Quantification of nitrogen excretion rates for three lumbricid earthworms using 15N

 Nitrogen excretion rates of ¹⁵N-labeled earthworms and contributions of ¹⁵N excretion products to organic (dissolved organic N) and inorganic (NH₄-N, NO₃-N) soil N pools were determined at 10  °C and 18  °C under laboratory conditions. Juvenile and adult Lumbricus terrestris L., pre-clitellate and adult Aporrectodea tuberculata (Eisen), and adult Lumbricus rubellus (Hoffmeister) were labeled with ¹⁵N by providing earthworms with ¹⁵N-labeled organic substrates for 5–6 weeks. The quantity of ¹⁵N excreted in unlabeled soil was measured after 48 h, and daily N excretion rates were calculated. N excretion rates ranged from 274.4 to 744 μg N g^–1 earthworm fresh weight day^–1, with a daily turnover of 0.3–0.9% of earthworm tissue N. The N excretion rates of juvenile L. terrestris were significantly lower than adult L. terrestris, and there was no difference in the N excretion rates of pre-clitellate and adult A. tuberculata. Extractable N pools, particularly NH₄-N, were greater in soils incubated with earthworms for 48 h than soils incubated without earthworms. Between 13 and 40% of excreted ¹⁵N was found in the ¹⁵N-mineral N (NH₄-N+NO₃-N) pool, and 13–23% was in the ¹⁵N-DON pool. Other fates of excreted ¹⁵N may have been incorporation in microbial biomass, chemical or physical protection in non-extractable N forms, or gaseous N losses. Earthworm excretion rates were combined with earthworm biomass measurements to estimate N flux from earthworm populations through excretion. Annual earthworm excretion was estimated at 41.5 kg N ha^–1 in an inorganically-fertilized corn agroecosystem, and was equivalent to 22% of crop N uptake. Our results suggest that the earthworms could contribute significantly to N cycling in corn agroecosystems through excretion processes. Received: 12 April 1999     

Earthworm populations are stable in temperate agricultural soils receiving wood-based biochar

The application of decomposable organic residues such as manure and crop litter is generally beneficial to earthworms. There is an emerging interest in applying biochar, a carbonaceous product of pyrolysis, to temperate agricultural soils. The slow decomposition rate of biochar, which also contains ash and combustion byproducts, could be detrimental to earthworms. The objective of this study was to describe the earthworm populations in biochar-amended soils on a dairy farm in the St. Francis River watershed, Quebec, Canada. Earthworms were collected from replicated field plots under cereal production. Site A received three wood-based biochar types at two application rates(5 and 10 t ha~(-1) biochar) plus an unamended control, while Site B received wood-based biochar(7.5 t ha~(-1) biochar), dairy cattle slurry(10 t ha~(-1) manure), or a combination of the biochar and manure rates plus an unamended control. Earthworms were collected by hand sorting and formaldehyde expulsion from soil pits. Three species, Aporrectodea turgida, Aporrectodea tuberculata, and Lumbricus rubellus, were found at the sites, and Aporrectodea was the dominant genus. Biochar sources, rates, and application with dairy slurry did not affect the earthworm population, which had 52–218 individuals m~(-2) in 2010 and 4–96 individuals m~(-2) in 2011. The seasonal variation in earthworm population may be due to flooding in the spring of 2011, which apparently interfered with earthworm reproduction. The similarity in earthworm abundance and biomass in plots with and without wood-based biochar leads to the conclusion that earthworm populations are stable in biochar-amended soils in this cold, humid temperate region.     

Species-specific earthworm population responses in relation to flooding dynamics in a Dutch floodplain soil

Earthworms dominate the animal biomass in moist floodplain soils. They are known to survive long periods in aerated water, but little is known about earthworm population dynamics in floodplain systems with changing inundation frequencies. This study determined earthworm population dynamics in a floodplain system, in relation to frequency and duration of flooding events. From October 2000 to May 2003 earthworms were hand sorted in the ‘Afferdensche en Deestsche Waarden’, a floodplain on the south bank of the river Rhine, near Druten, The Netherlands. Earthworm numbers and biomasses per age class (adult, subadult, juvenile) were recorded. Numbers and biomasses tend to decrease during flooding. Lumbricus terrestris was found in high numbers (>10/m²) only at the end of a flooding period. Allolobophora chlorotica was hardly affected by flooding; their biomass remained stable during the year. Aporrectodea caliginosa showed fluctuating numbers and biomasses during the sampling period that did not correlate with flooding frequency. Numbers and biomasses of Lumbricus rubellus were strongly reduced at the end of each flooding event, but their population densities fully recovered until next flooding event. Earthworm populations in floodplains fluctuate in time, depending on the season and on the time, duration and frequency of flooding. Different earthworm species react differently towards these flooding dynamics.     

