Relationships between earthworm communities and burrow numbers under different land use systems

This study addresses the influence of three different land use systems (continuous maize, pasture/maize rotation, permanent pasture) on the relationships between earthworm populations and the number of earthworm burrows quantified in a soil profile. Quantified burrows were limited to those observable by the naked eye (i.e. >2 mm in diameter) and enumerated earthworms were limited to those which could have created the observable burrows (i.e. >0.3 g).The results were combined with data from the literature coming from different geographical regions. This study showed that earthworm abundance decreased with the increasing land management intensity (maize crop vs. pasture), while the number of burrows could be higher or similar under maize compared to pasture. Under maize, despite lower earthworm abundance and the annual destruction of the burrows by tillage, the number of burrows was almost as high as under pasture. This absence of a relationship between burrow numbers and earthworm abundance was observed in the soil profile and for each layer of the profile for each land use system. Furthermore, the burrow number/m⁻² per earthworm strongly varied depending on land use and was far higher under maize when compared to pasture (74 vs. 7). Therefore, a power-law type relationship was clearly established between burrow number/m⁻² per earthworm and earthworm abundance. This power type relationship was also observed when including data from the literature although it followed a different mathematical model. These results were explained by (i) increased earthworm burrowing activity (i.e. an increase in the number of burrows produced by each earthworm) under maize, and (ii) the dynamics of burrow number under pasture (i.e. decreased burrowing activity and burrow destruction process); both results of food accessibility combined with inter-individual competition. The results of the study suggest that farmers should not use the number of pores as an indicator of earthworm abundance, but as an indicator of earthworm activity, which could be integrated in an indicator of soil quality.     

Relationships between earthworm communities and burrow numbers under different land use systems

This study addresses the influence of three different land use systems (continuous maize, pasture/maize rotation, permanent pasture) on the relationships between earthworm populations and the number of earthworm burrows quantified in a soil profile. Quantified burrows were limited to those observable by the naked eye (i.e. >2 mm in diameter) and enumerated earthworms were limited to those which could have created the observable burrows (i.e. >0.3 g).The results were combined with data from the literature coming from different geographical regions. This study showed that earthworm abundance decreased with the increasing land management intensity (maize crop vs. pasture), while the number of burrows could be higher or similar under maize compared to pasture. Under maize, despite lower earthworm abundance and the annual destruction of the burrows by tillage, the number of burrows was almost as high as under pasture. This absence of a relationship between burrow numbers and earthworm abundance was observed in the soil profile and for each layer of the profile for each land use system. Furthermore, the burrow number/m^?2 per earthworm strongly varied depending on land use and was far higher under maize when compared to pasture (74 vs. 7). Therefore, a power-law type relationship was clearly established between burrow number/m^?2 per earthworm and earthworm abundance. This power type relationship was also observed when including data from the literature although it followed a different mathematical model. These results were explained by (i) increased earthworm burrowing activity (i.e. an increase in the number of burrows produced by each earthworm) under maize, and (ii) the dynamics of burrow number under pasture (i.e. decreased burrowing activity and burrow destruction process); both results of food accessibility combined with inter-individual competition. The results of the study suggest that farmers should not use the number of pores as an indicator of earthworm abundance, but as an indicator of earthworm activity, which could be integrated in an indicator of soil quality.     

Diversity and abundance of earthworms across an agricultural land-use intensity gradient

Understanding how communities of important soil invertebrates vary with land use may lead to the development of more sustainable land-use strategies. We assessed the abundance and species composition of earthworm communities across six replicated long-term experimental ecosystems that span a gradient in agricultural land-use intensity. The experimental systems include a conventional row-crop agricultural system, two lower-intensity row-crop systems (no-till and tilled organic input), an early successional old-field system, a 40–60 years old coniferous forest plantation, and an old-growth deciduous forest system. Earthworm populations varied among systems; they were lowest in the most intensively managed row-crop system (107 m⁻²) and coniferous forest (160 m⁻²); intermediate in the old-field (273 m⁻²), no-till (328 m⁻²) and tilled organic (344 m⁻²) cropping systems; and highest in the old-growth deciduous forest system (701 m⁻²). Juvenile Aporrectodea species were the most common earthworms encountered in intensively managed systems; other species made up a larger proportion of the community in less intensively managed systems. Earthworm community biomass and species richness also varied and were lowest in the conventional row-crop system and greatest in the old-growth forest system. These results suggest that both land-use intensity and land-use type are strong drivers of the abundance and composition of earthworm communities in agricultural ecosystems.     

