Biometric relationships in earthworms (Oligochaeta)

Digging and hand-sorting of soil blocks is a very widespread method in the study of earthworm communities. One disadvantage of this method is that it is very time consuming and often many earthworms are incomplete because they were cut by the digging tools. When authors report earthworm biomass, no mention is made of the assessment of any relationship between the mass of those cut earthworms and their overall weight. In such cases, biomass is generally underestimated. In this paper, our objective was to propose a new method to estimate the weight of incomplete earthworms on the basis of preclitellar diameter and its usefulness for studying the dynamics of earthworm populations. Complete earthworms were collected from samplings performed in native savannahs and man-made pastures of the eastern plains of Colombia and from a poplar grove (Populus sp.) in Central Spain. A strong correlation between the preserved fresh weight and the maximum preclitellar diameter was found for all the species studied. Three types of models have provided a convenient method to estimate earthworm biomass: (i) linear for almost all the species; (ii) exponential for a large Neotropical anecic species, Martiodrilus carimaguensis (Glossoscolecidae); and (iii) second degree polynomic equation.     

不同土地利用方式下的蚯蚓种群特征及其与土壤生物肥力的关系

采用样方法对华北平原(河北曲周)盐渍化改造区7种土地利用方式下的蚯蚓种群进行详细调查,并通过培养实验研究了蚯蚓种群特征对若干土壤生物学指标的影响。结果表明:(1)在7种土地利用调查样地中共存在蚯蚓有3个科,5个属,5个种,其中赤子爱胜蚓(Eisenia fetida)占调查样地总个体数的60%以上,梯形流蚓(Aporrectodea trapezoides)和赤子爱胜蚓两个种在本地区广泛分布,样点出现频率分别为74%和44%,为该地区的优势种;(2)不同土地利用方式的蚯蚓种群密度及生物量变化趋势是:庭院菜地>直立免耕>清茬免耕>商品菜地>传统玉米地>果园>原貌地。其中庭院菜地蚯蚓种群的平均密度和生物量分别达到272 Ind.m-2和68.04gm-2;(3)蚯蚓种群密度和物种数等种群特征与土壤基础呼吸强度、微生物生物量碳含量成显著正相关(p<0.01),与土壤基础呼吸商成显著负相关(p<0.01);(4)不同土地利用方式下,蚯蚓的种群密度、生物量等种群特征对土壤中微生物群落的影响作用显著。蚯蚓生物量越大、种群越丰富的土壤有机质、氮、磷、钾等有效成分越高,反之则相反。室内培养实验表明,随着蚯蚓个体数量增加土壤原生动物总丰度、微生物生物量碳、氮也存在升高的趋势,与用土壤生物学特性指标及土壤化学特性指标评价的结果基本一致。     

Impact of earthworms on decomposition of garden refuse

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

Enhanced microbial degradation of pentachlorophenol from soil in the presence of earthworms: Evidence of functional bacteria using DNA-stable isotope probing

This study investigated the effect of two earthworm species (Amynthas robustus Perrier and Eisenia fetida Savigny) on the soil microbial degradation of pentachlorophenol (PCP). PCP-degrading microbes were identified using DNA-stable isotope probing (SIP). The results showed that adsorption and fixing to soil particles and organic fractions dominated the fate of PCP in soil without any amendments. The inoculation of both earthworm species significantly enhanced soil PCP disappearance and basal respiration. The DNA-SIP results revealed that Klebsiella, Cupriavidus, Aeromonas, and Burkholderia spp. were present at higher relative abundances in [¹³C]-labeled-PCP-amended soil microcosms than [¹²C]-PCP-amended soil in the presence of A. robustus, indicating that these bacterial species were responsible for PCP assimilation. Cupriavidus and Aeromonas sp. were also detected in the earthworm gut before inoculation, and their relative abundance was affected by earthworms. These results demonstrated that earthworms can introduce functional bacteria into soils and increase the population of PCP-degrading bacteria, thereby accelerating soil PCP degradation.     

