Endemic Collembola, privileged bioindicators of forest management

Our study compared the soil collembolan community at three semi-natural sites (a beech forest, a beech–fir forest and a fir stand) and three managed sites (Norway spruce, beech–fir and fir). Collembola were extracted from a total of 60 samples with a Berlese–Tullgren funnel, counted and identified to species level. A total of 7187 specimens, representing 51 species and 37 genera, were collected. There were significant differences between managed and natural forests (t-test, P=0.000). The communities and their population densities were significantly lower in the managed site: 51 species with 708,498 ind m⁻² in the semi-natural forests to 36 species and 306,042 ind m⁻² in the managed stands. The endemic component suffered a particularly severe decrease in species richness and abundance (57% and 71% lower in the managed forests, respectively). These species with narrow distribution and small local populations are doubly vulnerable to alterations of environmental conditions. They are very sensitive to loss of their natural habitat owing to human intervention and thus represent useful indicators of tolerance to environmental stress. We highlight the need to take endemic species into consideration in studies on the conservation of biodiversity because they are most at risk of extinction. Semi-natural forests are refuges for endemic species and should be protected.     

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.     

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.     

A multi-scale study of soil macrofauna biodiversity in Amazonian pastures

The influence of three spatially hierarchical factors upon soil macrofauna biodiversity was studied in four pasture plots in eastern Amazonia. The first factor was the local depth of the soil. The second factor was the ground cover type on the soil samples (bare ground, grass tufts, dead trees lying on the ground). The third factor was the dimensions of the grass tufts sampled (size and shape). The effect of each factor upon the morphospecies richness and density of total soil macrofauna was analysed. Detailed results are given for earthworms, termites, ants, beetles and spiders. All factors significantly affected the morphospecies richness and/or density of the soil macrofauna. The type of ground cover had the strongest influence, affecting the total richness and density of the soil macrofauna and of almost all the groups represented. The soil depth affected only the density of the termites and the global morphospecies richness. Interactions between soil depth and ground cover type affected the total macrofauna morphospecies richness and the density of the earthworms. The dimensions of the grass tuft influenced the global morphospecies richness, the morphospecies richness of the ants and the density of the spiders.     

Long-term land-use effects on soil invertebrate communities in Southern Piedmont soils,USA

Historically, a large percentage of land area in the Piedmont of the southeastern USA was under intensive agricultural management for the production of cotton. This intensive farming resulted in massive erosion, and general degradation of soil resources until insect pests and poor economic conditions forced large-scale abandonment of farmland around the 1930s. In subsequent decades, there have been four predominant land-uses in the region, and we sampled soil macroinvertebrates from three replicate sites of cultivated fields, grass-dominated fields, loblolly pine stands, and remnant hardwood stands for a period of 2 years, with the objective of examining soil invertebrate community composition in relation to these long-term land-uses. At each site we dug three or four soil pits that were 30 × 30 cm to a depth of 15 cm, and sorted the soil volume by hand for a time not more than 1 person h, collecting all invertebrates ~5 mm in length or larger. We recorded abundance data for all invertebrate taxa collected, and we calculated community indices including diversity, evenness, rank abundance and percent similarity in order to identify patterns of community assemblage within each land-use type. Results suggest that soils in hardwood stands support the most taxonomically diverse macroinvertebrate communities followed by pine stands, pastures, and cultivated fields in order of decreasing diversity. For earthworms, Diplocardia spp. (North American megascolecids) were most abundant in the hardwood stands, but sometimes made up a substantial fraction of the community in other land-uses; whereas lumbricid earthworms (primarily introduced Apporectodea spp.) were most abundant in the cultivated and pasture soils, or showed no consistent habitat preference (native Bimastos spp.). Scarab beetles (larvae and adults) were common in all four systems, but reached the highest densities in cultivated and grass sites. Carabid beetle larvae were collected most often from cultivated soils. Several taxa were collected either exclusively or predominantly from forested sites, including diplopods, chilopods, gastropods, and several taxa of Diptera. These results indicate that long-term soil disturbance and the attendant differences in vegetation structure have profoundly influenced the community composition of invertebrates in Southern Piedmont soils, and that more intense disturbance results in a less diverse invertebrate community composed of a few, frequently non-native, disturbance-tolerant taxa.     

