Interactions between residue placement and earthworm ecological strategy affect aggregate turnover and N2O dynamics in agricultural soil

Previous laboratory studies using epigeic and anecic earthworms have shown that earthworm activity can considerably increase nitrous oxide (N₂O) emissions from crop residues in soils. However, the universality of this effect across earthworm functional groups and its underlying mechanisms remain unclear. The aims of this study were (i) to determine whether earthworms with an endogeic strategy also affect N₂O emissions; (ii) to quantify possible interactions with epigeic earthworms; and (iii) to link these effects to earthworm-induced differences in selected soil properties. We initiated a 90-day ¹⁵N-tracer mesocosm study with the endogeic earthworm species Aporrectodea caliginosa (Savigny) and the epigeic species Lumbricus rubellus (Hoffmeister). ¹⁵N-labeled radish (Raphanus sativus cv. Adagio L.) residue was placed on top or incorporated into the loamy (Fluvaquent) soil. When residue was incorporated, only A. caliginosa significantly (p < 0.01) increased cumulative N₂O emissions from 1350 to 2223 μg N₂O-N kg⁻¹ soil, with a corresponding increase in the turnover rate of macroaggregates. When residue was applied on top, L. rubellus significantly (p < 0.001) increased emissions from 524 to 929 μg N₂O-N kg⁻¹, and a significant (p < 0.05) interaction between the two earthworm species increased emissions to 1397 μg N₂O-N kg⁻¹. These effects coincided with an 84% increase in incorporation of residue ¹⁵N into the microaggregate fraction by A. caliginosa (p = 0.003) and an 85% increase in incorporation into the macroaggregate fraction by L. rubellus (p = 0.018). Cumulative CO₂ fluxes were only significantly increased by earthworm activity (from 473.9 to 593.6 mg CO₂-C kg⁻¹ soil; p = 0.037) in the presence of L. rubellus when residue was applied on top. We conclude that earthworm-induced N₂O emissions reflect earthworm feeding strategies: epigeic earthworms can increase N₂O emissions when residue is applied on top; endogeic earthworms when residue is incorporated into the soil by humans (tillage) or by other earthworm species. The effects of residue placement and earthworm addition are accompanied by changes in aggregate and SOM turnover, possibly controlling carbon, nitrogen and oxygen availability and therefore denitrification. Our results contribute to understanding the important but intricate relations between (functional) soil biodiversity and the soil greenhouse gas balance. Further research should focus on elucidating the links between the observed changes in soil aggregation and controls on denitrification, including the microbial community.     

Effect of tillage on earthworms over short- and medium-term in conventional and organic farming

Earthworms play an important role in many soil functions and are affected by soil tillage in agricultural soils. However, effects of tillage on earthworms are often studied without considering species and their interactions with soil properties. Furthermore, many field studies are based on one-time samplings that do not allow for characterisation of temporal variation. The current study monitored the short (up to 53 days) and medium term (up to 4 years) effects of soil tillage on earthworms in conventional and organic farming. Earthworm abundances decreased one and three weeks after mouldboard ploughing in both conventional and organic farming, suggesting direct and indirect mechanisms. However, the medium-term study revealed that earthworm populations in mouldboard ploughing systems recovered by spring. The endogeic species Aporrectodea caliginosa strongly dominated the earthworm community (76%), whereas anecic species remained <1% of all earthworms in all tillage and farming systems over the entire study. In conventional farming, mean total earthworm abundance was not significantly different in reduced tillage (153 m⁻²) than mouldboard ploughing (MP; 130 m⁻²). However, reduced tillage in conventional farming significantly increased the epigeic species Lumbricus rubellus from 0.1 m⁻² in mouldboard ploughing to 9 m⁻² averaged over 4 years. Contrastingly, in organic farming mean total earthworm abundance was 45% lower in reduced tillage (297 m⁻²) than MP (430 m⁻²), across all sampling dates over the medium-term study (significant at 3 of 6 sampling dates). Reduced tillage in organic farming decreased A. caliginosa from 304 m⁻² in mouldboard ploughing to 169 m⁻² averaged over 4 years (significant at all sampling dates). Multivariate analysis revealed clear separation between farming and tillage systems. Earthworm species abundances, soil moisture, and soil organic matter were positively correlated, whereas earthworm abundances and penetration resistance where negatively correlated. Variability demonstrated between sampling dates highlights the importance of multiple samplings in time to ascertain management effects on earthworms. Findings indicate that a reduction in tillage intensity in conventional farming affects earthworms differently than in organic farming. Differing earthworm species or ecological group response to interactions between soil tillage, crop, and organic matter management in conventional and organic farming has implications for management to maximise soil ecosystem functions.     