Using FAME profiles for the characterization of animal wastes and vermicomposts

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

蚯蚓对红壤有机质分解的影响

The earthworms Pheretima carnosa, Drawida gisti and Eisenia foetida were studied to compare their contributions to the decomposition of various organic materials surface-applied on red soil in a 165-day greenhouse experiment. The native species Pheretima carnosa and Drawida gisti were equally effective in accelerating the decomposition of maize residue, according to fresh body weight, while commercial species Eisenia foetida had no significant influence on dry mass loss of maize residue. Liming with CaCO3 or CaO showed little effect on maize residue breakdown involved by Pheretima carnosa, but it inhibited this process involved by Drawida gisti. The capability of Pheretima carnosa to the decomposition of five kinds of organic materials was thoroughly examined. The dry mass losses in worm treatments were in the order of soybean residue ＞ maize residue ＞ pig manure ＞ semi-decayed maize ＞ ryegrass. However, the relative contributions of the earthworm to dry mass loss were in the order of pig manure (89.8%) ＞ semi-decayed maize residue (49.1%) ＞ maize residue (29.4%) ＞ soybean residue (20.9%) ＞ ryegrass residue (16.5%). Pheretima carnosa consumed 20～120 mg dry weight of organic material per gram fresh weight of biomass per day.     

Riboflavin storage in earthworm chloragocytes and chloragocyte-derived eleocytes and its putative role as chemoattractant for immunocompetent cells

We have shown previously that riboflavin is stored in chloragosomes of chloragocyte-derived earthworm eleocytes what may have an adaptive value for worms vulnerable to soil-derived pathogen invasion. Thus, it is important to determine whether riboflavin stores are also present in earthworm species with low eleocyte counts, especially in the chloragocytes of their chloragogen tissue that surrounds the coelomic site of the intestine, and find out its putative roles in earthworm immunity. Experiments were performed on adult worms from 6 species. Freely floating coelomocytes (consisting of amoebocytes plus species-specific numbers of eleocytes) were extruded by mild electric shock and then chloragocytes from the same anaesthetized worm were mechanically detached into buffer. Both cell suspensions were analyzed by spectrofluorometry for riboflavin content. It turned out that riboflavin is stored in species-specific quantities in all investigated species. Riboflavin storage predominates in free coelomocytes of eleocyte-rich species (Eisenia andrei, Dendrobaena veneta, Allolobophora chlorotica) while in chloragocytes of species with few eleocytes (Lumbricus terrestris, L. rubellus, and Aporrectodea caliginosa). Upon a massive microbial impact, the coelomocytes (both amoebocytes and eleocytes) are involved in the formation of multicellular bodies encapsulating soil-derived pathogens, what is connected with enhanced coelomocyte mobility. Thus the second aim of the present investigation was to check if riboflavin can participate in coelomocyte accumulations. Tests performed in a 48-well chemotaxis chamber revealed that riboflavin behave as chemoattractant for coelomocytes of all investigated earthworm species, which may be one of mechanisms underpinning its immunomodulatory functions.     

Development of QSAR's in soil ecotoxicology: Earthworm toxicity and soil sorption of chlorophenols,chlorobenzenes and chloroanilines

Soil adsorption and the toxicity of four chloroanilines for earthworms were investigated in two soil types. The toxicity tests were carried out with two earthworm species, Eisenia andrei and Lumbricus rubellus. LC₅₀ values in mg kg^?1 dry soil were recalculated towards molar concentrations in pore water using data from soil adsorption experiments. An attempt has been made to develop Quantitative Structure Activity Relationships (QSAR's) using these results and data on five chlorophenols and dichloroaniline in four soils and five chlorobenzenes in two soils published previously (Van Gestel and Ma, 1988, 1990; Van Gestel et al., 1991). Significant QSAR relationships were obtained between 1) adsorption coefficients (log K _om ) and the octanol/water partition coefficient (log k _ow ), and 2) LC₅₀ values (in itμmol L^?1 soil pore water) and log K _ow . It can be concluded that both earthworm species tested are equally sensitive to chlorobenzenes and chloroanilines, E. andrei is more sensitive than L. rubellus to chlorophenols.     