The effect of tillage type and cropping system on earthworm communities, macroporosity and water infiltration

To test the assumption that changes to earthworm communities subsequently affect macroporosity and then soil water infiltration, we carried out a 3 year study of the earthworm communities in a experimental site having six experimental treatments: 2 tillage management systems and 3 cropping systems. The tillage management was either conventional (CT; annual mouldboard ploughing up to −30 cm depth) or reduced (RT; rotary harrow up to −7 cm depth). The 3 cropping systems were established to obtain a wide range of soil compaction intensities depending on the crop rotations and the rules of decision making. In the spring of 2005, the impact of these different treatments on earthworm induced macroporosity and water infiltration was studied. During the 3 years of observation, tillage management had a significant effect on bulk density (1.27 in CT and 1.49 mg m⁻³ in RT) whereas cropping system had a significant effect on bulk density in RT plots only. Tillage management did not significantly affect earthworm abundance but significantly influenced the ecological type of earthworms found in each plot (anecic were more abundant in RT). On the contrary cropping system did have a significant negative effect on earthworm abundance (104 and 129 ind. m⁻² in the less and most compacted plots, respectively). Significantly higher numbers of Aporrectodea giardi and lower numbers of Aporrectodea caliginosa were found in the most compacted plots. CT affected all classes of porosity leading to a significant decrease in the number of pores and their continuity. Only larger pores, with a diameter superior to 6 mm, however, were adversely affected by soil compaction. Tillage management did not change water infiltration, probably because the increase in macroporosity in RT plots was offset by a significant increase in soil bulk density. However, cropping system had a significant effect on water infiltration (119 vs 79 mm h⁻¹ in the less and most compacted plots, respectively). In RT plots, a significant correlation was observed between larger macropores (diameter > 6 mm) and water infiltration illustrating the potential positive effect of earthworms in these plots.     

Earthworms of different functional groups affect the fate of the Bt-toxin Cry1Ab from transgenic maize in soil

The fate of the insecticidal Cry1Ab protein from crop residues (leaves and roots) of the transgenic maize variety MON810 was studied in the presence and absence of two earthworm species (Lumbricus terrestris, Aporrectodea caliginosa; separate incubations) in soil microcosms. The recombinant Cry1Ab protein was quantified using a highly sensitive ELISA. Control microcosms received corresponding non-transgenic plant material. All earthworms survived in the microcosms over a period of 5 weeks, irrespective of whether they received MON810 or non-transgenic plant material. Weight loss was observed for both earthworm species, independent of the plant material or transgenic modification. A strong decline of immunoreactive Cry1Ab in plant residues (mean initial concentration approx. 5000 ng g⁻¹) of MON810 was observed in all treatments, but in microcosms with earthworms this decline was significantly higher with less than 10% of the initial Cry1Ab concentration remaining after 5 weeks. Cry1Ab concentrations in casts were only 0.1% of those found in remaining plant material of the respective microcosms. No immunoreactive Cry1Ab proteins were found in earthworm tissues (threshold of detection: 0.58 ng g⁻¹ fresh weight). No further decline was found for Cry1Ab concentrations in casts of A. caliginosa during a subsequent period of 3 months of incubation in bulk soil (<0.1 ng g⁻¹) after removal of the earthworms from the microcosms, while in casts of L. terrestris the concentration decreased from 0.4 to below 0.1 ng g⁻¹. In conclusion, this study demonstrates that earthworms enhance the decline of immunoreactive Cry1Ab proteins from maize residues.     

Earthworms change the abundance and community structure of nematodes and protozoa in a maize residue amended rice-wheat rotation agro-ecosystem