Impact of ecologically different earthworm species on soil water characteristics

A laboratory experiment was performed to assess the impact of ecologically different earthworm species on soil water characteristics, such as soil tension, water content, and water infiltration rate. Three earthworm species (Lumbricus rubellus, Aporrectodea caliginosa, Lumbricus terrestris) were exposed in soil columns (diameter 30 cm, height 50 cm) for 100 days with a total fresh earthworm biomass of 22.7 ± 0.4 g per column, each in duplicate. Each column was equipped with tensiometers at 10 and 40 cm and FD-probes at 10 cm depth, to continuously measure the temporal development of soil tension and soil moisture. Additionally, 30 g of sieved and rewetted horse manure was placed on the soil surface as a food source. Precipitation events (10 mm) were simulated at day 28 and day 64. At the end of the experiment the water infiltration rate and the runoff at 55 cm depth were determined.The results showed considerable evidence, that ecologically different earthworms modify soil water characteristics in different ways. The anecic L. terrestris and the endogeic A. caliginosa showed the tendency to enhance the drying of the topsoil and subsoil. Their intensive and deep burrowing activity might enhance the exchange of water vapor due to a better aeration in soil. In contrast, the epigeic L. rubellus tended to enhance the storage of soil moisture in the topsoil, which might be linked to lower rates of litter loss from soil surface and thus a thicker litter layer remaining. A. caliginosa led to considerable higher water infiltration rates and faster water discharges in the subsoil, relative to the other species, probably due to a high soil dwelling activity.     

Endogeic and anecic earthworm abundance in six Midwestern cropping systems

Endogeic and juvenile anecic earthworm abundance was measured in soil samples and anecic populations were studied by counting midden numbers at the sites of two long-term cropping systems trials in South-central Wisconsin. The three grain and three forage systems at each site were designed to reflect a range of Midwestern USA production strategies. The primary objectives of this work were to determine if the abundance of endogeic or anecic earthworms varied among cropping systems or crop phases within a cropping system and were there specific management practices that impacted endogeic or anecic earthworm numbers. The earthworms present in the surface soil were: Aporrectodea tuberculata (Eisen), A. caliginosa (Savigny), A. trapezoides (Dugés); and juvenile Lumbricus terrestris (L.). True endogeic abundance was greatest in rotationally grazed pasture [188 m^?2 at Arlington (ARL) and 299 m^?2 at Elkhorn (ELK)], and smallest in conventional continuous corn (27 m^?2 at ARL and 32 m^?2 at ELK). The only type of anecic earthworm found was L. terrestris L. There was an average of 1.2 middens per adult anecic earthworm and the population of anecics was greatest in the no-till cash grain system (28 middens m^?2 at ARL, 18 m^?2 at ELK) and smallest in the conventional continuous corn system (3 middens m^?2 at ARL, 1 m^?2 at ELK). Earthworm numbers in individual crop phases within a cropping system were too variable from year-to-year to recommend using a single phase to characterize a whole cropping system. Indices for five management factors (tillage, manure inputs, solid stand, pesticide use, and crop diversity) were examined, and manure use and tillage were the most important impacting earthworm numbers across the range of cropping systems. Manure use was the most important management factor affecting endogeic earthworm numbers; but no-tillage was the most important for the juvenile and adult anecic groups and had a significantly positive influence on endogeic earthworm counts as well. The pesticides used, which were among the most commonly applied pesticides in the Midwestern USA, and increasing crop diversity did not have a significant effect on either the endogeic or anecic earthworm groups in this study. Consequently, designing cropping systems that reduce tillage and include manure with less regard to omitting pesticides or increasing crop diversity should enhance earthworm populations and probably improve sustainability.     

