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Livestock farming produces slurry, which is normally recycled by land-spreading; but this raises questions about the long-term impact of slurry on soil fauna and soil processes. To examine the long-term effects of slurry on earthworms, cattle and pig excreta slurries and inorganic fertiliser were applied to a ryegrass silage sward for 33 years in a replicated experiment, and earthworms sampled for four consecutive years at three seasonal intervals (March, May and October). Animal slurries were applied at three rates (50, 100 and 200 m3 ha−1 yr−1) and inorganic fertiliser as 200 kg N, 32 kg P and 160 kg K ha−1 yr−1. Epigeic earthworms, Lumbricus rubellus, Lumbricus juveniles, and the endogeic Allolobophora chlorotica responded strongly and positively to cattle slurry inputs. In the case of L. rubellus, biomass was five times greater in the highest input cattle slurry plots than in the untreated controls. In contrast to Al. chlorotica adults, other endogeic earthworms and juvenile Allolobophora were more associated with the inorganic fertiliser treatment. Aporrectodea juveniles, in particular, had significantly greater biomass in the inorganic fertiliser treatment. However, no such response was demonstrated by adult Aporrectodea caliginosa, the commonest species found, suggesting niche separation between adults and juveniles. Pig slurry applications produced relatively little change in earthworm biomass compared to controls, despite residual copper concentrations; suggesting that organic matter input was counterbalancing the sublethal effects of copper.

International audience; Recent EFSA (European Food Safety Authority) reports highlighted that the ecological risk assessment of pesticides needed to go further by taking more into account the impacts of chemicals on biodiversity under field conditions. We assessed the effects of two commercial formulations of fungicides separately and in mixture, i.e., Cuprafor Micro® (containing 500 g kg−1 copper oxychloride) at 4 (C1, corresponding to 3.1 mg kg−1 dry soil of copper) and 40 kg ha−1 (C10), and Swing® Gold (50 g L−1 epoxiconazole EPX and 133 g L−1 dimoxystrobin DMX) at one (D1, 5.81 10−2 and 1.55 10−1 mg kg−1 dry soil of EPX and DMX, respectively) and ten times (D10) the recommended field rate, on earthworms at 1, 6, 12, 18 and 24 months after the application following the international ISO standard no. 11268-3 to determine the effects on earthworms in field situations. The D10 treatment significantly reduced the species diversity (Shannon diversity index, 54% of the control), anecic abundance (29% of the control), and total biomass (49% of the control) over the first 18 months of experiment. The Shannon diversity index also decreased in the mixture treatment (both fungicides at the recommended dose) at 1 and 6 months after the first application (68% of the control at both sampling dates), and in C10 (78% of the control) at 18 months compared with the control. Lumbricus terrestris, Aporrectodea caliginosa, Aporrectodea giardi, Aporrectodea longa, and Allolobophora chlorotica were (in decreasing order) the most sensitive species to the tested fungicides. This study not only addressed field ecotoxicological effects of fungicides at the community level and ecological recovery, but it also pinpointed some methodological weaknesses (e.g., regarding fungicide concentrations in soil and statistics) of the guideline to determine the effects on earthworms in field situations.

Earthworms are widely known to impact soil health, having a keyrole in nutrient cycling and are often referred to as soil engineers.They are vital for soil microbial assemblages particularly through theirfeeding and burrowing activity in soil. Earthworms feed on soil organicmatter and litter, and the resulting casts alter the soil microbialcommunity. However, the gut microbiome of earthworms remains less known.In this study, we used amplicon sequencing of the 16S rRNA gene forbacteria and 18S rRNA gene for eukaryotes to assess the gut communityassemblages of earthworm species within three genera Aporrectodea,Allolobophora and Lumbricus that represent different life forms sharingthe same habitat. The objective was to compare the gut microbiomeprofiles of eukaryotic and prokaryotic organisms to assess significanceof earthworm life forms, and to explore the cross kingdom networks in anattempt to identify keystone species. We found a high eukaryoticdiversity with a dominance of the SAR supergroup along with fungi andmetazoan in the earthworm gut. The bacterial community were dominated bymembers of Proteobacteria, Acidobacteria, Actinobacteria, Firmicutes, andVerrucomicrobia. The eukaryotic and prokaryotic communities showedsimilar differences in alpha diversity, being lowest in Lumbricusherculeus. The beta diversity showed earthworm species as a key factor inshaping gut microbiomes with L. herculeus harboring distinct microbiomescompared to species of Aporrectodea caliginosa, A. longa, A. tuberculataand Allolobophora chlorotica. Cross kingdom networks showed highinteractions between several protist and bacterial OTUs. In conclusion,this study suggested that the community assemblages of gut microbiomeswas shaped by earthworm species and life form, and such assemblageconsist of cross kingdom interactions among eukaryotes and prokaryotes.