Impact of five different tillage systems on soil organic carbon content and the density,biomass, and community composition of earthworms after a ten year period

To assess the impact of different types of soil tillage on the density, biomass, and community composition of earthworms, a long-term field study was performed in which soils were tilled in different ways for ten years. This study included five different types of tillage: (i) plough, (ii) grubber, (iii) disc harrow, (iv) mulch sowing, and (v) direct sowing. At the end of the experiment the earthworm density, biomass, and community composition, and the SOC (soil organic carbon) content were determined. The results show that density, biomass, and community composition of earthworm populations varied in relation to the type of soil tillage used. The density of anecic earthworm species decreased when soils were managed by conventional ploughing, relative to reduced tillage practices, whereas conversely the density of endogeic species increased. Additionally, the varying types of soil tillage influenced the abundance and biomass of different earthworm species in different ways. The density of Aporrectodea caliginosa was positively influenced by ploughing, whereas Aporrectodea longa, Lumbricus castaneus, and Satchellius mammalis showed a positive relationship to the grubber and Allolobophora chlorotica to direct sowing. We attribute these changes to modifications in the vertical distribution of SOC and varying potentials for mechanical damage of earthworms by tillage. A decrease in tillage intensity modified the vertical SOC distribution in the topsoil and consequently revealed positive effects on earthworm biodiversity, thus sustaining soil functioning.     

Earthworms as soil quality indicators in Brazilian no-tillage systems

It is well known that earthworm populations tend to increase under no-tillage (NT) practices, but abundances tend to be highly variable. In the present study, data from the literature together with those on earthworm populations sampled in six watersheds in SW Paraná State, Brazil, were used to build a classification of the biological soil quality of NT systems based on earthworm density and species richness. Earthworms were collected in 34 farms with NT aging from 3 to 27 yr, in February 2010, using an adaptation of the TSBF (Tropical Soil Biology and Fertility) Program method (hand sorting of five 20 cm × 20 cm holes to 20 cm depth). Six forest sites were also sampled in order to compare abundances and species richness with the NT systems. Species richness in the 34 NT sites and in the 6 forests ranged from 1 to 6 species. Most earthworms encountered were exotics belonging to the genus Dichogaster (D. saliens, D. gracilis, D. bolaui and D. affinis) and native Ocnerodrilidae (mainly Belladrilus sp.), all of small individual size. In a few sites, individuals of the Glossoscolecidae (P. corethrurus, Glossoscolex sp., Fimoscolex sp.) and Megascolecidae (Amynthas gracilis) families were also encountered, in low densities. Urobenus brasiliensis (Glossoscolecidae) were found only in the forest fragments. In the NT farms, earthworm abundance ranged from 5 to 605 ind m⁻² and in the forest sites, from 10 to 285 ind m⁻². The ranking of the NT soil biological quality, based on earthworm abundance and species richness was: poor, with <25 individuals per m⁻² and 1 sp.; moderate, with ≥25–100 individuals per m⁻² and 2–3 sp.; good, with >100–200 individuals per m⁻² and 4–5 sp.; excellent, with >200 individuals per m⁻² and >6 sp. About 60% of the 34 farms fell into the poor to moderate categories based on this classification, so further improvements to the NT farm's management system are needed to enhance earthworm populations. Nevertheless, further validation of this ranking system is necessary to allow for its wider-spread use.     

Movement response of Collembola to the excreta of two earthworm species: Importance of ammonium content and nitrogen forms

Several studies reported variable effects of earthworms on microarthropod density and variety. The present study tests the attraction of seven collembolan species belonging to four families, to the excreta of two earthworm species belonging to two families and two ecological categories, Aporrectodea giardi and Hormogaster elisae. Our objectives were (1) to better understand the impact of earthworms on the composition and density of Collembola communities, and (2) to dissect mechanisms involved in the attraction. Experiments were performed in Petri dishes containing two half-disks of filter paper, one with earthworm excreta, i.e. casts or a mix of mucus and urine, and the other with natural soil aggregates or water, respectively. Collembola were introduced half-way between the two half-disks and their number was counted on each half-disk and compared over 140 min. The content of ammonium in casts and mucus-urine of both earthworm species was analyzed to determine whether it altered the responses of Collembola faced with different types of earthworm excreta. The behaviour of Collembola varied strongly among the seven collembolan species, and with type of excreta and earthworm species. Six collembolan species were attracted to the mucus and urine of at least one earthworm species. The mucus-urine mixture of A. giardi, with low ammonium content, was generally more attractive than that of H. elisae, which was even repulsive in some cases, probably because of high levels of ammonium. The attraction to casts of the two earthworm species was less frequent and more variable. Folsomia candida was neither attracted to the casts nor to the mucus and urine of any earthworm species. Therefore, (1) earthworm species with different ecology, and different nitrogen excretion pathway impact differently the behaviour of collembolan species belonging to the same family or arising from the same habitat, and (2) variations in the sensitivity to ammonium among collembolan species partially explain the variable response of Collembola to earthworm excreta.     

