by plant transpiration, the earthworm population was controlled by water stress and not by food stress. We therefore conclude that high water consumption by productive crops may degrade the habitat for geophagous earthworms.     

Restoration of giant cardon cacti in barren desert soil amended with common compost and inoculated with Azospirillum brasilense

Barren desert soil that otherwise could not support perennial plant growth was amended with six levels of common agricultural compost. Seedlings of the giant cardon cactus, one of the primary plant species responsible for soil stabilization in the southern Sonoran Desert, were inoculated with the plant-growth-promoting bacterium Azospirillum brasilense Cd, planted, and grown for 18 months under nursery conditions typical for slow-growth cacti. Control plants were grown without compost amendment, without inoculation (negative control), or in fertile, rare “resource island” soil preferred by cardon seedlings (positive control). During the prolonged growth period, the decisive factor in seedling growth in barren soil was the addition of small amounts of common compost; 6 to 25% of the growth substrate volume gave the best growth response and, to a lesser extent, so did inoculation with A. brasilense Cd. Although the bacteria significantly affects plant growth when amended with “resource island” soil and added to barren soil, its effect on plant growth was far smaller than when compost alone was added. Compost added to barren soil significantly increased the dry weight parameters of the plant to almost similar levels obtained by the “resource island” soil; however, the compost amendment supports a more voluminous and greener plant with elevated pigment levels. This study shows that barren soil supplemented with compost can replace the rare “resource island” soil for cardon nurseries destined to abate soil erosion in the desert.     

Influence of farm yard manure,poultry manure and forest litter on copper solubility in soil and uptake by Phaseolus vulgaris

Long term and intensive use of copper‐based fungicides on coffee farms may contaminate soils with copper. The legacy of copper pollution may pose the risk of contaminating food crops cultivated on these soils. A randomized block design field experiment at Kilimanjaro, Tanzania was designed to investigate the effects of different application rates of cattle manure, poultry manure and forest litter on aqua regia, EDTA and CaCl₂ extractable copper in soils and copper uptake by bean plants grown on this long‐term copper‐contaminated soil (more than 50 yr of copper application). It was important to examine the potential of the organic amendments in mobilizing or immobilizing copper and assess the risks of contaminating bean crops at a farm, where the application of organic amendments was common practice. At harvest, rhizosphere soils were collected and analysed. The soils were found to have large concentrations of copper, greatly exceeding international standard levels. Poultry manure applied at 40 tons/ha significantly increased CaCl₂ extractable copper compared with the control treatment. Organic carbon, cation exchange capacity, EDTA extractable copper, aqua regia extractable copper and copper concentrations in bean seeds or leaves were not significantly changed by the organic matter treatments. The concentration of copper was significantly less in bean seeds than in bean leaves (P < 0.01). The bean plants did not take up excessive quantities of copper, and therefore, the risk of copper contamination of bean crops in this farm appears to be small.     

Natural N abundance of plant and soil inorganic-N as evidence for over-fertilization with compost

To test the hypothesis that N isotope composition can be used as evidence of excessive compost application, we measured variation in patterns of N concentrations and corresponding δ¹⁵N values of plants and soil after compost application. To do so, a pot experiment with Chinese cabbage (Brassica campestris L. cv. Maeryok) was conducted for 42 days. Compost was applied at rates of 0 (SC0), 500 (SC1), 1000 (SC2), and 1500 mg N kg⁻¹ soil (SC3). Plant-N uptake linearly increased with compost application (r² = 0.956, P < 0.05) with an uptake efficiency of 76 g N kg⁻¹ of compost-N at 42 days after application, while dry-mass accumulation did not show such linear increases. Net N mineralized from compost-N increased linearly (r² = 0.998, P < 0.01) with a slope of 122 g N kg⁻¹ of compost-N. Plant-δ¹⁵N increased curvilinearly with increasing compost application, but this increase was insignificant between SC2 and SC3 treatments. The δ¹⁵N of soil inorganic-N (particularly NO₃⁻-N) increased with compost application. We found that plant-δ¹⁵N reflected the N isotope signal of soil NO₃⁻-N at each measurement during plant growth, and that δ¹⁵N of inner leaves and soil NO₃⁻-N was similar when initial NO₃⁻ in the compost was abundant. Therefore, we concluded that δ¹⁵N of whole plant (more obviously in newer plant parts) and soil NO₃⁻-N could reveal whether compost application was excessive, suggesting a possible use of δ¹⁵N in plants and soil as evidence of excess compost application.     

The effect of lime on microbial activity in spruce (Picea abies L.) forests

Summary The use of lime increased heat output and decreased the C:N ratio (global indicators of biological activity) in the organic horizons of a spruce forest (Picea abies L.) soil. These changes were still present after 18 years. During the same period, the muramic acid content increased slightly, while the concentration of both ergosterol and glucosamine decreases. The ratios of ergosterol or glucosamine to muramic acid decreased significantly after 3 years in the plots that had been irrigated and limed, and after 8 years in the limed non-irrigated plots.     