Earthworm collection from agricultural fields: Comparisons of selected expellants in presence/absence of hand-sorting

The role earthworms play in soil fertility is under increasing scientific scrutiny, especially in light of the fact that farmers are seeking to reduce soil tillage. However, there are many discrepancies in earthworm sampling methods. The aim of this study is to compare the efficiency of three chemical expellants (formaldehyde, commercial ‘hot’ mustard and allyl isothiocyanate, abbreviated AITC), with two sampling methods: (i) a simple method that consisted in spreading an expellant solution on the soil and retrieving earthworms that emerged at the soil surface, and (ii) a combined method that consisted in applying a chemical to expel earthworms and then hand-sorting the remaining earthworms from the block of soil. Sampling efficiency was measured in terms of earthworm density and biomass, for endogeic and anecic earthworms.With the simple method, a higher density of earthworms was sampled using formaldehyde and AITC than with mustard. Formaldehyde, AITC and mustard expelled not significantly different biomasses of 47.7, 31.9 and 20.5 g m^?2, respectively, on average over the three plots. The combined method did not yield a significantly different density or biomass with the different chemicals.Formaldehyde is toxic and commercial ‘hot’ mustard is difficult to standardise and inefficient when used without hand-sorting. Accounting for the accuracy of the sampling methods as well as the toxicity of the chemicals to users and soil organisms, AITC appears to be a reliable and promising chemical expellant whether or not in combination with hand-sorting. Its use would be a step towards standardizing earthworm sampling methods.     

Combinations of ripping depth and tine spacing for compacted sandy and clayey soils

Soil compaction is one of the major problems facing modern intensive agriculture. To remove soil compaction and restore soil productivity soil must be ripped to loosen it. Ripping is a costly process involving high fuel consumption, as well as depreciation of the implements through wear and tear. This article shows research into some combinations of tine spacing and ripping depth and their consequences for soil properties and grain yields. Three sites were chosen for these experiments on clayey and sandy soils. Treatments were a factorial of three tine spacings (20, 30 and 40 cm) by three ripping depths (15, 30 and 40 cm) together with the control.Commercial gypsum at 2.5 t/ha was applied to all treatments to maintain soil structure after ripping and the treatments were treatments were monitored for two seasons under wheat and barley crops.The highest grain yield in sandy soil was found with the combination of 40 cm ripping depth and 20 cm tine spacing. In clayey soils tine spacings of 20 cm and 30 cm in combination with 40 cm ripping depth, were equally effective for grain yield. The shallowest depth treatment, 15 cm, did not significantly affect grain yields regardless of tine spacing. It seems that the best practical compromise of tine spacing and ripping depth is 30 cm × 30 cm. The highest stored soil water was obtained from the deepest ripping and the widest tine spacing (40 cm × 40 cm) treatments and the lowest was obtained from the shallowest depth and narrowest spacing (15 cm × 20 cm) treatments which was still higher than the control treatment. However, due to soil re-settlement and re-compaction, the soil water storage obtained in the year after ripping to 40 cm depth was in many cases only equal to that obtained from 30 cm ripping depth. Soils ripped at 30 cm or deeper had significantly higher water infiltration rate than soils ripped at 15 cm depth. Soil bulk density, though decreased significantly in all ripping treatments relative to the controlled treatments in the first year, showed no stable pattern of change in the second year. All shallow ripping treatments (15 cm) regardless of tine spacing had similar soil strength and were not significantly different from the control. The other two ripping depths in general were equal, and significantly better than the controls. It is concluded that ripping to 30 or 40 cm depth in combination with 30 or 40 cm tine spacing was most effective for treating compacted soils.     