Earthworm channels in cultivated clayey and loamy Norwegian soils

Studies of earthworm species and their activity, expressed as channels, on cultivated loamy (Humic Cryaquept) and clayey soils (Typic Endoaqualf and Typic Cryaquept) were conducted in southern and central Norway before conversion from conventional to organic cropping systems. At all the three study areas: Landvik (Grimstad), Voll (Ås) and Kvithamar (Stjørdal), the earthworm species Aporrectodea caliginosa, Aporrectodea rosea, Allolobophora chlorotica and Lumbricus rubellus were found. At Landvik, where the deep-burrowing species Aporrectodea longa, A. caliginosa and Lumbricus terrestris dominated, medium (4–6 mm) and coarse (>6 mm) earthworm channels were most numerous below the plough layer. Almost no coarse pores were found at Voll and Kvithamar. The volume of earthworm channels below the plough layer was 0.6–0.8% of total soil volume at Landvik and Voll and 0.3–0.4% at Kvithamar. Earthworm channels >6 mm below the plough layer were useful to identify present and previous activity of deep-burrowing earthworms such as L. terrestris. Below the plough layer, roots were almost entirely restricted to earthworm channels and interaggregate spaces.     

Earthworm species composition affects the soil bacterial community and net nitrogen mineralization

Knowledge of the effects of species diversity within taxonomic groups on nutrient cycling is important for understanding the role of soil biota in sustainable agriculture. We hypothesized that earthworm species specifically affect nitrogen mineralization, characteristically for their ecological group classifications, and that earthworm species interactions would affect mineralization through competition and facilitation effects. A mesocosm experiment was conducted to investigate the effect of three earthworm species, representative of different ecological groups (epigeic: Lumbricus rubellus; endogeic: Aporrectodea caliginosa tuberculata; and anecic: Lumbricus terrestris), and their interactions on the bacterial community, and on nitrogen mineralization from ¹⁵N-labelled crop residue and from soil organic matter.Our results indicate that L. rubellus and L. terrestris enhanced mineralization of the applied crop residue whereas A. caliginosa had no effect. On the other hand, L. rubellus and A. caliginosa enhanced mineralization of the soil organic matter, whereas L. terrestris had no effect. The interactions between different earthworm species affected the bacterial community and the net mineralization of soil organic matter. The two-species interactions between L. rubellus and A. caliginosa, and L. rubellus and L. terrestris, resulted in reduced mineral N concentrations derived from soil organic matter, probably through increased immobilization in the bacterial biomass. In contrast, the interaction between A. caliginosa and L. terrestris resulted in increased bacterial growth rate and reduced total soil C. When all three species were combined, the interaction between A. caliginosa and L. terrestris was dominant. We conclude that the effects of earthworms on nitrogen mineralization depend on the ecological traits of the earthworm species present, and can be modified by species interactions. Knowledge of these effects can be made useful in the prevention of nutrient losses and increased soil fertility in agricultural systems, that typically have a low earthworm diversity.     

Earthworms increase soil microbial biomass carrying capacity and nitrogen retention in northern hardwood forests

Earthworms have been shown to produce contrasting effects on soil carbon (C) and nitrogen (N) pools and dynamics. We measured soil C and N pools and processes and traced the flow of ¹³C and ¹⁵N from sugar maple (Acer saccharum Marsh.) litter into soil microbial biomass and respirable C and mineralizable and inorganic N pools in mature northern hardwood forest plots with variable earthworm communities. Previous studies have shown that plots dominated by either Lumbricus rubellus or Lumbricus terrestris have markedly lower total soil C than uncolonized plots. Here we show that total soil N pools in earthworm colonized plots were reduced much less than C, but significantly so in plots dominated by contain L. rubellus. Pools of microbial biomass C and N were higher in earthworm-colonized (especially those dominated by L. rubellus) plots and more ¹³C and ¹⁵N were recovered in microbial biomass and less was recovered in mineralizable and inorganic N pools in these plots. These plots also had lower rates of potential net N mineralization and nitrification than uncolonized reference plots. These results suggest that earthworm stimulation of microbial biomass and activity underlie depletion of soil C and retention and maintenance of soil N pools, at least in northern hardwood forests. Earthworms increase the carrying capacity of soil for microbial biomass and facilitate the flow of N from litter into stable soil organic matter. However, declines in soil C and C:N ratio may increase the potential for hydrologic and gaseous losses in earthworm-colonized sites under changing environmental conditions.     