Earthworm activity as a determinant for N2O emission from crop residue

Earthworm activity may have an effect on nitrous oxide (N₂O) emissions from crop residue. However, the importance of this effect and its main controlling variables are largely unknown. The main objective of this study was to determine under which conditions and to what extent earthworm activity impacts N₂O emissions from grass residue. For this purpose we initiated a 90-day (experiment I) and a 50-day (experiment II) laboratory mesocosm experiment using a Typic Fluvaquent pasture soil with silt loam texture. In all treatments, residue was applied, and emissions of N₂O and carbon dioxide (CO₂) were measured. In experiment I the residue was applied on top of the soil surface and we tested (a) the effects of the anecic earthworm species Aporrectodea longa (Ude) vs. the epigeic species Lumbricus rubellus (Hoffmeister) and (b) interactions between earthworm activity and bulk density (1.06 vs. 1.61 g cm⁻³). In experiment II we tested the effect of L. rubellus after residue was artificially incorporated in the soil. In experiment I, N₂O emissions in the presence of earthworms significantly increased from 55.7 to 789.1 μg N₂O-N kg⁻¹ soil (L. rubellus; p<0.001) or to 227.2 μg N₂O-N kg⁻¹ soil (A. longa; p<0.05). This effect was not dependent on bulk density. However, if the residue was incorporated into the soil (experiment II) the earthworm effect disappeared and emissions were higher (1064.2 μg N₂O-N kg⁻¹ soil). At the end of the experiment and after removal of earthworms, a drying/wetting and freezing/thawing cycle resulted in significantly higher emissions of N₂O and CO₂ from soil with prior presence of L. rubellus. Soil with prior presence of L. rubellus also had higher potential denitrification. We conclude that the main effect of earthworm activity on N₂O emissions is through mixing residue into the soil, switching residue decomposition from an aerobic and low denitrification pathway to one with significant denitrification and N₂O production. Furthermore, A. longa activity resulted in more stable soil organic matter than L. rubellus.     

Emission of nitrous oxide from hydrocarbon contaminated soil amended with waste water sludge and earthworms

Soils in Mexico are often contaminated with hydrocarbons and addition of waste water sludge and earthworms accelerates their removal. However, little is known how contamination and subsequent bioremediation affects emissions of N₂O and CO₂. A laboratory study was done to investigate the effect of waste water sludge and the earthworm Eisenia fetida on emission of N₂O and CO₂ in a sandy loam soil contaminated with the polycyclic aromatic hydrocarbons (PAHs): phenanthrene, anthracene and benzo(a)pyrene. Emissions of N₂O and CO₂, and concentrations of inorganic N (ammonium (NH₄⁺), nitrite (NO₂^?) nitrate (NO₃^?)) were monitored after 0, 5, 24, 72 and 168 h. Adding E. fetida to the PAHs contaminated soil increased CO₂ production rate significantly 2.0 times independent of the addition of sludge. The N₂O emission rate from unamended soil expressed on a daily base was 5 μg N kg^?1 d^?1 for the first 2 h and increased to a maximum of 325 μg N kg^?1 d^?1 after 48 h and then decreased to 10 μg N kg^?1 d^?1 after 168 h. Addition of PAHs, E. fetida or PAHs + E. fetida had no significant effect on the N₂O emission rate. Adding sludge to the soil sharply increased the N₂O emission rate to >400 μg N kg^?1 d^?1 for the entire incubation with a maximum of 1134 μg N kg^?1 d^?1 after 48 h. Addition of E. fetida, PAHs or PAHs + E. fetida to the sludge-amended soil reduced the N₂O emission rate significantly compared to soil amended with sludge after 24 h. It was found that contaminating soil with PAHs and adding earthworms had no effect on emissions of N₂O. Emission of N₂O, however, increased in sludge-amended soil, but addition of earthworms to this soil and contamination reduced it.     