有机管理对不同土地利用方式下土壤质量的影响

许多研究发现有机管理可以改善农田土壤质量,但是否不同土地利用方式下都存在此结论尚未明确。为探究有机管理对不同土地利用方式土壤质量的影响,本研究基于一个多土地利用方式的有机管理农场及其附近常规管理农田进行土壤质量调查,对比不同管理措施及大棚菜地、果园、露天菜地、农田边界、稻田田埂5种土地利用方式下农田土壤质量的差异,并对不同管理措施下农田土壤养分含量、重金属含量、动物数量共计20个指标进行方差分析和主成分分析。研究发现虽然总体上(综合5种土地利用方式的均值)有机管理的土壤pH显著高于常规农田;但针对一种土地利用方式,只有露天菜地和农田边界的土壤pH显著提高。总体上有机管理下土壤全磷、全钾、有效磷含量显著降低,但露天菜地土壤有机质和全氮含量均显著高于常规管理农田。有机管理在总体上显著降低了土壤Cr、Cu、Ni、Zn含量;但针对一种土地利用方式,只显著降低了大棚菜地土壤Cr、Ni、Zn含量,果园和稻田田埂土壤Cu含量,露天菜地和农田边界土壤Zn含量。有机管理虽然总体上均显著增加了土壤中蜘蛛目、倍足纲、步甲、蚯蚓的数量;但针对一种土地利用方式,只显著增加了露天菜地地表蜘蛛目、倍足纲、步甲,稻田田埂地表步甲、土壤蚯蚓以及果园倍足纲数量。主成分分析结果表明,不同管理方式下土壤质量差异明显,有机管理下蜘蛛目、倍足纲、步甲和蚯蚓等土壤动物较多,土壤pH较高,土壤P含量较低,重金属Cu、Pb、Zn、Cr含量较低,但并非在所有土地利用方式下都成立。由此可见,有机管理虽然总体上可以改善土壤质量,但受具体管理措施、人为投入品数量和有机种植时间等因素影响,并非在所有土地利用方式下效果都显著,且在不同土地利用方式下显著改善的指标也各不相同,因此需要针对不同土地利用方式和不同指标采取针对性的改善措施。     

Earthworm extraction with onion solution

Earthworm populations can be assessed using various extraction solutions of which the most widely used is formaldehyde (0.5%). Although formaldehyde extraction has been proposed as a standard method by ISO, its use has been questioned due to its toxic and carcinogenic properties, and the associated risks to human health and the environment. Alternative, safer methods are needed, so the present study was undertaken to evaluate the efficiency of a low cost and non-toxic onion (Allium cepa L.) extract solution in collecting earthworms in two soil types in Southern Brazil. The study was divided in three parts. In the first, we examined the efficiency of solutions with 60, 80, 100, 170 and 200 g onion extract L⁻¹ H₂O in extracting earthworms from a sandy Ultisol underlying a citrus orchard. The second experiment evaluated the effectiveness of solutions of 25, 75, 125 and 175 g onion extract L⁻¹ H₂O compared with the standard formaldehyde solution (0.5%) in extracting earthworms from a clayey Oxisol cultivated with corn. The third experiment compared the onion extract solution at concentration of 175 g L⁻¹ with formaldehyde (0.5%) in extracting earthworms from the sandy Ultisol. Increasing concentrations of onion extract solution led to higher extraction of earthworms in both soil types, but the highest numbers were obtained with 175 g L⁻¹ H₂O, a concentration that had similar efficiency to the standard solution of formaldehyde (0.5%). The onion extract solution is a low cost and non-toxic alternative to the use formaldehyde in extracting worms from soils, although further research is warranted to assess its use in other soils, climates and vegetation conditions.     