Plant influences on native and exotic earthworms during secondary succession in old tropical pastures

Land-use changes can drastically alter earthworm communities. Native species are often lost and few exotic species, such as Pontoscolex corethrurus, rapidly prevail when tropical forests are converted to pastures. However, this process can be reversed when forests recover from abandoned pastures through secondary succession. We hypothesized (1) that the formation of forest floor mass during secondary succession in pastures promotes the recovery of native, anecic earthworms and (2) that the shift from grass vegetation in pastures to woody plants in secondary forests decreases the abundance and biomass of the exotic, endogeic P. corethrurus. To test the first hypothesis, we developed a litter manipulation experiment by removing and adding plant litter in plots of mature secondary forests in the Cayey Mountains, Puerto Rico. To test the second hypothesis we performed a greenhouse experiment to examine the influence of a pasture grass species Axonopus compressus and a dominating woody species Miconia prasina of the secondary forests on the number and biomass of the earthworm P. corethrurus. We found in the litter manipulation experiment that earthworm diversity, density and fresh weight were not affected by litter input. However, in the greenhouse experiment, A. compressus increased the number and biomass of P. corethrurus, whereas M. prasina decreased the exotic, endogeic earthworm. Our results suggest that the quantity of litter does not promote rapid changes (<1 year) in native, anecic earthworm diversity, and that the exotic, endogeic P. corethrurus is favored by grass A. compressus compared to the woody plant M. prasina. The shift in vegetation from grass to woody plants promotes the decrease in the density and biomass of the exotic, endogeic P. corethrurus during secondary succession in old tropical pastures.     

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

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

Vermicomposting of Vegetable Waste

? Vermicomposting is the degradation of organic waste through earthworm consumption, which converts the material into worm castings. Vermicomposting of vegetable waste was examined in order to identify suitable worm species and efficient levels of temperature and moisture. Laboratory experiments were conducted under controlled conditions using commonly available species: Pheretima sp.; Eisenia sp. and P. excavatus. Worms survived in the moisture range of 20-80 percent and the temperature range of 20°-40°C. Worm survival in decomposed and undecomposed organic wastes was also studied. Trials indicate that P. excavatus is the appropriate species for vegetable waste vermicomposting.     

Possible use of earthworm Eisenia fetida (Sav.) biomass for breeding aquarium fish

The aim of this research was to find further arguments advocating the idea of vermicomposting in household and school conditions. An alternative use of Eisenia fetida biomass was feeding it to aquarium fish (Poecilia reticulata). Fish were bred in two groups, each with six specimens of both genders. The control group consisted fish fed on standard food whilst the treatment group was fed on earthworm biomass. Compared to those fed on a standard food, earthworm-fed fish increased brood number significantly and produced twice the offspring (P < 0.001).     

Phosphatase activity in earthworm faeces

Wormcasts of Allolobophora caliginosa have been shown to have higher phosphatase activity than occurs in uningested soil, resulting in an increase in inorganic P released by mineralization of organic P. This paper describes similar observations on organic wastes which may be used for vermiculture. Besides A. caliginosa, three species which will feed on dung or other organic substrates were investigated, Eisenia foetida, Dendrobaena veneta and Lumbricus rubellus. When cow dung was used as a substrate, background phosphatase activity was too high for the effects of earthworm activity to be detected and a culture medium was therefore developed from sterilized paper waste sludge with added phytin. Phosphatase activity, assayed by a modified Hoffman method, was higher in the presence of all four earthworm species than in controls. It showed two peaks in relation to pH, at 3–5 and 9–10. The former is attributed to microbial activity and the latter to microbial activity or earthworm alkaline phosphatase. After 1 month, cultures of paper waste sludge with phytin contained about twice as much water soluble P when worms were present than in their absence.     