Factors influencing earthworm communities in post-industrial areas of Krakow Soda Works

Application of earthworm in soil re-cultivation and re-creation in post-industrial ecosystems make a big challenge for temporal applied zoology. The sediments of the Krakow Soda Works “Solvay” have undergone land reclamation in different ways: older sediments traps were left without any re-cultivation practices; meanwhile the newest ones were reclaimed using standard method (new soil cover planted with combination of grass and leguminous plants). The effect of different treatments on community and population structure of earthworm was estimated during consecutive years 1999–2000. Six localities differing in time of establishment, reclamation processes, vegetation type and soil properties were chosen. Nine species were recorded, among which Aporrectodea caliginosa occurred in all localities, being also the most abundant. Two other species, Lumbricus rubellus and Dendrobaena octaedra, which are epigeic species, become most important in forest assemblages and were characteristic for communities of older succession processes. Abundance of adult forms as well as total biomass were significantly affected by soil depth (r = 0.75, P < 0.05, r = 0.917, P < 0.001, respectively). Species richness however was connected with higher amount of macroelements and average plant height. Shannon diversity index and its evenness negatively correlated only with forestation (r = –0.67, P < 0.05, r = –0.68, P < 0.05, respectively). Niche overlap (α Pianka) for all analysed species extracted two groups differing in environmental requirements. First contained epigeic earthworms, closely related to plant succession (PCA results), the other one grouped endogeic and anecique species correlated significantly with soil depth and plant density. Community structure of earthworms do not reflects succession changes in post-industrial habitats, but is strongly affected by microhabitat factors in local scale (mainly soil depth and plant density).     

The efficiency of soil hand-sorting in assessing the abundance and biomass of earthworm communities. Its usefulness in population dynamics and cohort analysis studies

Pit digging and manually revising soil blocks is a frequently used method used for field studies of earthworm communities. The aim of this study was to compare the efficiency of hand-sorting (HS) to extract small earthworms, ca. 0.2 g, and the usefulness in studies of population dynamics and cohort analysis. Many earthworms are not recovered when revising manually the soil. Factors include soil characteristics, i.e. moisture, texture, etc. and also a human factor, which is more relevant if the study is conducted in the long-term. We used data collected in a field study of earthworm communities during 2 years in the savannas of Colombia. Small soil blocks (20 × 20 × 20 cm) were dug out in order to collect the smallest earthworms by washing-sieving (WS) and compare the results with the standard HS of large monoliths (100 × 100 × 50 cm). In fact, this methodology has rarely been addressed in earthworm population field studies. Our results showed that HS efficiency varied owing to the species and ranged from 31.4% up to 100% in the savanna and from 44% to 80% in the pasture, for two small species, i.e Aymara n. sp. (epigeic) and Ocnerodrilidae sp. (endogeic). In the case of the Glossodrilus n. sp. (endogeic) these values were similar, i.e. 51.7% and 58.1%, in the savanna and pasture, respectively. We also used frequency tables to calculate the average efficiency of HS 1 m² soil cores for each weight class in each species in order to obtain a population density correction factor. This allowed us to make corrections in earthworm density in the histograms for population dynamics analysis. We conclude that this method should be the modus operandi in long-term earthworm demography studies.     

Effects of cropping and tree density on earthworm community composition and densities in central Cameroon

Earthworms can have positive effects upon crop growth in the tropics. If soils are to be managed sustainably, then more attention should be paid to the effects of cultivation and cropping practices upon earthworms. When forest vegetation is cleared, slashed, burned and land is tilled and cultivated, earthworm abundance, diversity and activity are reduced. Conversely, retaining trees in agroecosystems may maintain earthworm populations during the cropping phase.Here, we assessed the impact on earthworm species diversity and densities of crop cultivation in the understorey of timber plantations thinned to two tree densities and compared these with uncropped, undisturbed timber plantation controls. The plots were reassessed after two and a half years of fallow to see whether populations had recovered. The experiment was in central Cameroon.Seventeen earthworm species were recorded from Eudrilidae subfamilies Eudrilinae and Pareudrilinae, Ocnerodrilidae and Acanthodrilidae, most of which were endemics. This included two new species from two new genera from the sub-family Pareudrilinae, one new species from one new genus of Ocnerodrilidae, two new species of Dichogaster and one new species of Legonodrilus. Ten species were epigeic, six were endogeic and one was anecic.Generally, earthworm densities were lower in cropped plots than in the undisturbed plantation control. The most abundant species was a Legonodrilus sp. nov. with average densities of 49 individuals m⁻² in the crop phase and 80 ind. m⁻² in the fallow phase. By the fallow phase, densities in the low tree density (120 ind. m⁻²) were higher than in the high density (40 ind. m⁻²). The densities of the epigeic Acanthodrilidae were significantly reduced to 7 ind. m⁻² in the cropped plots compared with 42 ind. m⁻² in the control plots. The effects of cropping were thus species-specific and more work is required to identify which of these endemics are the ecosystem engineers in the system.     