Degradation and small-scale spatial homogenization of topsoils in intensively-grazed steppes of Northern China

Overgrazing has led to severe degradation and desertification of semi-arid grasslands in Northern China over the last decades. Despite the fact that vegetation is often heterogeneously distributed in semi-arid steppes, little attention has been drawn to the effect of grazing on the spatial distribution of soil properties. We determined the spatial pattern of soil organic carbon (SOC), total nitrogen (N_tot), total sulphur (S_tot), bulk density (BD), pH, Ah thickness, and carbon isotope ratios (δ¹³C) at two continuously grazed (CG) and two ungrazed (UG79 = fenced and excluded from grazing in 1979) sites in Leymus chinensis and Stipa grandis dominated steppe ecosystems in Inner Mongolia, Northern China. Topsoils (0–4 cm) were sampled at each site using a large grid (120 m × 150 m) with 100 sampling points and a small plot (2 m × 2 m) with 40 points. Geostatistics were applied to elucidate the spatial distribution both at field (120 m × 150 m grid) and plant (2 m × 2 m plot) scale. Concentrations and stocks of SOC, N_tot, S_tot were significantly lower and BD significantly higher at both CG sites. At the field scale, semivariograms of these parameters showed a heterogeneous distribution at UG79 sites and a more homogeneous distribution at CG sites, whereas nugget to sill ratios indicated a high small-scale variability. At the plant scale, semivariances of all investigated parameters were one order of magnitude higher at UG79 sites than at CG sites. The heterogeneous pattern of topsoil properties at UG79 sites can be attributed to a mosaic of vegetation patches separated by bare soil. Ranges of autocorrelation were almost congruent with spatial expansions of grass tussocks and shrubs at both steppe types. At CG sites, consumption of biomass by sheep and hoof action removed vegetation patches and led to a homogenization of chemical and physical soil properties. We propose that the spatial distribution of topsoil properties at the plant scale (<2 m) could be used as an indicator for degradation in semi-arid grasslands. Our results further show that the maintenance of heterogeneous vegetation and associated topsoil structures is essential for the accumulation of SOM in semi-arid grassland ecosystems.     

Spatial patterns of grasses influence soil macrofauna biodiversity in Amazonian pastures

Grasslands are often characterized by small-scale spatial heterogeneity due to the juxtaposition of grass tufts and bare ground. Although the mechanisms generating plant spatial patterns have been widely studied, few studies concentrated on the consequences of these patterns on belowground macrofauna. Our objective was to analyze the impact of grass tuft (Brachiaria bryzantha cv. marandu) spatial distribution on soil macrofauna diversity in Amazonian pastures, at a small scale (less than 9 m²). Soil macrofauna was sampled among B. bryzantha tufts, which showed a variable spatial distribution ranging from dense to loose vegetation cover. The vegetation configuration explained 69% of the variation in total soil macrofauna density and 68% of the variation in total species richness. Soil macrofauna was mainly found in the upper 10 cm of soil and biodiversity decreased with increasing distances to the nearest grass tuft and increased with increasing vegetation cover. The size of the largest grass tuft and the micro-landscape connectivity also had a significant effect on biodiversity. The density and species richness of the three principal soil ecological engineers (earthworms, ants and termites) showed the best correlations with vegetation configuration. In addition, soil temperature significantly decreased near the plants, while soil water content was not influenced by the grass tufts. We conclude that soil macrofauna diversity is low in pastures except close to the grass tufts, which can thus be considered as biodiversity hotspots. The spatial arrangement of B. bryzantha tussocks influences soil macrofauna biodiversity by modifying soil properties in their vicinity. The possible mechanisms by which these plants could affect soil macrofauna are discussed.     

Quantitative extraction of macro-invertebrates from temperate and tropical leaf litter and soil: efficiency and time-dependent taxonomic biases of the Winkler extraction