The influence of earthworms on nematodes and protozoan communities was determined during the wheat phase of a six year rice-wheat rotation agro-ecosystem. Experimental plots in the rotation had five treatments, i.e. incorporation or mulching of maize residues with or without added earthworms and a control. The addition of maize residues to soil strongly affected the abundance and community structure of nematodes and protozoa in the absence of earthworms. The presence of earthworms gave significantly lower total nematode numbers at all soil depths following maize residue incorporation than the same treatment without earthworms, and also gave lower (although not significantly) total nematode numbers in the upper soil layer following maize residue mulching than the same treatment without earthworms. This was mainly due to a significant decrease in bacterial-feeding nematode numbers. Earthworms also strongly affected the distribution of the number of total nematodes and two trophic groups (bacterial and plant feeders) with soil depth. In the presence of earthworms, total protozoan and flagellate numbers significantly increased at all soil depths following both incorporation and mulching of maize residues, while numbers of amoebae increased only when maize residues were mulched. Additionally, in earthworm casts total nematode numbers (mainly bacterial and fungal feeders) were significantly higher, whereas total protozoa numbers (mainly flagellates and amoebae) were significantly lower than that in soil from 0 to 5 cm layer.These results indicated that earthworm activity could affect the abundance and community structure of microfauna, and change their distribution between soil layers and cast material, depending on the mode of application of organic residues.     

Effects of earthworms on soil enzyme activity in an organic residue amended rice–wheat rotation agro-ecosystem

The effect of earthworms on soil hydrolases (protease, urease, invertase, and alkaline phosphatase) and dehydrogenase activities was investigated in maize residue amended rice–wheat rotation agro-ecosystem. Experimental plots in the rotation had five treatments, i.e. incorporation or mulching of maize residues with or without added earthworms and an untreated control. The application of maize residues to soil without earthworms significantly enhanced the five soil enzyme activities compared with the control treatment during rice and wheat cultivation. The presence of earthworms further significantly enhanced protease activity in the soils with both incorporated and mulched maize residues during two cultivation seasons, but only significantly increased alkaline phosphatase activity in the soil with incorporated maize residue during the rice cultivation season. Invertase activity was significantly enhanced by the presence of earthworms in the soil with maize residue incorporation during two cultivation seasons. There were no changes in dehydrogenase activity when earthworms were present. Additionally, the five enzyme activities in earthworm casts were significant higher than those in the surrounding soil, especially dehydrogenase and invertase activities. Whatever the treatment, the values obtained for the enzyme activities in both soil and casts, except for dehydrogenase activity in earthworm casts, were significantly higher under wheat than those in rice-cultivated soil. These results indicate that the presence of earthworms strongly affected soil enzyme activities, depending on the method of organic residue application, and the enhanced enzyme activities of earthworm casts probably contributed to the surrounding soil enzyme activities.     

Earthworm burrowing in laboratory microcosms as influenced by soil temperature and moisture

Earthworm burrows contribute to soil macroporosity and support diverse microbial communities. It is not well known how fluctuations in soil temperature and moisture affect the burrowing activities of earthworms. The objective of this experiment was to evaluate the maximum depth and length of burrows created by the endogeic earthworm Aporrectodea caliginosa (Savigny) and the anecic earthworm Lumbricus terrestris L. for a range of temperatures (5–20 °C) and soil water potentials (−5 and −11 kPa). The laboratory microcosm was a plexiglass chamber (45 cm high, 45 cm wide) containing 0.14 m² of pre-moistened soil and litter, designed to house a single earthworm for 7 days. Earthworm mass, surface casting and burrowing activities were affected significantly by soil temperature, moisture and the temperature×moisture interaction. Burrow length and maximum burrow depth increased with increasing temperature, but there was less burrowing in wetter soil (−5 kPa) than drier soil (−11 kPa). Weight gain and surface casting, however, were greater in soil at −5 kPa than −11 kPa. Our results suggest more intensive feeding and limited burrowing in wetter soil than drier soil. Earthworms inhabiting the non-compacted, drier soil may have pushed aside particles without ingesting them to create burrows. The result was that earthworms explored a larger volume of soil, deeper in the chamber, when the soil was drier. How these burrowing activities may affect the community structure and activity of soil microorganisms and microfauna in the drilosphere remains to be determined.     

Growth rates of Aporrectodea caliginosa (Oligochaetae: Lumbricidae) as influenced by soil temperature and moisture in disturbed and undisturbed soil columns