Population and behavioural level responses of arable soil earthworms to boardmill sludge application

The response of earthworms to soil application of boardmill waste sludge was quantified in field and laboratory experiments. The influence of one application of 6×10⁴ tonnes ha⁻¹ of unamended sludge was tested against no application on silty-clay arable soil. After 2 years, results in stubble-cultivated soil showed a 1.7 times lower density of earthworms where sludge was added, whereas there was no difference in average earthworm fresh weight. In direct-drilled soil, there was an indication of lower average numbers of Lumbricus terrestris middens when sludge was applied. Negative impacts in the field may have been due to indirect negative effects of the sludge application. In the laboratory, habitat choice tests were undertaken with two common species from the field site using sludge-based mulch and fertiliser products. Aporrectodea caliginosa showed no discernible preference between soil and soil mixed with unamended sludge (mulch), but favoured soil over soil mixed with sludge and chicken manure (fertiliser). Tests with L. terrestris showed a similar pattern. The preference of L. terrestris for the two products was compared with that for chopped barley straw by direct observation of foraging behaviour. Differences in foraging time of L. terrestris for different feeds were not significant, but mass of straw collected was significantly greater compared with either type of sludge treatment.     

Impact of landspread sewage sludge and earthworm introduction on established earthworms and soil structure

Summary Sewage sludge was applied to twelve 4-m² plots in two forest (mixed hardwood, Norway spruce plantation) site and one old field site. The earthworm Eisenia fetida was introduced to half the control and half the treated plots. Earthworm populations were sampled by formalin extraction and hand-sorting five times in the year following treatment. One year after treatment, soil samples were wet-sieved and water-stable aggregate size-class arrays were determined.The dominant earthworm in the study site, Lumbricus terrestris, increased in density and mean individual biomass in response to sludge treatment in mixed hardwood and old field plots. In the Norway spruce plots, L. terrestris increased in individual biomass but decreased in density following sludge application. The density of the introduced E. fetida rapidly declined in all control plots. One year after introduction, E. fetida was found only in the sludge-treated Norway spruce plot. The introduction of E. fetida with sludge decreased the density and biomass of L. terrestris in the hardwood plots.Sludge treatment increased the percentages of 4-mm diameter water-stable aggregates in old field and hardwood plots. The addition of E. fetida with sludge in the hardwood plots generated no increase in 4-mm water-stable aggregates. In the old field, sludge + E. fetida increased the 4-mm water-stable aggregates. Little change in water-stable aggregates in response to either treatment combination was seen in the Norway spruce site.     

Growth of Lumbricus terrestris and Aporrectodea caliginosa in an acid forest soil, and their effects on enchytraeid populations and soil properties

A laboratory experiment was carried out to test the hypothesis that the earthworms Lumbricus terrestris and Aporrectodea caliginosa are able to maintain their populations and reproduce in the acid forest soil of a deciduous forest where no lumbricids were found in the field. The experiment was conducted in 45-l containers in which layers of mineral subsoil, humus and organic topsoil collected from the site were established. Both species survived and at least L. terrestris reproduced during the 60 weeks’ incubation. Burrows and middens of L. terrestris were recorded and quantities of litter were consumed. The presence of lumbricids increased the organic matter content of humus, reduced the acidity of the topsoil and humus layers, and suppressed the population of the enchytraeid Cognettia sphagnetorum. A dense population of Enchytraeus albidus was found in L. terrestris middens. It is concluded that edaphic factors do not explain the absence of earthworms, but isolation from cultural landscapes and lack of opportunity to colonize the site from the surroundings is the decisive factor.     

Peat amendment and production of different crop plants affect earthworm populations in field soil

A field experiment was conducted to study the effects of peat amendment and crop production system on earthworms. The experiment was established on a field previously cultivated with oats and with silt as the main soil type. Perennial crops strawberry, timothy and caraway, and annual crops rye, turnip rape, buckwheat, onion and fiddleneck were cultivated with conventional methods. All the crops were grown with and without soil amendment with peat. Earthworms were sampled twice: 4 and 28 months after establishment of the experiment. In the former case part of the experimental plots were soil sampled and hand sorted for estimation of earthworms. In the latter case all experimental plots were sampled and both soil sampling and mustard extraction was carried out. Soil organic carbon and microbial biomass was measured at 14 and 28 months. Peat increased the abundance of juvenile Aporrectodea caliginosa by 74% in three growing seasons, but had no effect on adult numbers. Lumbricus terrestris numbers were not increased by peat treatment. Three season cultivation of caraway favoured both A. caliginosa and L. terrestris. An equal abundance of A. caliginosa was also found in plots cultivated with turnip rape and fiddleneck. Total earthworm and especially A. caliginosa numbers were very small in plastic-mulched strawberry beds. This was mainly attributed to repeated use of the insecticide endosulfan. With the strawberry plots omitted there was a significant correlation between soil microbial N measured at 14 months and juvenile Aporrectodea spp. and Lumbricus spp. numbers measured at 28 months. Adult earthworm numbers were not associated with either soil organic C or microbial biomass.     

Influence of cropping system and nitrogen input on soil fauna and microorganisms in a Swedish arable soil

Summary The development of a number of components was analysed in an agro-ecosystem study with four cropping regimens, barley without and with N fertilization, grass ley, and lucerne. A great variation in N inputs (1–39 g N m^-2 year^-1) and cropping systems produced a variation in primary production (260–790 g C m^-2 year^-1) and input of organic material to the soil (150–270 g C m^-2 year^-1). This was reflected in variations of total soil animal biomass (1.6–5.1 g C m^-2) and in variations in the abundance of various animal groups, nematodes (5.6–9.8×10⁶m^-2), micro- (2.6–4.8×10^-4 m^-2), and macroarthropods (0.9–4.2×10³ m^-2). In contrast, total bacteria, fungi, flagellates, and amoebae varied quite independently of the organic matter input. Mineralization processes covaried more with C and N inputs and total animal biomass than with microbial biomass. it is suggested that the rather constant microbial biomass was a result of an adjustment in the grazing pressure of microbial-feeding animals to the level of microbial production.Dedicated to the late Prof. Dr. W. Kühnelt