Winkler extractors, a simple device presumed to extract macro-invertebrates efficiently from soil and litter samples, is being used increasingly in ecological surveys and functional studies of soil macro-invertebrate communities. In this study the extraction efficiency and taxonomic bias of the Winkler extraction are evaluated for extraction periods of 3 h up to 7 weeks, calibrated by hand-sorting after 7 weeks. The method extracts most macro-invertebrates completely or to a proportion of over 90% except Isopoda, Diplopoda and Mollusca. However, for an exhaustive result, a long extraction period of several weeks is necessary. For the most speciose group (adult beetles) and for the commonly most abundant group (ants), a short extraction of 3 days was sufficient to get 70% of the individuals and nearly all species. Three days was also sufficient to recover the rank abundance order of beetle families, while for ‘higher taxa’ and for Chilopoda species, 4 and 3 weeks were necessary, respectively. Optimum extraction times for the abundant macro-invertebrate groups and possible adjustment factors for the soil macro-invertebrates of temperate woodlands are proposed to compensate the taxonomic bias caused by short extraction periods. However, for recording an accurate snapshot of the soil and litter fauna at a particular time, shorter extraction periods are advisable because of the short life cycle of many soil invertebrates causing emergence of later stages or a second generation during longer extraction periods. The problem of contamination of samples is also discussed.     

Earthworms in Chromolaena odorata (L.) King and Robinson (Asteraceae) fallows along a chronosequence: Changes in community structure and identification of persistent and indicator species

The species Chromolaena odorata (Asteraceae) is a notorious invasive shrub spreading throughout West and Central Africa and as such, there is a need to determine its environmental impact, particularly on soil biodiversity and functioning. Indeed, soil organisms such as earthworms are known to strongly influence soil properties and biogeochemical cycles. This study, conducted in Central Côte d’Ivoire, aims to investigate the temporal dynamics of earthworm communities in C. odorata fallows of different ages and to identify associated indicators and persistent species. Three distinct classes of fallows identified by local farmers, were considered: young (1–3 years, C1), medium-aged (4–8 years, C2) and old (>9 years, C3). Each of the classes included four plot replicates where earthworms were sampled using the Tropical Soil Biology and Fertility (TSBF) 25 cm × 25 cm × 30 cm soil monolith method. The study of earthworm communities was focused on density, biomass, diversity and complementarity. Indicator values (IndVals) were used to identify indicator species of the classes of fallows. The shrub exerted a mixed influence on earthworms depending on the functional group, with litter feeders and polyhumics declining over time as a result of a reduction of the litter availability on the soil surface. The species richness was significantly greater in C1 than in the other classes although the Shannon–Weaver's index did not vary significantly. However, a cluster analysis performed on densities highlighted marked differences between C2 and the two other classes in terms of community composition. Indicator species were found for C1 and C2. The geophagous Millsonia omodeoi has emerged as a persistent species as its density and biomass steadily increased so that it became the dominant species in old fallows. The roles of litters and soil parameters in influencing earthworm communities are discussed.     

Inability of Near Infrared Reflectance Spectroscopy (NIRS) to identify belowground earthworm casts in no-tillage soil

Several studies have emphasised the ability of Near Infrared Reflectance Spectroscopy (NIRS) to identify surface earthworm casts in the field. However, less is known about casts deposited within the soil, which usually represent the majority found in the field. This study tested the ability of NIRS to identify belowground casts in agricultural systems. Casts and surrounding soils were sampled at depths of 20–30 cm in a loamy soil under no tillage for 12 years. To distinguish different types of cast, sizes and orientations relative to the horizontal plane were measured. NIRS analyses and analyses of carbon and nitrogen content were also performed to compare casts to surrounding soils. Casts were classified into 4 size classes, with no preferential orientation. Cast carbon and nitrogen content were not influenced by their size and did not differ from surrounding soils. PCAs performed on the NIRS data did not allow casts to be differentiated from surrounding soils, regardless of size class. However, soil aggregates were clearly differentiated probably due to their spatial distribution in the soil. Although this study did not identify specific NIRS signatures for casts, it shows the utility of this method to investigate the origin of the soil consumed by earthworms. In our case, NIRS analyses suggest that the high bulk density of the soil (1.42 g cm⁻³) forced ingestion by endogeic earthworms, simply to move around, without preferential selection for organic matter. Consequently, their casts were deposited a few mm from where they had ingested soil with similar organic matter quality.     