Riboflavin content in the coelomocytes of contrasting earthworm species is differentially affected by edaphic variables including organic matter and metal content

The riboflavin content in extruded coelomocyte lysates derived from Dendrodrilus rubidus may serve as a sensitive bioindicator of soil metal pollution: the vitamin (B2) content has previously been found to be high in worms from unpolluted soil but low in worms inhabiting Zn/Pb mine soils, aerially deposited Ni-contaminated soil, and in worms experimentally transferred from clean soil to the metalliferous field soils. The aim of the present work was to extend these observations by comparing the number and riboflavin composition of coelomocytes retrieved from three lumbricid species (Allolobophora chlorotica, Dendrobaena veneta, Eisenia andrei) after 4-week exposures to an unpolluted commercial soil, two geochemically contrasting unpolluted field soils, and two different Zn/Pb/Cd-polluted soils from the Bukowno district in South Poland. Whilst eco-physiologically contrasting, these three earthworm species share the trait of possessing relatively high numbers of eleocytes, a category of immune-competent coelomocyte rich in autofluorescent riboflavin. Spectroflurometric analysis indicated that coelomocyte riboflavin content in worms maintained in strongly metalliferous soils or in unpolluted sandy-clay and loamy-sand soils was increased in coleomocytes from epigeic D. veneta and E. andrei species, whilst was decreased in endogeic A. chlorotica. In conclusion, the riboflavin content of earthworm coelomocytes is affected in species-specific ways by edaphic variables, including organic matter and metal pollution.     

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.     

Species richness and zoogeographic affinities of earthworms in Cyprus

A new survey taken in the East Mediterranean island of Cyprus showed an increase in the number of earthworm species (from the six recorded earlier to 16). These species belong to two earthworm families: Lumbricidae (15 species) and Acanthodrilidae (one species). The majority of these species are autochthonous (57–64%) and five to seven species (36–44%) were introduced. The autochthonous species are (i) endemic of the Levant (Dendrobaena semitica), (ii) distributed in Anatolia and the Caucasus (D. pentheri), (iii) distributed from Europe up to Anatolia (Allolobophora nematogena, D. pantaleonis, Proctodrilus tuberculatus) (iv) distributed from Europe up to the Levant (Murchieona minuscula), (v) distributed in the Transcaucasian region and in the East Mediterranean (Helodrilus patriarchalis), or (vi) widely distributed along the Mediterranean basin (D. byblica, Eiseniella neapolitana, and Octodrilus complanatus). The data obtained indicate that the Cypriote earthworm fauna is significantly richer in species than previously thought and that it has more zoogeographic affinities with the Anatolian fauna than with those of Levant.     

The interactions between soil type and earthworm species determine the properties of earthworm casts

Earthworms are recognized to increase soil porosity, reorganize soil structure, and stimulate soil microflora and nutrient mineralization. The properties of earthworm casts should depend both on earthworm species or ecological group and on soil properties. Interactions between earthworm species and soil types have been suggested, but only poorly demonstrated. In order to better understand those interactions, two hypotheses led our study: (1) Soil type has a greater influence on cast properties than earthworm; (2) Earthworms from different species influence cast properties differently; (3) The intensity and direction of the impact of each earthworm species on cast properties vary with soil properties. Fifteen physical and chemical variables (N–NH₄⁺, N–NO₃⁻, total organic C and N, C/N ratio, CaCO₃, pH, P, K⁺, Mg²⁺, Mn²⁺, Na⁺, CEC, moisture, wettability) were measured in casts of three earthworm species (Lumbricus terrestris, Allolobophora chlorotica and Aporrectodea rosea) produced in three temperate soils. Univariate and multivariate analyses showed that earthworm species and soil types significantly impacted cast properties. pH, N_t, K and Mg contents were interactively altered by both factors. Multivariate analysis showed that a difference of soil type had a major impact on casts properties (62%) compared to the impact of a difference of earthworm species (10%). Cast properties were most impacted by L. terrestris, then by A. chlorotica and last by A. rosea. The response ratio (ratio of the properties of the casts to the properties of the bulk soil) was used to quantify the effect of earthworm species compared to the control soil. It showed a higher response of variables in casts in nutrient-rich soils, especially in casts of L. terrestris. The interactions between earthworm species and soil types on cast properties were discussed with regards to earthworm ecology, properties of the soil, and earthworm modifications of cast microflora.