Allolobophora chlorotica chlorotica (Savigny, 1826) Enteriоn chlоrоticum Savigny, 1826: 182. Allоlоbоphоra chlоrоtica: ��apkarev 1980: 166; Stojanović 1996: 26; Stojanović & Karaman 2005a: 128; Milutinović et al. 2010: 629. Allоlоbоphоra chlоrоtica chlоrоtica: Mr��ić 1991: 211; Stojanović et al. 2008: 60; Csuzdi 2012: 97; Stojanović & Milutinović 2014: 308. E. Vаllеy mеаdоw, аnthrоpоgеnic biоtоpеs (Stоjаnоvić 1996); fоrеst sоils, cultivаtеd sоils, mеаdоws (Stоjаnоvić et al. 2008; Milutinоvić et al. 2010). D. Pаnnоniаn pаrt���Pаlić, Pаnčеvо, Zrеnjаnin (��аpkаrеv 1978); Bеlа Crkvа (��аpkаrеv 1978, 1980); Fru��kа gоrа, Plаtičеvо, Kаrаvukоvо, Turskе livаdе (Stоjаnоvić & Milutinоvić 2014). Bаlkаn pаrt: Bеоgrаd (��аpkаrеv 1972); Grоckа, Kаrаburmа, (��аpkаrеv 1980, 1989; Stоjаnоvić 1996); Mlаdеnоvаc, Kоrаćicа, Krаgujеvаc, Lаzаrеvаc, Tоpоlа, Arаnđеlоvаc (��аpkаrеv 1989); Knjаžеvаc, Nоvi Pаzаr, Ni��, Svrljig (��аpkаrеv 1980; Stоjаnоvić 1996); Kru��еvаc, Pirоt, Klisurе nеаr Vlаsinа, Lеpеnicа, Pеtrоvаčkа rеkа, Mоrаvа, Hоmоljе, Sjеnicа (Stоjаnоvić 1996); Jаstrеbаc (Stоjаnоvić & Kаrаmаn 2005а; Milutinоvić et al. 2010). Z. Pеrеgrinе spеciеs nаtivе in thе Pаlеаrctic., Published as part of Stojanovi��, Mirjana, Sekuli��, Jovana & Traki��, Tanja, 2018, Checklist of earthworms (Oligochaeta: Lumbricidae) from Serbia: a review, pp. 124-155 in Zootaxa 4496 (1) on page 126, DOI: 10.11646/zootaxa.4496.1.9, http://zenodo.org/record/1447067, {"references":["Savigny, J. C. (1826) Analyse des Travaux de l'Academie royale des Sciences, pendant l'annee 1821, partie physique. In: Cuvier, G. (Ed.), Memoires de l'Academie des Sciences de l'Institut de France, Paris, 5, pp. 176 - 184.","Sapkarev, J. (1980) Prilog poznavanju kisnih glista (Lumbricidae) SR Srbije. Zbornik radova faune Srbije, 1, 165 - 179.","Stojanovic, M. & Karaman, S. (2005 a) Further contribution to the knowledge of the earthworms of Sumadija. Archives of biological sciences, Belgrade, 57, 127 - 133.","Milutinovic, T., Avramovic, A., Pesic, S., Blesic, B., Stojanovic, M. & Bogdanovic, A. M. (2010) Contribution to the knowledge of pedofauna in Sumadija (Central part of Serbia). Biotechnology & Biotechnological Equipment, 24 (2), 628 - 635. https: // doi. org / 10.1080 / 13102818.2010.10817911","Mrsic, N. (1991) Monograph on earthworms (Lumbricidae) of the Balkans I, II. Slovenska Akademija Znanosti in Umetnosti, Razred za Naravoslovne Vede, Opera, Ljubljana, 31, 1 - 757.","Stojanovic, M., Milutinovic, T. & Karaman, S. (2008) Earthworm (Lumbricidae) diversity in the Central Balkans: An evaluation of their conservation status. European Journal of Soil Biology, 44, 54 - 67. https: // doi. org / 10.1016 / j. ejsobi. 2007.09.005"]}

