Recovery of enchytraeid populations after severe drought events

The potential impact of changes in precipitation patterns associated with climate changes was investigated in Enchytraeidae (Oligochaeta) in a Danish heathland. The amount of precipitation was manipulated during spring and summer in an experimental field site in order to reveal effects of three different drought regimes: weak drought (WD), medium drought (MD) and high drought (HD). Enchytraeids were sampled every six to eight weeks (0–9 cm depth) for more than the eight months and soil water potential (SWP) and soil water content (SWC) was measured on a regular basis for five months.The enchytraeid communities were generally reduced due to a natural drought spell. The HD treatment significantly reduced the moisture level of the soil further with SWP below −15 bar (5 and 10 cm depth) and SWC around 5% (v/v) for more than two months. As a result almost no enchytraeids were found in HD plots after two months with high drought stress. Nevertheless, the HD-treated enchytraeid communities recovered within two months, as there was no significant difference in biomass and density of the different treatments at that time. During periods with extreme low SWP enchytraeids were practically absent in the top soil (0–3 cm), but a few animals were found in 3–6 cm. During this period SWP was around −15 bar even in 20 cm depth, indicating that active stages could not have survived. Thus, we suggest that the species present must be dependent on a drought tolerant stage, as vertical migration could not have supported the observed recovery.SWP and SWC were both significantly correlated with the total density and biomass of enchytraeids. However, density was better correlated with the SWP and SWC compared to biomass, which can be due to hatching of cocoons and increased fragmentation rate.     

Population dynamics of Enchytraeidae (Oligochaeta) in different agricultural systems

Organic matter and abiotic conditions seem to influence distribution patterns of Enchytraeidae. In this study effects of changes in management practices on population dynamics of enchytraeids were determined. At two sites (in Athens and Griffin, GA) parts of a fescue (FE) field were converted into conventional tillage (CT) and no-tillage (NT) plots and changes in densities and depth distribution of enchytraeids were followed for 3 years. A site at Watkinsville, containing various soil textures and characterised by very low organic carbon content, which was converted into no-tillage 4 years earlier, was also sampled. Significant reductions in enchytraeid densities, after conversion of fescue into CT, were only found at Griffin. The management practices affected the vertical distribution of enchytraeids. In fescue and NT more enchytraeids were found in the 0–5cm than in the 5–15cm layer. In conventional tillage fields enchytraeids were more evenly distributed over the profile or more abundant in the 5–15cm layer. Management also affected the timing of population dynamics in the different plots. At two sample dates high abundances were found in CT plots only, not in any of the other plots. Enchytraeids were larger at Athens than at Griffin and Watkinsville and contained more soil particles in their gut. At Athens enchytraeids will presumably contribute more to the development of soil structure than at the other sites. We conclude that management affects vertical enchytraeid distributions in soil and changes the timing of population dynamics. Received: 7 January 1996     

Do enchytraeid worms and habitat corridors facilitate the colonisation of habitat patches by soil microbes?

Due to their high abundance and ubiquitous existence, microbes are considered to be efficient colonisers of newly established habitats. To shed light on the dispersal mechanisms of soil microbes, a controlled microcosm experiment was established. In these microcosms, the dispersal of microbes from a source humus patch to originally sterile humus patches (embedded in a mineral soil matrix) was followed for 16 months, applying 16S and 18S ribosomal DNA-based PCR-DGGE molecular methods. Specifically, the role of enchytraeid worms and habitat (humus) corridors as possible facilitators of microbe dispersal was studied. The results showed that enchytraeid worms function efficiently as vectors for horizontal dispersal of saprophytic fungi in soil. Some of the fungi also proved to disperse through the corridors by vegetative growth, although this dispersal was inefficient as compared to dispersal with the enchytraeids. Virtually no saprophytic fungi were able to disperse through the mineral soil matrix in the absence of both enchytraeid worms and corridors. Unlike soil fungi, the dispersal of soil bacteria was not affected by any of the studied factors. The results of the present experiment provide direct evidence of the crucial role of soil fauna in aiding the horizontal dispersal of soil fungi. The role of enchytraeids as a functionally important species in boreal forest soils is further emphasized, since bringing microbes into contact with new resources is likely to enhance the rate of decomposition in soils.     