Earthworm growth is affected by fluctuations in soil temperature and moisture and hence, may be used as an indicator of earthworm activity under field conditions. There is no standard methodology for measuring earthworm growth and results obtained in the laboratory with a variety of food sources, soil quantities and container shapes cannot easily be compared or used to estimate earthworm growth in the field. The objective of this experiment was to determine growth rates of the endogeic earthworm Aporrectodea caliginosa (Savigny) over a range of temperatures (5–20 °C) and soil water potentials (−5 to−54 kPa) in disturbed and undisturbed soil columns in the laboratory. We used PVC cores (6 cm diameter, 15 cm height) containing undisturbed and disturbed soil, and 1 l cylindrical pots (11 cm diameter, 14 cm height) with disturbed soil. All containers contained about 500 g of moist soil. The growth rates of juvenile A. caliginosa were determined after 14–28 days. The instantaneous growth rate (IGR) was affected significantly by soil moisture, temperature, and the temperature×moisture interaction, ranging from −0.092 to 0.037 d⁻¹. Optimum growth conditions for A. caliginosa were at 20 °C and −5 kPa water potential, and they lost weight when the soil water potential was −54 kPa for all temperatures and also when the temperature was 5 °C for all water potentials. Growth rates were significantly greater in pots than in cores, but the growth rates of earthworms in cores with undisturbed or disturbed soil did not differ significantly. The feeding and burrowing habits of earthworms should be considered when choosing the container for growth experiments in order to improve our ability to extrapolate earthworm growth rates from the laboratory to the field.     

Contrasting response to cropping of populations of earthworms and predacious nematodes in four soils

The activities of many soil animals make a positive contribution to soil processes and they should be considered for inclusion in indices of ‘soil quality'. To assess the potential use of nematodes and earthworms as indicators, the relationships between populations of earthworms (Lumbricidae), total number of nematodes and predacious nematodes (Mononchoidea) and six soil physical factors, soil carbon and pH were investigated in four New Zealand soils. In each, soil treatments ranged from 5–90 year pastures to cropping with maize or barley for 11–29 years. With increasing cultivation, trends in bulk density, total porosity, aggregate stability and concentration of total carbon were similar in all four soils. In Manawatu (Dystric Fluventic Eutrochrept) and Kairanga (Typic Endoaquept) soils earthworm populations were negligible under continuous cropping while Mononchoidea were abundant (11 600 and 34 100 m⁻²). In contrast, in Moutoa (Fluvaquentic Endoaquoll) and Wakanui (Aquic Ustochrept) soils earthworms persisted under cultivation, while Mononchoidea were less abundant (300 and 2500 m⁻²). At these two latter sites, aggregate stability was higher (1.14 and 0.92 mm mean weight diameter (MWD)) than in Manawatu and Kairanga soils (0.38 and 0.35 mm MWD). These relationships between aggregate stability, earthworm abundance and predacious nematodes show not only that some potential indicators may have a local rather than national application, but also that there are important interactions between soil physical properties and soil fauna which require further investigation.     

Earthworm composition,diversity and biomass under three land use systems in south-eastern Australia

In south-eastern Australia, strips of planted native trees and shrubs (shelterbelts) are frequently established to restore ecosystem services altered by agriculture. Despite their wide use, little is known about the effects of establishing shelterbelts on soil macro invertebrates, especially earthworms, which are of major importance in soil processes. We assessed earthworm composition, diversity and biomass in three land use systems: native shelterbelts dominated by Acacia and Eucalyptus species, agricultural pastures and native remnant woodland fragments dominated by Eucalyptus blakelyi and/or Eucalyptus melliodora. Earthworm communities differed significantly among systems, with abundance, biomass and diversity greatest under pasture. Within shelterbelts we saw a shift from high earthworm biomass and density to low with increasing time after establishment. Soil edaphic variables did not correlate strongly with earthworm biomass or density, but were correlated with earthworm community composition. Overall the introduction of native woody vegetation was associated with a decline in density and biomass of earthworms, including a decrease in the relative abundance of exotic species. As such shelterbelts can be used to promote native earthworm relative abundance, which may be important for local diversity, soil function and landscape connectivity.     

Role of earthworms in regenerating soil structure after compaction in reduced tillage systems

New non-tillage or reduced tillage agricultural practises are being increasingly adopted but generally result in higher soil compaction. Due to their recognised physical influence mainly through burrow creation, it is often claimed that earthworm activity could alleviate soil compaction in these systems. To put this assumption to the test, an experimental compaction event was carried out on one plot of arable land. The abundance and biomass of earthworms were evaluated in compacted (under wheel tracks) and non-compacted (between wheel tracks) zones, seven times over a two-year period. In addition, the functional consequences of earthworm activity, defined by burrow abundance assessed in 2D and 3D and water infiltration, were measured three times over the same period. The short-term (less than three months) effects of the compaction were clear: soil bulk density increased from 1.46 to 1.57 g cm⁻³, the abundance and biomass of earthworms were greatly reduced (−40% and −70% respectively) and the number and continuity of macroporosity were lower under wheel tracks at least until a depth of 30 cm. After these initial detrimental effects, we observed a rapid recovery of earthworm populations with no statistical difference between compacted and control zones more than three months after the compaction. However, the recovery of soil functional properties linked to earthworm activity, macroporosity and water infiltration, was much slower and took between 12 and 24 months. Despite these modifications, there were no significant changes in soil bulk density with time during the two-year period. This study demonstrates that earthworms are important actors in the regeneration of compacted soil. Although the complete regeneration of compacted soil by earthworms is a slow process, agricultural practises that promote earthworm density and activity should be encouraged in reduced or minimum tillage systems.     