Earthworm response to rotation and tillage in a Missouri claypan soil

 Agricultural management practices affect earthworm populations. A field experiment was conducted to determine the effect of two rotations and two tillage systems on earthworm population density and biomass in a claypan soil. The rotations were soybean/corn and wheat/corn, and the tillage systems were conventional tillage (chisel plowed and disked) and no-tillage. Earthworm and soil samples were collected in fall 1995, spring 1996, and fall 1996. Aporrectodea trapezoides and Diplocardia singularis were the species identified at the site. A. trapezoides accounted for 92–96% of the total earthworm population density and D. singularis accounted for only 4–8%. In a no-till system, soybean/corn rotation resulted in significantly greater population density of A. trapezoides compared with the wheat/corn rotation. Crop residue quality (low C:N ratio) and quantity were important factors in increasing A. trapezoides population density and biomass. Conventional tillage markedly decreased population density and biomass of both earthworm species. Our results suggest that rotation and tillage significantly affect earthworm population density and biomass. Received: 6 June 1998     

Earthworm feeding activity and development of the humus profile

Earthworm casts and digestive tract contents were simultaneously examined, using the same methods, in a recently formed humus profile in a mountain spruce forest. Earthworm species had distinct diets and an earthworm foodweb could be distinguished. Lumbricus terrestris and Aporrectodea icterica were distinct from the other species examined: the former to some extent as a litter consumer, and both species because they excavated mineral material which was deposited within new top layers of the mull humus. Aporrectodea nocturna and Aporrectodea caliginosa both had a non-specific soil feeding mode. Most of the species enriched the humus profile with amorphous organic matter finely incorporated within a mineral matrix. Besides different food selection, a network of burrows was produced as a consequence of the different burrowing behaviour of each earthworm species. Received: 2 January 1997     

Impact of earthworms on decomposition of garden refuse

Summary Decomposition of garden refuse was studied in containers with and without the earthworm species Eisenia andrei and Lumbricus rubellus. The reduction of cellulose and hemicellulose was greater where earthworms were present. Respiration was similar regardless of the presence of earthworms, whereas dry matter reduction, on average, was greatest without earthworms. The earthworm biomass decreased during the 58 days of the experiment; E. andrei increased in biomass, whereas L. rubellus died out.     

Effects of pasture improvement and intensive cultivation on microbial biomass,enzyme activities,and composition and size of earthworm populations

We investigated the quantity and distribution of organic C, microbial biomass C, protease, arylsulphatase and arylphosphatase activity, and earthworm numbers and biomass in the soil from a 37-year-old grazed pasture supplied with superphosphate at rates of 0, 188, and 376 kg ha^-1 annually. The results were compared with a non-irrigated wilderness site which had not been used for agriculture and an arable site that had been intensively cultivated for 11 consecutive years. In the 0- to 5-cm layer, organic C followed the trend arableAporrectodea caliginosa (77–89% of total numbers) although Lumbricus rubellus made an increasing contribution to the population with increasing superphosphate rates. In the unirrigated wilderness site the population consisted of 56% A. caliginosa and 44% L. rubellus. While Octolasion cyaneum and A. rosea made up a small proportion of the population in the improved pasture sites, they were not present in the wilderness or arable sites. A. caliginosa was the only species present in the arable site. The mean fresh weight of individuals followed the order arable相似文献   

Earthworms and pH affect communities of nematodes and enchytraeids in forest soil

In northern boreal forests the occurrence of endogeic and anecic earthworms is determined by soil pH. Increasing evidence suggests that large detritivorous soil animals such as earthworms can influence the other components of the decomposer community. To study the effects of earthworms and pH on soil nematode and enchytraeid communities, a factorially designed experiment was conducted with Lumbricus rubellus and/or Aporrectodea caliginosa. Earthworms were added to "mesocosms" containing unlimed (pH 4.8) or limed (pH 6.1) coniferous mor humus with their natural biota of micro-organisms. In the absence of earthworms, nematodes were significantly more abundant in limed than in unlimed humus. Earthworms markedly decreased the numbers of nematodes both in unlimed and limed soils. Earthworm activities eliminated enchytraeids in unlimed soil, but liming improved the survival of some species. It was concluded that liming of soil, either alone or mediated by the earthworm populations, is likely to affect soil nematode and enchytraeid community and mineralisation.     