Influence of warming and enchytraeid activities on soil CO2 and CH4 fluxes

To determine the sum of ‘direct’ and ‘indirect’ effects of climatic change on enchytraeid activity and C fluxes from an organic soil we assessed the influence of temperature (4, 10 and 15 °C incubations) on enchytraeid populations and soil CO₂ and CH₄ fluxes over 116 days. Moisture was maintained at 60% of soil dry weight during the experimental period and measurements of enchytraeid biomass and numbers, and CO₂ and CH₄ fluxes were made after 3, 16, 33, 44, 65, 86 and 116 days. Enchytraeid population numbers and biomass increased in all temperature treatments with the greatest increase produced at 15 °C (to over threefold initial values by day 86). Results also showed that enchytraeid activity increased CO₂ fluxes by 10.7±4.5, 3.4±4.0 and 26.8±2.6% in 4, 10 and 15 °C treatments, respectively, with the greatest CO₂ production observed at 15 °C for the entire 116 day incubation period (P<0.05). The soil respiratory quotient analyses at lower temperatures (i.e. 4-10 °C) gave a Q₁₀ of 1.7 and 1.9 with and without enchytraeids, respectively. At temperatures above 10 °C (i.e. 10-15 °C) Q₁₀ significantly increased (P<0.01) and was 25% greater in the presence of enchytraeids (Q₁₀=3.4) than without (Q₁₀=2.6). In contrast to CO₂ production, no significant relationships were observed between net CH₄ fluxes and temperature and only time showed a significant effect on CH₄ production (P<0.01).Total soil CO₂ production was positively linked with enchytraeid biomass and mean soil CO₂-C production was 77.01±6.05 CO₂-C μg mg enchytraeid tissue⁻¹ day⁻¹ irrespective of temperature treatment. This positive relationship was used to build a two step regression model to estimate the effects of temperature on enchytraeid biomass and soil CO₂ respiration in the field. Predictions of potential CO₂ production were made using enchytraeid biomass data obtained in the field from two upland grassland sites (Sourhope and Great Dun Fell at the Moor House Nature Reserve, both in the UK). The findings of this work suggest that a 5 °C increase in atmospheric temperature above mean ambient temperature could have the potential to produce a significant increase in enchytraeid biomass resulting in a near twofold increase in soil CO₂ release from both soil types. The interaction between temperature and soil biology will clearly be an important determinant of soil respiration responses to global warming.     

Effects of artificial acidification and liming on the activity of digestive enzymes in Cognettia sphagnetorum (Vejdovsky´, 1878) (Annelida, Enchytraeidae)

The effects of simulated acid rain and acidification, combined with liming, on amylolytic, laminarinolytic and xylanolytic activity in whole body homogenates of enchytraeids Cognettia sphagnetorum were studied under field conditions. Simulated acid rain (pH 2.5) and simulated acid rain with subsequent liming (CaCO₃) were applied to experimental plots in a mixed forest soil. The pH of the soil was lowered by acid treatment (4.3), while the pH increased after liming (6.3) in comparison with the control (4.5). Acidification of soil caused a decrease in enchytraeid body mass and amylolytic activity. In acidified plots after liming, amylolytic activity and laminarinolytic activity increased, while live body mass decreased. The enzymatic activity of enchytraeids depended on season and also indirectly on individual mean mass. Received: 12 February 1996     

Earthworm invasion alters enchytraeid community composition and individual biomass in northern hardwood forests of North America

European earthworms are invading many ecosystems worldwide and fundamentally transform habitats by acting as dominant ecosystem engineers. However, there is little knowledge of the consequences of earthworm invasion on the composition and diversity of native soil organisms. Particularly functionally similar groups, such as enchytraeids (Annelida: Enchytraeidae), may be affected through changes in the chemical and physical properties of the soil, but also due to competition for resources. In 2010–2011, we studied the impact of earthworm invasion on enchytraeids at two sites in the northern hardwood forests of North America: one site within the Chippewa National Forest in northern Minnesota and one site in the Chequamegon-Nicolet National Forest in northern Wisconsin, USA. At each site, three plots were sampled along a transect, representing (1) a non-invaded or very slightly invaded area, (2) the leading edge of earthworm invasion and (3) a heavily invaded area with an established population of the anecic earthworm Lumbricus terrestris (among other species). In total, 29 enchytraeid (morpho)species were identified (some yet to be formally described, several first or second records for the continent); of those 24 occurred at the Minnesota site and 17 at the Wisconsin site. The structure of enchytraeid assemblages differed significantly among the three invasion stages, although this was not equally pronounced at the two sites. Each stage was characterized by one or several indicator species. Mean enchytraeid densities (10,700–30,400 individuals/m²) did not differ significantly among the invasion stages, but were lowest at the leading edge of earthworm invasion at both sites. In the heavily invaded plot at the Minnesota site, the mean enchytraeid density and biomass in L. terrestris middens were significantly higher than in soil in-between the middens. This was due to a pronounced effect of L. terrestris middens in the uppermost 3 cm of soil. Differences in biomass among earthworm invasion stages were most apparent for mean individual biomass. This was significantly higher in the heavily invaded area than at the leading edge or in the non-invaded area at the Minnesota site. Compositional changes of the enchytraeid assemblage are likely to result in changes in the functioning of soil foods webs. Our results suggest that earthworm invasions can cause a loss of native species in soil, including heretofore unknown ones, that might go unnoticed.     