Allolobophora chlorotica chlorotica (Savigny, 1826) Enterion chloroticum Savigny, 1826: 182. Allolobophora chlorotica: Zicsi & Michalis 1981: 248; 1993: 302. Michalis 1982: 349; 1987: 62. Michalis et al. 1985: 38; 1989: 5. Szederjesi & Csuzdi 2012: 29. Allolobophora chlorotica chlorotica: Szederjesi 2017. Widely introduced peregrine species (Csuzdi & Zicsi 2003). Distribution in Greece. Mainland: Thessaloniki, Kastoria, Naoussa, Imathia, Florina, Grevena, Kilki, Velvendos, Aiginio, Katerini (Michalis 1972). Florina (��apkarev 1972). Kaligoni, Patras (Zicsi 1973). Konitsa, Filiates, Metsovo, Megalo Peristeri (Zicsi 1974). Veroia (Michalis 1975a). Trikala, Agiokampos, Sykourion, Karditsa, Mouzaki (Michalis 1976). Lyggiades, Stavraki, Mpizani, Katsikas, Perama, Agios Ioannis, Vryssi Pappa, Eleousa, Kastritsa, Anatoli, Ioannina, Korinthos, Meteora, Deskati, Foteina, Nea Pella, Trikala, Edessa, Agras (Zicsi & Michalis 1981). Kato Milea, Palaeogratsano (Michalis 1983). Agia Foteini, Kerassies, Giannakochori, Agios Nikolaos, Agios Athanasios, Militsa, Rhodochori, Naoussa, Karydies, Fyteia (Michalis et al. 1985). Soufli, between Rizia and Orestiada (Michalis 1987). Aridea, Mavrovouni, Polycarpi, Sevastiana, Agras, Limnochori, Edessa, Logos (Michalis et al. 1989). Meteora, Kastraki, Korydallos, Larissa, Metsovo, Ktima Despoti, Lyggiades, Karyes, Konitsa, Kalpaki, Voidomatis, Nea Apollonia (Zicsi & Michalis 1993). Athamania (Szederjesi & Csuzdi 2012). Lefkas: Kaligoni, Fryni (Zicsi 1973). Kos: Asfendiou (Michalis 1975b, 1983). Crete: Garipa, Skalani (Szederjesi 2017)., Published as part of Szederjesi, T��mea, Vavoulidou, Evangelia, Chalkia, Christina, D��nyi, L��szl�� & Csuzdi, Csaba, 2017, An annotated checklist of earthworms of Greece (Clitellata: Megadrili), pp. 57-82 in Zootaxa 4272 (1) on page 58, DOI: 10.11646/zootaxa.4272.1.3, http://zenodo.org/record/583379, {"references":["Savigny, J. C. (1826) G. Cuvier: Analyse des Travaux de l'Academie royale des Sciences, pendant l'annee 1821, partie physique. Memoires de l'Academie des Sciences de l'Institut de France Paris, 5, 176 - 184.","Zicsi, A. & Michalis, K. (1981) Ubersicht der Regenwurm-fauna Griechenlands (Oligochaeta: Lumbricidae). Acta Zoologica Hungarica, 27, 239 - 264.","Michalis, K. (1982) Katalog der Oligochaetenfauna Griechenlands. Biologia Gallo-Hellenica, 9, 343 - 362.","Michalis, K., Fragoulis, A. & Panidis, S. (1985) Earthworms (Oligochaeta: Lumbricidae) from the montainous mas of Pella province: southwest region, and Imathias province, northwest region: a systematical and ecological study. Ecologia Mediterranea, 11 (4), 37 - 42.","Szederjesi, T. & Csuzdi, Cs. (2012) New and little known earthworm species form Greece (Oligochaeta: Lumbricidae, Acanthodrilidae). Zootaxa, 3304, 25 - 42.","Csuzdi, Cs. & Zicsi, A. (2003) Earthworms of Hungary (Annelida: Oligochaeta; Lumbricidae). In: Csuzdi, Cs. & Mahunka, S. (Eds.), Pedozoologica Hungarica. 