Impact of earthworms on the diversity of microarthropods in a vertisol (Martinique)

In a study of a 15-year-old pasture in Martinique (French West Indies), abundance and organization of microarthropod communities were correlated with the spatial distribution of the earthworm Polypheretima elongata (Megascolecidae). In patches of high earthworm density (133 individuals m^–2), microarthropod density was significantly higher (80000 individuals m^–2) than in patches with few earthworms (31 worms m^–2 and 49000 microarthropods m^{– 2}). The diversity of microarthropod communities followed a similar pattern, the Shannon index for Collembola communities being, respectively, 3.12 and 1.82 in and outside earthworm patches. These results suggest that mesofauna abundance and diversity might be at least partly determined by the activity of larger invertebrates, as a result of the dramatic effects that the latter group exerts upon soil structure, pore distribution and food resources. Received: 7 February 1997     

Earthworm populations in soils disturbed by trampling

Heavy trampling by livestock in pasture entrances considerably reduced earthworm density and biomass, as did behicular compaction on a human path through pasture. Surface species were most severely affected, whilst the deep burrowing Allolobophora longa proved particularly resistant. All species showed a downward shift in vertical distribution. A laboratory experiment demonstrated the great effectiveness of A. longa in ameliorating the effects of soil compaction. Earthworm population changes on trampling are likely to facilitate recovery of soil drainage once trampling ceases.     

Organic resources and earthworms affect phosphorus availability to sorghum after phosphate rock addition in semi-arid West Africa

A field experiment was laid out in Burkina Faso (West Africa) on an Eutric Cambisol to investigate the interaction of organic resource quality and phosphate rock on crop yield and to assess the contribution of earthworms (Millsonia inermis Michaelsen) to P availability after phosphate rock application. Organic resources of different quality were applied at a dose equivalent to 40 kg N ha^–1 with or without phosphate rock from Kodjari (Burkina Faso) at a dose equivalent to 25 kg P ha^–1, and were compared with control and single phosphate rock treatments in a factorial complete block design with four replicates. Sorghum (Sorghum bicolor L. Moench) variety SARIASSO 14 was grown. Sheep dung had the highest impact on earthworm casting intensity followed by maize straw. Combining organic resources with phosphate rock reduced earthworm casting activities compared to a single application of organic resources or phosphate rock. Addition of phosphate rock to maize straw reduced P availability in earthworm casts whereas combining sheep dung or compost with phosphate rock increased P availability. The contribution of earthworms to Kodjari phosphate rock solubilisation mainly occurred through their casts, as the available P content of casts was 4 times higher than that of the surrounding soil.     

Earthworm populations under tropical maize cultivation: the effect of mulching with velvetbean

Earthworm populations were studied in three tropical agroecosystems of southern Mexico: improved maize with a Mucuna pruriens cover crop (MM), continuous conventional maize (CM) and pastures (P). Three replicates and six monoliths were sampled in each agroecosystem. Three earthworm species were found, two native (Balanteodrilus pearsei, Larsonidrilus orbiculatus) and one exotic (Polypheretima elongata). In all systems, the dominant species was B. pearsei, with negligible presence of the exotic species in MM and P plots. Total abundance was significantly higher in MM than in CM; and earthworm biomass was also higher in MM than in CM and P. Juveniles of both native species dominated, mainly concentrated in the top 20 cm of soil. B. pearsei and L. orbiculatus displayed different preferences (within each agroecosystem) for soil organic matter, N and temperature. Further experiments are required to investigate whether mulching with M. pruriens results in an increased earthworm abundance and biomass through a N-improvement effect or as a result of microclimatic changes and to study the extent to which earthworms and M. pruriens are synergistic in enhancing maize growth.     