Increased frequency of drought reduces species richness of enchytraeid communities in both wet and dry heathland soils

Studies of biological responses in the terrestrial environment to rapid changes in climate have mostly been concerned with above-ground biota, whereas less is known of belowground organisms. The present study focuses on enchytraeids (Oligochaeta) of heathland ecosystems and how the enchytraeid community has responded to simulated climate change in a long-term field experiment. Either increased temperature or repeated drought was applied for 13 years to field plots located in Wales, The Netherlands and Denmark representing a gradient in precipitation and annual temperature fluctuations thereby providing an opportunity to study biological responses on a local (within sites) and regional scale. Warming treatments increasing night-time temperature (0.5–1 °C higher than ambient at 5 cm soil depth) had no detectable effects on the enchytraeid communities. Increased intensity and frequency of drought had rather weak persistent effects on total enchytraeid abundance suggesting that ecosystem functions of enchytraeids may only be transiently impacted by repeated spring or summer drought. However, drought treatment had persistent negative effects on species richness and community structure across sites. Drought treated plots harboured only 35–65% of the species present in control plots, and the reduction of species richness was most pronounced at the driest sites. It is discussed that soil invertebrates, due to their weak migratory potential, may be more liable to extinction under changing climatic conditions than above-ground species, and therefore consequences of climate change to soil organisms need particular attention in future research.     

Influence of Cognettia sphagnetorum (Oligochaeta: Enchytraeidae) on nitrogen mineralization in homogenized mor humus

Summary The influence of the enchytraeid species Cognettia sphagnetorum on N mineralization in homogenized mor humus was examined in a laboratory study. The mor humus was incubated in containers (150 ml) for 8 months at various temperatures and with different moisture levels. Two series were used, one with C. sphagnetorum and one without. The presence of enchytraeids in the cultures increased the level of NH₄ ⁺ and NO₃ ^- by about 18% compared with the cultures without enchytraeids. Almost 40% of this difference was explained by the decomposition of dead enchytraeids. Temperature and soil moisture were the most important factors controlling the mineralization rate. The optimum moisture for N mineralization was between pF 1.6 and 1.1.Dedicated to the late Prof. Dr. W. Kühnelt     

Evaluating the efficiency of sampling methods in assessing soil macrofauna communities in arable systems

The soil fauna is often a neglected group in many large-scale studies of farmland biodiversity due to difficulties in extracting organisms efficiently from the soil. This study assesses the relative efficiency of the simple and cheap sampling method of handsorting against Berlese–Tullgren funnel and Winkler apparatus extraction. Soil cores were taken from grassy arable field margins and wheat fields in Cambridgeshire, UK, and the efficiencies of the three methods in assessing the abundances and species densities of soil macroinvertebrates were compared. Handsorting in most cases was as efficient at extracting the majority of the soil macrofauna as the Berlese–Tullgren funnel and Winkler bag methods, although it underestimated the species densities of the woodlice and adult beetles. There were no obvious biases among the three methods for the particular vegetation types sampled and no significant differences in the size distributions of the earthworms and beetles. Proportionally fewer damaged earthworms were recorded in larger (25 × 25 cm) soil cores when compared with smaller ones (15 × 15 cm). Handsorting has many benefits, including targeted extraction, minimum disturbance to the habitat and shorter sampling periods and may be the most appropriate method for studies of farmland biodiversity when a high number of soil cores need to be sampled.     

A method of processing soil core samples for root studies by subsampling

Root studies are generally believed to be very important in ecological research. Soil coring is a valuable approach to root research, but it requires a very large amount of processing time. We present here a method for processing soil cores consisting of the combination and homogenization of several soil cores from a plot, with subsequent subsampling for root extraction. The required subsample size was determined for a topsoil and a subsoil sample from a groundnut field and was found to be 5–10% of the total soil sample. Advantages and limitations of the method are discussed.     