Interactive effects of functionally different earthworm species on aggregation and incorporation and decomposition of newly added residue carbon

The interactive effects of two functionally different earthworm species (Aporrectodea caliginosa (endogeic species) and Lumbricus rubellus (epigeic species)) on the incorporation of fresh residue into large macroaggregates and formation of microaggregates within these large macroaggregates were investigated during a short-term laboratory experiment using ¹³C-labelled sorghum (Sorghum bicolor (L.) Moench) residues. Soil was collected from a long-term no-tillage agricultural field, crushed through a 250-μm sieve and incubated under laboratory conditions. The following earthworm treatments were applied: (i) soil+¹³C-labelled residue+A. caliginosa; (ii) soil+¹³C-labelled residue+L. rubellus; (iii) soil+¹³C-labelled residue+A. caliginosa+L. rubellus and; (iv) soil+¹³C-labelled residue. Two residue placement treatments (i.e. surface and incorporated) were superimposed on the earthworm treatments. Earthworms were added after 8 days of incubation. Aggregate size distribution and total C and ¹³C were measured after 22 days. Microaggregates, fine inter-microaggregate particulate organic matter (inter-POM) and intra-microaggregate POM (intra-POM) were isolated from macroaggregates. Earthworms had a greater stimulating effect on the formation of large macroaggregates (>2000 μm) and microaggregates within large macroaggregates when residue was incorporated in the soil, especially in the presence of A. caliginosa. When residue was placed on the surface, residue-derived intra-POM C was highest when L. rubellus was present and significantly lower in the presence of A. caliginosa. Residue-derived inter-POM C was highest when a mix of both species was present. These results indicate that earthworm species differentially affect incorporation of fresh organic matter into stable microaggregates within macroaggregates, and that interactive effects of earthworm species might have important consequences for the incorporation and protection of C inside of microaggregates within macroaggregates especially when residues are placed on the soil surface.     

A simple and effective method to keep earthworms confined to open-top mesocosms

Earthworms can have a profound effect on a myriad of soil physical, chemical and microbial parameters. To better understand their role in the soil, they are often studied under controlled conditions. However, a persistent problem in such controlled experiments is the ability of earthworms to escape from experimental units with open tops (e.g. for plant growth). Here, we tested whether adhesive hook tape applied to the inside of mesocosms is effective in confining them to their experimental units. A mesocosm study was set up with hook tape treatments (control, one layer, two layers), mesocosm material (polyvinylchloride – PVC, polypropylene – PP) and earthworm species (Lumbricus rubellus (Hoffmeister), Aporrectodea caliginosa (Savigny), Lumbricus terrestris (L.) + Aporrectodea longa (Ude)) as different factors to study the escape of earthworms during 24 h. In the treatments without hook tape, individuals of L. rubellus and A. caliginosa escaped, with highest escape rates (80%) for L. rubellus from the PP mesocosms, and lowest escape rates (20%) for A. caliginosa from the PVC mesocosms. When hook tape was applied, in either one or two layers, no individuals of those species escaped. The two anecic earthworm species, L. terrestris and A. longa did not escape from any mesocosms, irrespective of the presence of hook tape. As not a single earthworm escaped from the hook tape treatments, we conclude that applying hook tape is a simple, inexpensive and effective method to keep earthworms confined to experimental units.     

Nitrous oxide production in turfgrass systems: Effects of soil properties and grass clipping recycling