Deep-burrowing earthworm additions changed the distribution of soil organic carbon in a chisel-tilled soil

We investigated the influence of earthworms on the three-dimensional distribution of soil organic carbon (SOC) in a chisel-tilled soil. By burrowing, foraging, and casting at the surface and throughout the soil, anecic earthworms such as Lumbricus terrestris L. may play a major role in regulating the spatial distribution of organic matter resources both at the surface and within the soil. In the fall of 1994, we manipulated ambient earthworm communities, which were without deep burrowing species, by adding 100 earthworm individuals m⁻² in spring and fall for 3 years. Overall, the biomass of L. terrestris was increased with earthworm additions and total earthworm biomass declined compared with ambient control treatments. To investigate the spatial variability in soil organic carbon due to this shift in earthworm community structure, we sampled soil on a 28×24 cm grid from the surface to 40 cm in four layers, 10 cm deep. Samples were analyzed for total carbon. We found that additions of anecic earthworms significantly increased average soil organic carbon content from 16.1 to 17.9 g C kg⁻¹ for the 0–10 cm soil, and from 12.4 to 14.7 g kg⁻¹ at 10–20-cm depth, and also changed the spatial distribution of soil organic carbon from uniform to patchy, compared with the ambient treatment.     

A method of extracting earthworms from cores of soil with minimum damage to the soil

 Lumbricid earthworms have often been shown to increase the growth of plants. The earthworms and plants were generally grown together in the same soil, although sometimes earthworms were reluctant to enter the soil. It was not possible to isolate the mechanism for the increased growth, as no method was available to extract the earthworms with no damage to the soil before the plants were grown. A method is described which enabled Aporrectodea caliginosa, but not A. trapezoides, to enter all cores, and which extracted A. caliginosa from the cores probably with minimum damage to the soil. Received: 20 October 1998     

Inter- and intra-specific interactions of Lumbricus rubellus (Hoffmeister, 1843) and Octolasion lacteum (Örley, 1881) (Lumbricidae) and the implication for C cycling

Earthworms are important engineering species of many terrestrial ecosystems as they play a significant role in regulating C turnover. The effects of earthworms on moderating C decomposition processes differ across species and with interactions between species, which is not fully understood. We carried out an experiment to study the interactions of Lumbricus rubellus and Octolasion lacteum, and their effects on soil respiration. Laboratory mesocosms were set up using tulip poplar (Liriodendron tulipifera) leaf litter and varying densities of earthworms in single and combined species treatments. CO₂ efflux rate was used as an indicator of C decomposition rates, and measured with CO₂ sensors every five days over one month. L. rubellus induced higher leaf consumption rate and higher CO₂ efflux than O. lacteum; meanwhile O. lacteum grew more than L. rubellus. Both litter consumption rate and growth rate of earthworms decreased with increasing earthworm density. Soil CO₂ efflux increased with increasing earthworm density (from ∼1-2 μg CO₂ g⁻¹ hr⁻¹ with no earthworms to ∼ 4 μg CO₂ g⁻¹ hr⁻¹ with 8 earthworms). Combining the two species had a synergistic effect on leaf litter consumption, and neutralizing effects on soil respiration. The data suggest that the strength of intra- and inter-specific interactions among earthworm ecological groups varies at different absolute and relative densities, leading to altered leaf litter decomposition and C cycling.     

Short-term responses of two contrasting species of earthworms in an agricultural soil amended with coal fly-ash

With the renewed interest in the use of coal fly-ash for amendment of agricultural soils in Australia, we assessed how earthworms, as indicators of soil health, responded to this ameliorant. We assessed survival, weight, burrowing and elemental concentrations for earthworms of a native unnamed Megascolecid species and of exotic Aporrectodea trapezoides in intact soil cores treated with an alkaline fly-ash at rates equivalent to 0, 5 and 25 t/ha over 6 weeks. Fly-ash did not affect survival, growth, number of burrows created or phosphorus solubilisation. Transfer of the earthworms to the new environment having vastly different pH from where they were collected, and possibly overcrowding, caused mortality in the soil cores for all treatments. A. trapezoides that had smaller individuals suffered mortality of 12% compared with 23% for the larger earthworms of Megascolecids. Earthworms of Megascolecids each increased their weight by 0.24g (25% of their original weight) while those of A. trapezoides lost 0.18g each (21% of their original weight). The difference in growth between the two earthworms was associated with grazing habit and probably with the large difference in the pH between source soil and that of the core soil. Megascolecids appeared to minimize grazing on ash-tainted soil and so ingested less Zn, which was more abundant in the fly-ash than in the soil, compared with A. trapezoides that had elevated concentration of this metal. Extractable P in the soil was increased with both species of earthworms, more so with the exotic species that solubilized 11% more P than the native Megascolecids. The benign influence of fly-ash on survival and growth of worms was associated with the pH of soil remaining unchanged during the six weeks of incubation.     