Dynamics and stratification of Enchytraeidae in the organic layer of a Scots pine forest

In a Dutch Scots pine forest an experiment was conducted to quantify the role of soil biota in the functioning of the soil ecosystem, and the effects of enhanced nitrogen deposition. For this, the site was sampled at 8-week intervals during 2.5 years. This paper reports on the population dynamics of enchytraeids in the field and in stratified litterbags. Mean yearly abundance of the enchytraeid community in the field was 47 600 m^–2, or 0.70 g (dry weight) m^–2. The community consisted mainly of three species: Cognettia sphagnetorum, Marionina clavata and Achaeta eiseni, of which C. sphagnetorum was dominant. The enchytraeid populations showed a marked stratification in the same sequence. Freshly fallen pine needles were colonized by C. sphagnetorum, while other species followed much later. It was found that data from the litterbags were reasonably comparable with field data, when expressed per gram of dry substrate, but less so when expressed per square metre. Multiple regression analysis of the data showed that the population dynamics in the litter layer could largely be explained by temperature and moisture fluctuations; in deeper layers other factors, such as the stage of decomposition, were probably more important. Received: 26 June 1997     

Diversity of terrestrial Enchytraeidae (Oligochaeta) in Latin America: Current knowledge and future research potential

This article reviews the current knowledge on terrestrial enchytraeid diversity in Latin America and it outlines the research potential that this group offers. Enchytraeids occur worldwide in all soils with sufficient moisture, oxygen and nutrient supply, but knowledge on their diversity and functioning in the tropics is practically non-existent, except in Latin America. Here, taxonomic efforts and research projects have led to the knowledge of currently 62 terrestrial or semi-aquatic species. Abundance of enchytraeids could be determined at the species level, and differences in the ecological behaviour of species were detected concerning factors such as soil type and land use. Especially South America harbours a rich and idiosyncratic enchytraeid fauna, dominated by genera or species absent or rare in other regions of the world, which is interesting from the phylogenetic point of view. However, only a minute fraction of the actual diversity is known to-date. Seventeen species are possibly peregrines. Densities appear to be comparable to those in temperate regions, from below 10,000 to 270,000 ind. m⁻², suggesting that enchytraeids may be as important for soil processes as in temperate regions. Site-specific species richness makes enchytraeids good biological indicators. Enchytraeids provide a widely open field for research in Latin America ranging from taxonomy and faunistics over biogeography, phylogenetics, comparative morphology, and developmental biology, to fundamental and applied ecology. The widespread ignorance concerning enchytraeids among scientists and naturalists as well as the current taxonomic impediment should be overcome by sound taxonomical and ecological work, training courses, identification manuals and popularization.     

Spatial distribution of isopods in an oak–beech forest

Soil macroinvertebrates were studied in a Mediterranean-type forest on brown-pebble forest soils in southern Russia. At the site, 144 intact soil cores (76 cm² each) forming a grid of 24 × 6 units were taken in order to determine animal spatial distribution. Abundance of isopods was 166.3 ± 16.0 indiv. m^–2 and they constituted about 12% of the total macrofaunal abundance. Biomass of isopods was 3.5 g m^–2, or about 21% of the total biomass of macrofauna. Three woodlice genera (Armadillidium, Cylisticus, and Trachelipus) were found at the site. The two latter genera formed almost all (93%) of the isopod population. We found that spatial distribution of woodlice was heterogeneous: areas with 4–5 individuals per sample were neighboring those without animals. In order to study soil factors influencing isopod distribution in the brown-pebble forest soil, the size of a sample was artificially increased by combining adjacent sample units. Litter mass (r = 0.41) and loss on ignition (LOI) (r = –0.55) significantly influenced isopod distribution. Soil pH was near neutral (6.79), LOI was 8.39, and the water holding capacity was 70.9%. Pebbles comprised up to 84% of the sample's mass. Ca. 40 samples are recommended for estimation of isopod abundance in brown forest soil.     