Soil N₂O emissions can affect global environments because N₂O is a potent greenhouse gas and ozone depletion substance. In the context of global warming, there is increasing concern over the emissions of N₂O from turfgrass systems. It is possible that management practices could be tailored to reduce emissions, but this would require a better understanding of factors controlling N₂O production. In the present study we evaluated the spatial variability of soil N₂O production and its correlation with soil physical, chemical and microbial properties. The impacts of grass clipping addition on soil N₂O production were also examined. Soil samples were collected from a chronosequence of three golf courses (10, 30, and 100-year-old) and incubated for 60 days at either 60% or 90% water filled-pore space (WFPS) with or without the addition of grass clippings or wheat straw. Both soil N₂O flux and soil inorganic N were measured periodically throughout the incubation. For unamended soils, cumulative soil N₂O production during the incubation ranged from 75 to 972 ng N g⁻¹ soil at 60% WFPS and from 76 to 8842 ng N g⁻¹ soil at 90% WFPS. Among all the soil physical, chemical and microbial properties examined, soil N₂O production showed the largest spatial variability with the coefficient of variation ~110% and 207% for 60% and 90% WFPS, respectively. At 60% WFPS, soil N₂O production was positively correlated with soil clay fraction (Pearson's r = 0.91, P < 0.01) and soil NH₄⁺–N (Pearson's r = 0.82, P < 0.01). At 90% WFPS, however, soil N₂O production appeared to be positively related to total soil C and N, but negatively related to soil pH. Addition of grass clippings and wheat straw did not consistently affect soil N₂O production across moisture treatments. Soil N₂O production at 60% WFPS was enhanced by the addition of grass clippings and unaffected by wheat straw (P < 0.05). In contrast, soil N₂O production at 90% WFPS was inhibited by the addition of wheat straw and little influenced by glass clippings (P < 0.05), except for soil samples with >2.5% organic C. Net N mineralization in soil samples with >2.5% organic C was similar between the two moisture regimes, suggesting that O₂ availability was greater than expected from 90% WFPS. Nonetheless, small and moderate changes in the percentage of clay fraction, soil organic matter content, and soil pH were found to be associated with large variations in soil N₂O production. Our study suggested that managing soil acidity via liming could substantially control soil N₂O production in turfgrass systems.     

Soil–atmosphere greenhouse gas exchange in a cool,temperate Eucalyptus delegatensis forest in south-eastern Australia

Forests are the largest C sink (vegetation and soil) in the terrestrial biosphere and may additionally provide an important soil methane (CH₄) sink, whilst producing little nitrous oxide (N₂O) when nutrients are tightly cycled. In this study, we determine the magnitude and spatial variation of soil–atmosphere N₂O, CH₄ and CO₂ exchange in a Eucalyptus delegatensis forest in New South Wales, Australia, and investigate how the magnitude of the fluxes depends on the presence of N₂-fixing tree species (Acacia dealbata), the proximity of creeks, and changing environmental conditions. Soil trace gas exchange was measured along replicated transects and in forest plots with and without presence of A. dealbata using static manual chambers and an automated trace gas measurement system for 2 weeks next to an eddy covariance tower measuring net ecosystem CO₂ exchange. CH₄ was taken up by the forest soil (?51.8 μg CH₄-C m^?2 h^?1) and was significantly correlated with relative saturation (S_r) of the soil. The soil within creek lines was a net CH₄ source (up to 33.5 μg CH₄-C m^?2 h^?1), whereas the wider forest soil was a CH₄ sink regardless of distance from the creek line. Soil N₂O emissions were small (<3.3 μg N₂O-N m^?2 h^?1) throughout the 2-week period, despite major rain and snowfall. Soil N₂O emissions only correlated with soil and air temperature. The presence of A. dealbata in the understorey had no influence on the magnitude of CH₄ uptake, N₂O emission or soil N parameters. N₂O production increased with increasing soil moisture (up to 50% S_r) in laboratory incubations and gross nitrification was negative or negligible as measured through ¹⁵N isotope pool dilution.The small N₂O emissions are probably due to the limited capacity for nitrification in this late successional forest soil with C:N ratios >20. Soil–atmosphere exchange of CO₂ was several orders of magnitude greater (88.8 mg CO₂-C m^?2 h^?1) than CH₄ and N₂O, and represented 43% of total ecosystem respiration. The forest was a net greenhouse gas sink (126.22 kg CO₂-equivalents ha^?1 d^?1) during the 2-week measurement period, of which soil CH₄ uptake contributed only 0.3% and N₂O emissions offset only 0.3%.     