Earthworm populations in a northern U.S. Cornbelt soil are not affected by long-term cultivation of Bt maize expressing Cry1Ab and Cry3Bb1 proteins

Earthworms, which play a key role in biogeochemical processes in soil ecosystems, could be negatively affected by the cultivation of transgenic Bt crops. Studies to date have found few effects of Bt maize on earthworm species. If adverse effects occur, they are likely to be chronic or sub-lethal and expressed over large spatial and temporal scales. Our objective in the present study was to investigate potential effects on earthworm populations in soil cultivated with Bt maize in a large multiple-year field study. We surveyed the earthworm populations in 0.16-ha experimental field plots of two varieties of Cry1Ab Bt maize, one variety of Cry3Bb1 Bt maize, and three non-transgenic control varieties cultivated for four years. Four earthworm species were found in our sample: Aporrectodea caliginosa, Aporrectodea trapezoides, Aporrectodea tuberculata (collectively, the A. caliginosa species complex), and Lumbricus terrestris. We found no significant differences in the biomass of juveniles and adults for all four species between Bt and non-Bt maize varieties. From this and previous studies, we conclude that the effects of Cry1Ab and Cry3Bb1 Bt maize on the A. caliginosa species complex and L. terrestris are small. Nonetheless, general conclusions about the effects of Bt maize on earthworm populations are not warranted due to the small number of species tested. In future laboratory studies, earthworm species should be selected according to their association with a Bt crop and the impact of that species to valued soil ecosystem processes.     

Using FAME profiles for the characterization of animal wastes and vermicomposts

This study investigated the possibility of fingerprinting different organic wastes (cow, pig and horse manure) and the vermicomposts produced by different earthworm species (Eisenia andrei, Eudrilus eugeniae and Lumbricus rubellus) analyzing the profiles of fatty acid methyl esters (FAMEs). We found clear differences between their microbial communities, demonstrating the power and sensitivity of the total FAME analysis. In addition, qualitative and quantitative analyses of specific biomarkers permitted to determine differences between samples and to evaluate the effect of earthworms in the decomposition of organic matter. Fatty acid profiles were largely determined by the different vermicomposting earthworm species. Fatty acid 18:2ω6 increased significantly in horse manure vermicomposted by L. rubellus and in cow manure vermicomposted by the three earthworm species, whereas it decreased significantly in pig manure vermicomposted by L. rubellus and E. eugeniae. Fatty acid 20:4ω6 increased significantly in all vermicomposts obtained with the three earthworm species.     

Earthworm populations and growth rates related to long-term crop residue and tillage management

Conventional tillage creates soil physical conditions that may restrict earthworm movement and accelerate crop residue decomposition, thus reducing the food supply for earthworms. These negative impacts may be alleviated by retaining crop residues in agroecosystems. The objective of this study was to determine the effects of various tillage and crop residue management practices on earthworm populations in the field and earthworm growth under controlled conditions. Population assessments were conducted at two long-term (15+ years) experimental sites in Québec, Canada with three tillage systems: moldboard plow/disk harrow (CT), chisel plow or disk harrow (RT) and no tillage (NT), as well as two levels of crop residue inputs (high and low). Earthworm growth was assessed in intact soil cores from both sites. In the field, earthworm populations and biomass were greater with long-term NT than CT and RT practices, but not affected by crop residue management. Laboratory growth rates of Aporrectodea turgida (Eisen) in intact soil cores were affected by tillage and residue inputs, and were positively correlated with the soil organic C pool, suggesting that tillage and residue management practices that increase the soil organic C pool provide more organic substrates for earthworm growth. The highest earthworm growth rates were in soils from RT plots with high residue input, which differed from the response of earthworm populations to tillage and residue management treatments in the field. Our results suggest that tillage-induced disturbance probably has a greater impact than food availability on earthworm populations in cool, humid agroecosystems.     

Earthworm influence on carbon dioxide and nitrous oxide fluxes from an unfertilized corn agroecosystem

Earthworms modify the soil environment through their feeding, casting, and burrowing activities, which may lead to more decomposition and respiration in aerobic microsites and more denitrification in anaerobic microsites. The objective of this study was to determine whether earthworms increase CO₂ and N₂O fluxes from an unfertilized corn agroecosystem. Earthworm populations within field enclosures (2.9 m²) were reduced by repeatedly applying carbaryl insecticide, then single and mixed populations of Lumbricus terrestris L. and Aporrectodea caliginosa (Savigny) were added. Gas samples were collected once a week for 14 weeks, from June to September 2005. Carbaryl applications reduced, but did not eliminate earthworms from enclosures. The CO₂ and N₂O fluxes were affected by the sampling date, with peak gas fluxes after rainfall events. Mean CO₂ and N₂O fluxes during the study period tended to be greater from enclosures with added earthworms than the control (no earthworms added), but were not significantly affected by earthworm treatments due to the low survival rate of introduced earthworms. Better control of earthworm populations in the field is required to fully assess the impact of earthworms on CO₂ and N₂O fluxes from temperate agroecosystems.     