Effects of earthworms on nitrogen mineralization

The influence of earthworms (Lumbricus terrestris and Aporrectodea tuberculata) on the rate of net N mineralization was studied, both in soil columns with intact soil structure (partly influenced by past earthworm activity) and in columns with sieved soil. Soil columns were collected from a well drained silt loam soil, and before the experiment all earthworms present were removed. Next, either new earthworms (at the rate of five earthworms per 1200 cm³, which was only slightly higher than field numbers and biomass) were added or they were left out. At five points in time, the columns were analyzed for NH ₄ ⁺ , NO ₃ ^– , and microbial biomass in separate samples from the upper and lower layers of the columns. N mineralization was estimated from these measurements. The total C and N content and the microbial biomass in the upper 5 cm of the intact soil columns was higher than in the lower layer. In the homogenized columns, the C and N content and the microbial biomass were equally divided over both layers. In all columns, the concentration of NH ₄ ⁺ was small at the start of the experiment and decreased over time. No earthworm effects on extractable NH ₄ ⁺ were observed. However, when earthworms were present, the concentration of NO ₃ ^– increased in both intact and homogenized cores. The microbial biomass content did not change significantly with time in any of the treatments. In both intact and homogenized soil, N mineralization increased when earthworms were present. Without earthworms, both type of cores mineralized comparable amounts of N, which indicates that mainly direct and indirect biological effects are responsible for the increase in mineralization in the presence of earthworms. The results of this study indicate that earthworm activity can result in considerable amounts of N being mineralized, up to 90 kg N ha^–1 year^–1, at the density used in this experiment.     

Interaction between earthworms and arbuscular mycorrhizal fungi on the degradation of oxytetracycline in soils

The interactive impact of earthworms (Eisenia fetida) and arbuscular mycorrhizal fungi (Rhizophagus intraradices, AM fungi) on the degradation of oxytetracycline (OTC) in soils was studied under greenhouse conditions. Treatments included maize plants inoculated vs. not inoculated with AM fungi and treated with or without earthworms at low (1 mg kg⁻¹ soil DM) or high (100 mg kg⁻¹ soil DM) OTC rates. The root colonization rate, the hyphal density of mycorrhizae, the residual OTC concentration in soils, catalase, dehydrogenase, urease, soil microbial biomass C, Shannon–Wiener index (H) for microbial communities from T-RFLP profiles were measured at harvest. The results indicated that earthworms and AM fungi would individually or interactively enhance OTC decomposition and significantly decreased the residual OTC concentration at both high and low OTC rates. Both earthworms and AM fungi could promote the degradation of OTC by increasing soil microbial biomass C at both high and low OTC rates. The effect of soil enzyme activity and soil microbial diversity on OTC decomposition was different between high and low OTC rates. Hyphomicrobium and Bacillus cereus were dominant bacteria, and Thielavia and Chaetomium were dominant phyla of fungi at all occasions. Earthworm activity stimulated the growth of Hyphomicrobium and Thielavia, while AM fungi may stimulate B. cereus, Thielavia and Chaetomium, resulting in greater OTC decomposition. The interaction between earthworms and AM fungi in affecting the degradation of OTC may be attributed to different mechanisms, depending on soil microbial biomass, function (enzyme activity) and communities (the abundance of Hyphomicrobium, B. cereus, Thielavia and Chaetomium) in the soil.