Identifying earthworm's organic matter signatures by near infrared spectroscopy in different land-use systems in Tabasco,Mexico

In the state of Tabasco, South-eastern, Mexico, land-use changes such as the conversion of natural into agricultural systems, modify soil quality and the abundance of soil macrofauna, including earthworms. The aim of this study was to characterize by near-infrared spectroscopy (NIRS) the earthworms’ fingerprint in soil, in six sites including natural and agricultural ecosystems with low and high earthworm biomass and low and high earthworm diversity, in order to identify specific wavelengths that discriminate the presence/abundance of earthworm species and functional groups. The spectral region of 1860–1870 nm was significantly correlated with total earthworm density, particularly at one of the sites (Cedar polyculture; r = 0.8, p < 0.05). Earthworm biomass had a specific NIRS wavelength according to the earthworm species and feeding category: 1820 and1860–1870 nm wavelengths were significantly correlated with Polypheretima elongata (r² = 0.7, p < 0.05; mesohumic species) biomass and 2090 nm for biomass of all Lavellodrilus species (polyhumics). Two species had a much wider spectral range: L. bonampakensis and Dichogaster saliens (an epigeic worm; 1690–2300 nm, r² = 0.7, p < 0.05). Biomasses of Periscolex brachysistis and Diplotrema murchiei were not significantly correlated with any near infrared wavelength spectra analyzed. Combining a maximum of 4 species per wavelength, mesohumic earthworms had a wider wavelength spectrum than polyhumics. Therefore, earthworm species diversity, biomass and abundance are associated with soil quality (as measured by NIR spectra) and this relationship varies with species and ecological category. Sites with lower and higher earthworm diversity have lower and higher soil organic matter quality, respectively, as observed by the wider or narrower spectral range with which earthworm biomasses are correlated.     

Survival of Aporrectodea caliginosa and its effects on nutrient availability in biosolids amended soil

Few earthworms are present in production agricultural fields in the semi-arid plains of Colorado, where earthworm populations may be constrained by limited water and/or organic matter resources. We conducted a 12-week laboratory incubation study to determine the potential of a non-native endogeic earthworm (Aporrectodea caliginosa) to survive in a low-organic matter Colorado soil (1.4% organic C content), supplemented with or without biosolids, and to determine the effects of A. caliginosa on soil microbial biomass and soil nutrient availability. A factorial design with three main effects of A. caliginosa, biosolids addition, and time was used. Data was collected through destructively sampling at one, two, four, eight, and twelve weeks. During the 12-week study, 97.5% of the worms in the soil survived, and the survival of the earthworms was not significantly affected by the addition of biosolids. The addition of biosolids, however, did significantly reduce the gain in mass of the earthworms (8% mass gain compared to 18% in soil without biosolids). The presence of A. caliginosa significantly increased soil NH₄-N, and NO₃-N concentrations by 31% and 4%, respectively, which was less than the six fold increases in both soil NH₄-N, and NO₃-N concentrations supplied from biosolids. Microbial biomass carbon was not affected by A. caliginosa, but microbial biomass N was affected by an earthworm × biosolids interaction at week 1 and 12. We concluded that A. caliginosa can survive in a low-organic matter Colorado soil under optimal moisture content and that once established, A. caliginosa can provide modest increases in inorganic N availability to crops Colorado agroecosystems.     

The influence of soluble carbon and fertilizer nitrogen on nitric oxide and nitrous oxide emissions from two contrasting agricultural soils

Contradictory effects of simultaneous available organic C and N sources on nitrous oxide (N₂O), carbon dioxide (CO₂) and nitric oxide (NO) fluxes are reported in the literature. In order to clarify this controversy, laboratory experiments were conduced on two different soils, a semiarid arable soil from Spain (soil I, pH=7.5, 0.8%C) and a grassland soil from Scotland (soil II, pH=5.5, 4.1%C). Soils were incubated at two different moisture contents, at a water filled pore space (WFPS) of 90% and 40%. Ammonium sulphate, added at rates equivalent to 200 and 50 kg N ha^?1, stimulated N₂O and NO emissions in both soils. Under wet conditions (90% WFPS), at high and low rates of N additions, cumulative N₂O emissions increased by 250.7 and 8.1 ng N₂O–N g^?1 in comparison to the control, respectively, in soil I and by 472.2 and 2.1 ng N₂O–N g^?1, respectively, in soil II. NO emissions only significantly increased in soil I at the high N application rate with and without glucose addition and at both 40% and 90% WFPS. In both soils additions of glucose together with the high N application rate (200 kg N ha^?1) reduced cumulative N₂O and NO emissions by 94% and 55% in soil I, and by 46% and 66% in soil II, respectively. These differences can be explained by differences in soil properties, including pH, soil mineral N and total and dissolved organic carbon content. It is speculated that nitrifier denitrification was the main source of NO and N₂O in the C-poor Spanish soil, and coupled nitrification–denitrification in the C-rich Scottish soil.     