Effects of endogeic earthworms on soil processes and plant growth in coniferous forest soil

Summary The effects of the endogeic earthworm, Aporrectodea caliginosa tuberculata (Eisen) on decomposition processes in moist coniferous forest soil were studied in the laboratory. The pH preference of this species and its effects on microbial activity, N and P mineralization, and the growth of birch seedlings were determined in separate pot experiments. Homogenized humus from a spruce stand was shown to be too acid for A. c. tuberculata. After liming, the earthworms thrived in the humus and their biomass increased (at pH above 4.8). In later experiments in which the humus was limed, the earthworms positively influenced the biological activity in humus and also increased the rate of N mineralization. A. c. tuberculata increased the growth of birch seedlings, with increases observed in stems, leaves, and roots. Neither NH ₄ ⁺ -N fertilizer nor mechanical mixing with artificial worms affected seedling growth. No plant-growth-affecting compounds (e.g., hormone-like compounds) due to the earthworms were present in the humus. The shoot: root ratio in the birch seedlings was not affected by either the earthworms or the fertilizer. The experiments revealed the impact of earthworm activity on soil processes and plant growth.     

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.     

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

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

Earthworms increase the ratio of bacteria to fungi in northern hardwood forest soils, primarily by eliminating the organic horizon

The exotic earthworm invasion in hardwood forests of the northern United States is associated with many ecosystem-level changes. However, less is known about the effects of the invasion on the composition of the soil microbial community through which ecosystem-level changes are mediated. Further, earthworm effects on soil microbial community composition have not been well studied in the field. To evaluate changes in bacterial and fungal abundance associated with the earthworm invasion we quantified bacterial and fungal biomass by microscopic counts in paired earthworm-invaded (earthworm) and earthworm-free (reference) plots in five forest stands in central New York (USA). Earthworms significantly increased the ratio of bacteria to fungi on an area basis (per m²), by more than two times in mid-summer and early autumn. While this effect was associated primarily with the lack of the fungal-dominated organic horizon in earthworm plots, a higher ratio of bacteria to fungi in the surface 5 cm mineral soil also contributed as it developed between spring and mid-summer. Earthworm reduction of fungal biomass was confirmed by substantially lower growth of fungal hyphae into mesh sand bags in earthworm compared to reference plots. Burrowing activity by the earthworm Lumbricus terrestris increased the ratio of bacteria to fungi over the short-term within earthworm plots, introducing small-scale spatial heterogeneity associated with burrows. Our study suggests that the exotic earthworm invasion in these northern hardwood forests markedly increased the ratio of bacteria to fungi by eliminating the fungal-rich organic horizon, and was associated localized increases in bacterial vs. fungal abundance in mineral soil, setting the stage for future research into linkages between the earthworm invasion, bacterial and fungal abundance, and ecosystem processes.     

Changes in potassium availability and other soil properties due to soil ingestion by earthworms

An incubation experiment was conducted to study the changes that occur in potassium availability and other soil properties with ingestion of soil by earthworms. Two soils were used. Raumai soil with high non-exchangeable K and Milson soil with low non-exchangeable K were incubated with two species of earthworm, Aporrectodea caliginosa and Lumbricus rubellus, for 8 weeks. The casts and soil samples were analysed for exchangeable K, Ca, Mg, Na, and H, pH, organic C, and texture. The results indicated that in Raumai soil, the exchangeable K levels of the casts of both earthworm species were significantly higher than for the control soil, the effect being more marked for L. rubellus than for A. caliginosa. In Milson soil, the exchangeable K levels were significantly lower in the casts of both types of earthworm than in the control soil. The nitric acid-extractable K of the soil and casts was not markedly different for either soil type, but available non-exchangeable K values were significantly higher for the casts of L. rubellus from Milson soil than for the noningested Milson soil. In Raumai soil, the exchangeable Ca was higher in the casts of L. rubellus, exchangeable Mg and H were reduced, and exchangeable Na did not change markedly in the cast compared to the control soil. For Milson soil, the casts contained lower exchangeable Ca and H but higher Na and Mg than the control. The casts of both species of earthworm had significantly higher pH values for both soil types. There was no marked difference in the organic C content of the control soil and cast samples for Milson but a reduction in the casts of A. caliginosa for the Raumai soil. Finer fractions increased in the casts of both earthworm species in both soil types.