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

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

Effects of animal manure and mineral fertilizer on the total carbon and nitrogen contents of soil size fractions

Summary Soil was sampled in autumn 1984 in the 132 field (sandy loam soil) of the Askov long-term experiments (started in 1894) and fractionated according to particle size using ultrasonic dispersion and sedimentation in water. The unmanured plot and plots given equivalent amounts of N (1923–1984 annual average, 121 kg N/ha) in either animal manure or mineral fertilizer were sampled to a depth of 15 cm, fractionated and analysed for C and N. Mineral fertilizer and animal manure increased the C and N content of whole soil, clay (<2 m) and silt (2–20 m) size fractions relative to unmanured samples, while the C content of the sand size fractions (fine sand 1, 20–63 m; fine sand 2, 63–200 m; coarse sand, 200–2000 m) was less affected. Clay contained 58% and 65°70 of the soil C and N, respectively. Corresponding values for silt were 30% and 26%, while sand accounted for 10% of the soil C. Fertilization did not influence this distribution pattern. The C : N ratio of the silt organic matter (14.3) was higher and that of clay (10.6) lower than whole-soil C:N ratios (12.0). Fertilization did not influence clay and silt C : N ratios. Animal manure caused similar relative increases in the organic matter content of clay and silt size fractions (36%). In contrast, mineral fertilizer only increased the organic matter content of silt by 21% and that of clay by 14%.     

Seasonal pattern of denitrification under an irrigated wheat-maize cropping system fertilized with urea and farmyard manure in different combinations

Under semiarid subtropical field conditions, denitrification was measured from the arable soil layer of an irrigated wheat–maize cropping system fertilized with urea at 50 or 100 kg N ha^–1 year^–1 (U₅₀ and U₁₀₀, respectively), each applied in combination with 8 or 16 t ha^–1 year^–1 of farmyard manure (FYM) (F₈ and F₁₆, respectively). Denitrification was measured by acetylene inhibition/soil core incubation method, also taking into account the N₂O entrapped in soil cores. Denitrification loss ranged from 3.7 to 5.7 kg N ha^–1 during the growing season of wheat (150 days) and from 14.0 to 30.3 kg N ha^–1 during the maize season (60 days). Most (up to 61%) of the loss occurred in a relatively short spell, after the presowing irrigation to maize, when the soil temperature was high and a considerable NO₃^–-N had accumulated during the preceding 4-month fallow; during this irrigation cycle, the lowest denitrification rate was observed in the treatment receiving highest N input (U₁₀₀+F₁₆), mainly because of the lowest soil respiration rate. Data on soil respiration and denitrification potential revealed that by increasing the mineral N application rate, the organic matter decomposition was accelerated during the wheat-growing season, leaving a lower amount of available C during the following maize season. Denitrification was affected by soil moisture and by soil temperature, the influence of which was either direct, or indirect by controlling the NO₃^– availability and aerobic soil respiration. Results indicated a substantial denitrification loss from the irrigated wheat–maize cropping system under semiarid subtropical conditions, signifying the need of appropriate fertilizer management practices to reduce this loss.     

Enchytraeid communities in successional habitats (from meadow to forest)

The study was conducted in two successional series: natural on mesotrophic soils (a meadow, two birch woods and a mixed coniferous forest), and human-made on sandy soils (old field, birch plantation and pine forest). In both these series species richness and diversity of enchytraeid communities decreased with advancing succession. In the first two successional stages, enchytraeid communities showed a high degree of similarity (the meadow and 30-year-old birch thicket, ore the old field and 10-year-old birch plantation). Variation in soil in the same plant communities cause differences in animal communities. The abundance of enchytraeids varied from 14 to 51 × 10³ m⁻² individuals. In the majority of habitats C. sphagnetorum was the dominant species. Soil pH showed a positive correlation with species diversity and negative with number of C. sphagnetorum. The body size of this species was small at low pH (presumably as a result of frequent reproduction). It is suggested that the way of reproduction of the dominant species could confer a competitive advantage, thereby accounting for the simplification of enchytraeid communities in later stages of succession.     

Aggregate associated carbon, nitrogen and sulfur and their ratios in long-term fertilized soils