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.     

Removal of benzo (a) pyrene from soil using an endogeic earthworm Pontoscolex corethrurus ()

The endogeic earthworm Pontoscolex corethrurus (Müller, 1857) was the most abundant species (75%) in soil contaminated with hydrocarbons, mostly benzo(a)pyrene (BaP), in the state of Tabasco (Mexico). The earthworm P. corethrurus was tested for its capacity to remove 100 mg BaP kg⁻¹ from an Anthrosol soil (sterilized or not) and amended with legume Mucuna pruriens (L.) DC. var. utilis (Wall. ex Wight) Baker ex Burck (3%) or the grass Brachiaria humidicola (L.) DC (3%) (recently renamed as Urochloa humidicola (Rendle) Morrone & Zuloaga) in an aerobic incubation experiment. P. corethrurus removed 26.6 mg BaP kg⁻¹ from the sterilized soil and application of B. humidicola as feed increased this to 35.7 mg BaP kg⁻¹ and M. pruriens to 34.2 mg BaP kg⁻¹ after 112 days. The autochthonous microorganisms removed 9.1 mg BaP kg⁻¹ from the unsterilized soil and application of B. humidicola increased this to 18.0 mg BaP kg⁻¹ and M. pruriens to 11.2 mg BaP kg⁻¹. Adding P. corethrurus to the unsterilized soil accelerated the removal of BaP and 36.1 mg kg⁻¹ was dissipated from soil. It was found that the autochthonous microorganisms removed BaP from soil, but addition of P. corethrurus increased the dissipation 4-fold. The endogeic earthworm P. corethrurus can thus be used to remediate hydrocarbon-contaminated soils in tropical regions.     

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.     

N2O emissions from humid tropical agricultural soils: effects of soil moisture,texture and nitrogen availability

We studied soil moisture dynamics and nitrous oxide (N₂O) fluxes from agricultural soils in the humid tropics of Costa Rica. Using a split-plot design on two soils (clay, loam) we compared two crop types (annual, perennial) each unfertilized and fertilized. Both soils are of andic origin. Their properties include relatively low bulk density and high organic matter content, water retention capacity, and hydraulic conductivity. The top 2–3 cm of the soils consists of distinct small aggregates (dia. <0.5 cm). We measured a strong gradient of bulk density and moisture within the top 7 cm of the clay soil. Using automated sampling and analysis systems we measured N₂O emissions at 4.6 h intervals, meteorological variables, soil moisture, and temperature at 0.5 h intervals. Mean daily soil moisture content at 5 cm depth ranged from 46% water filled pore space (WFPS) on clay in April 1995 to near saturation on loam during a wet period in February 1996. On both soils the aggregated surface layer always remained unsaturated. Soils emitted N₂O throughout the year. Mean N₂O fluxes were 1.04±0.72 ng N₂O-N cm⁻² h⁻¹ (mean±standard deviation) from unfertilized loam under annual crops compared to 3.54±4.31 ng N₂O-N cm⁻² h⁻¹ from the fertilized plot (351 days measurement). Fertilization dominated the temporal variation of N₂O emissions. Generally fluxes peaked shortly after fertilization and were increased for up to 6 weeks (‘post fertilization flux’). Emissions continued at a lower rate (‘background flux’) after fertilization effects faded. Mean post-fertilization fluxes were 6.3±6.5 ng N₂O-N cm⁻² h⁻¹ while the background flux rate was 2.2±1.8 ng N₂O-N cm⁻² h⁻¹. Soil moisture dynamics affected N₂O emissions. Post fertilization fluxes were highest from wet soils; fluxes from relatively dry soils increased only after rain events. N₂O emissions were weakly affected by soil moisture during phases of low N availability. Statistical modeling confirmed N availability and soil moisture as the major controls on N₂O flux. Our data suggest that small-scale differences in soil structure and moisture content cause very different biogeochemical environments within the top 7 cm of soils, which is important for net N₂O fluxes from soils.