Decomposition and mineralization of energy crop residues governed by earthworms

Energy crops are increasingly cultivated in agricultural management systems world-wide. A substitution of food crops (e.g. cereals) by energy crops may generally alter the biological activity and litter decomposition in soil due to their varying structural and chemical composition and subsequently modify soil functioning. A soil microcosm experiment was performed to assess the decomposition and microbial mineralization of different energy crop residues in soil compared to a food crop, with or without earthworms. Residues of the energy crops winter rape (Brassica napus), maize (Zea mays), miscanthus (Miscanthus giganteus) and the food crop oat (Avena sativa) were each provided as food source for a mixed earthworm population, each consisting of one individual of Lumbricus terrestris, Aporrectodea caliginosa, and Octolasion tyrtaeum. After 6 weeks, the rate of litter loss from the soil surface, earthworm biomass, microbial biomass-C and -N, microbial activity, and enzyme activities were determined. The results emphasized, that litter loss and microbial parameters were predominantly promoted by earthworms and were additionally influenced by the varying structural and chemical composition of the different litter. Litter decay by earthworms was highest in N-rich maize litter treatment (C-N ratio 34.8) and lowest in the case of miscanthus litter (C-N ratio 134.4). As a consequence, the microbial biomass and basal respiration in soils with maize litter were higher, relative to other litter types. MBC-MBN ratio in soil increased when earthworms were present, indicating N competition between earthworms and microorganisms. Furthermore, enzyme activities responded in different ways on the varying types of litter and earthworm activity. Enzymes involved in the N-cycle decreased and those involved in the C-cycle tended to increase in the presence of earthworms, when litter with high C-N ratio was provided as a food source. Especially in the miscanthus treatments, less N might remain for enzymatic degradation, indicating that N competition between earthworms and microorganisms may vary between different litter types. Especially, an expansion of miscanthus in agricultural management systems might result in a reduced microbial activity and a higher N deficit for microorganisms in soil.     

Effects of traditional and biodynamic farmyard manure amendment on yields,soil chemical,biochemical and biological properties in a long-term field experiment

We studied the effects of applications of traditionally composted farmyard manure (FYM) and two types of biodynamically composted FYM over 9 years on soil chemical properties, microbial biomass and respiration, dehydrogenase and saccharase activities, decomposition rates and root production under grass-clover, activity and biomass of earthworms under wheat, and yields in a grass-clover, potatoes, winter wheat, field beans, spring wheat, winter rye crop rotation. The experiment was conducted near Bonn, on a Fluvisol using a randomised complete block design (n=6). Our results showed that plots which received either prepared or non-prepared FYM (30 Mg ha^–1 year^–1) had significantly increased soil pH, P and K concentrations, microbial biomass, dehydrogenase activity, decomposition (cotton strips), earthworm cast production and altered earthworm community composition than plots without FYM application. Application of FYM did not affect the soil C/N ratio, root length density, saccharase activity, microbial basal respiration, metabolic quotient and crop yields. The biodynamic preparation of FYM with fermented residues of six plant species (6 g Mg^–1 FYM) significantly decreased soil microbial basal respiration and metabolic quotient compared to non-prepared FYM or FYM prepared with only Achillea. The biodynamic preparation did not affect soil microbial biomass, dehydrogenase activity and decomposition during 62 days. However, after 100 days, decomposition was significantly faster in plots which received completely prepared FYM than in plots which received no FYM, FYM without preparations or FYM with the Achillea preparation. Furthermore, the application of completely prepared FYM led to significantly higher biomass and abundance of endogeic or anecic earthworms than in plots where non-prepared FYM was applied.     

Population dynamics of the earthworm Amynthas alexandri (Annelida: Megascolecidae) in a Kumaun Himalayan pasture soil

Ecological studies on earthworms were conducted in a Kumaun Himalayan pasture soil. The C:N ratio in the soil declined with increasing depth. A combination of hand-sorting and formalin application was used to sample the earthworms. Three species, Amynthas alexandri, A. diffringens (Megascolecidae), and Eisenia fetida (Lumbricidae) were found. Of the 13310 individuals collected, 99.9% were A. alexandri. The maximum density (138.8 m^-2) and biomass (25.2 g m^-2) were recorded in the wet season. More than 60% of the total earthworm numbers and biomass were recorded at 0–10 cm in depth. The mean yearly ratio of clitellate to aclitellate worms was 1:7.3.