Farmyard manure (FYM) and fertilizer applications are important management practices used to improve nutrient status and organic matter in soils and thus to increase crop productivity and carbon (C) sequestration. However, the long-term effects of fertilization on C, nitrogen (N) and sulfur (S) associated with aggregates, especially on S are not fully understood. We investigated the effects of more than 80 years of FYM (medium level of 40 Mg ka⁻¹ and high level of 60 Mg ka⁻¹) and mineral fertilizer (NPKS and NK) on the concentrations and pools of C, N, and S and on their ratios in bulk soil, dry aggregates and water stable aggregates on an Aquic Eutrocryepts soil in South-eastern Norway. A high level of FYM and NPKS application increased the proportion of small dry aggregates (<0.6 mm) by 8%, compared with the control (without fertilizer). However, both medium and high level of FYM application increased the proportion of large water stable aggregates (>2 mm) compared with mineral fertilizer (NPKS and NK). The total C and N pools in bulk soils were also increased in FYM treatments but no such increase was seen with mineral fertilizer treatments. The increased total S pool was only found under high level of FYM application. Water stable macroaggregates (>2 and 1–2 mm) and microaggregates (<0.106 mm) contained higher concentrations of C, N and S than the other aggregate sizes, but due to their abundance, medium size water stable aggregates (0.5–1 mm) contained higher total pools of all three elements. High level of FYM application increased the C concentration in water stable aggregates >2, 0.5–1 and <0.106 mm, and increased the S concentration in most aggregates as compared with unfertilized soils. Higher C/N, C/S and N/S ratios were found both in large dry aggregates (>20 and 6–20 mm) and in the smallest aggregates (<0.6 mm) than in other aggregate sizes. In water stable aggregates, the C/N ratio generally increased with decreasing aggregate size. However, macroaggregates (>2 mm) showed higher N/S ratios than microaggregates (<0.106 mm). We can thus conclude, that long-term application of high amounts of FYM resulted in C, N and S accumulation in bulk soil, and C and S accumulation in most aggregates, but that the accumulation pattern was dependent on aggregate size and the element (C, N and S) considered.     

Enchytraeid worms retard polycyclic aromatic hydrocarbon degradation in a coniferous forest soil

In this study the fate of naphthalene, fluorene and pyrene were investigated in the presence and absence of enchytraeid worms. Microcosms were used, which enabled the full fate of ¹⁴C-labelled PAHs to be followed. Between 60 and 70% of naphthalene was either mineralised or volatilised, whereas over 90% of the fluorene and pyrene was retained within the soil. Mineralisation and volatilisation of naphthalene was lower in the presence of enchytraeid worms. The hypothesis that microbial mineralisation of naphthalene was limited by enchytraeids because they reduce nutrient availability, and hence limit microbial carbon turnover in these nutrient poor soils, was tested. Ammonia concentrations increased and phosphorus concentrations decreased in all microcosms over the 56 d experimental period. The soil nutrient chemistry was only altered slightly by enchytraeid worms, and did not appear to be the cause of retardation of naphthalene mineralisation. The results suggest that microbial availability and volatilisation of naphthalene is altered as it passes through enchytraeid worms due to organic material encapsulation.     

Enchytraeid worms (Oligochaeta) enhance mineralization of carbon in organic upland soils

We investigated the functional role of enchytraeid worms (Oligochaeta) in organic upland soils experimentally, because that role of these animals is little known. We made microcosms of intact soil cores cut from two depths, 0–4 cm and 4–8 cm, of a Cambic Stagnohumic Gley from the Moor House National Nature Reserve (UK). Enchytraeids were added to half of the microcosms, resulting in four treatments: litter (L), litter + enchytraeids (L + E), soil (S) and soil + enchytraeids (S + E). Triplicates of each treatment were established, and all microcosms (60) were then incubated in the dark at 15°C, arranged in a fully randomized design. The experiment ran over 110 days, with five destructive harvests at days 10, 25, 50, 75 and 110, when microbial measurements (soil respiration and biomass C) as well as measures of decomposition (nutrient concentration in leachates) were made. Enchytraeids almost doubled the availability of organic carbon (measured as dissolved organic carbon in soil leachates) in the surface (0–4 cm) microcosms only. There were no effects of enchytraeids on the release of inorganic N or P from either soil horizon, although the release of ammonium and phosphate was correlated with the number of enchytraeids in the microcosms. The depth from which the soil was taken exerted a strong influence on nutrient leaching, with almost six times more ammonium and four times more carbon being leached from the surface (0–4 cm) layer than from the more decomposed (4–8 cm) horizon. There was little nitrate leaching from any of the treatments, with only one‐quarter as much nitrate leached from the surface (0–4 cm) as from the subsurface (4–8 cm) horizon. Enchytraeids had no detectable effect on microbial biomass, but they increased microbial respiration by 35% in the surface (0–4 cm) horizon. Because they enhanced microbial activity in this horizon we suggest that enchytraeids indirectly drive the processes of decomposition and nutrient mineralization in organic upland soils.