1. Hungarian Natural History Museum, Budapest, pp. 1 - 271.","Michalis, K. (1972) Symvoli is tin meletin to Oligochaeton tis Makedonias. PhD Dissertation, University of Thessaloniki, Thessaloniki, 98 pp.","Sapkarev, J. (1972) Beitrage zur Kenntnis der Lumbricidenfauna Griechenlands. Macedonian Academy of Sciences and Arts, 4, 31 - 36.","Zicsi, A. (1973) Regenwurmer (Oligochaeta: Lumbricidae) aus Griechenland. Opuscula Zoologica Budapest, 12, 99 - 103.","Zicsi, A. (1974) Ein neue Dendrobaena - Art (Oligochaeta: Lumbricidae) aus Griechenland. Acta zoologica hungarica, 20, 449 - 451.","Michalis, K. (1975 a) Neue Angaben zur Fauna der Oligochaeten von Mazedonien. Scientific Annals, Faculty of Physics and Mathematics, University of Thessaloniki, 15, 201 - 211.","Michalis, K. (1976) Beitrag zur Kenntnis der Oligochaetenfauna Thessaliens. Scientific Annals, Faculty of Physics and Mathematics, University of Thessaloniki, 16, 153 - 168.","Michalis, K. (1983) New biotopes of the species Allolobophora dofleini (Ude, 1922) and the subspecies Bimastus antiquus bouchei (Zicsi-Michalis, 1981) (Oligochaeta-Lumbricidae). Ecologia Mediterranea, 9, 57 - 61.","Michalis, K. (1987) Contribution to the systematic and ecological study of the earthworms (Oligochaeta, Lumbricidae) of the Western Thrace. Arquivos do Museu Bocage, 1 (5), 59 - 69.","Michalis, K., Fragoulis, A. & Panidis, S. (1989) Notes on the earthworms (Oligochaeta, Lumbricidae) from central west Macedonia. Acta zoologica cracoviensia, 32, 3 - 14.","Zicsi, A. & Michalis, K. (1993) Zwei neue Dendrobaena - Arten aus Griechenland (Oligochaeta: Lumbricidae). Acta Zoologica Hungarica, 39, 301 - 310.","Michalis, K. (1975 b) Die Oligochaeten der Insel Kos. Scientific Annals, Faculty of Physics and Mathematics, University of Thessaloniki, 15, 187 - 200."]}

The limited dispersal ability of earthworms is expected to result in marked genetic isolation by distance and remarkable spatial patterns of genetic variation. To test this hypothesis, we investigated, using microsatellite loci, the spatial genetic structure of two earthworm species, Allolobophora chlorotica and Aporrectodea icterica, in two plots of less than 1 ha where a total of 282 individuals were collected. We used spatial autocorrelation statistics, partial Mantel tests of isolation-by-distance (IBD) and isolation-by-resistance (IBR), and Bayesian test of clustering to explore recent patterns involved in the observed genetic structure. For A. icterica, a low signal of genetic structure was detected, which may be explained by an important dispersal capacity and/or by the low polymorphism of the microsatellite loci. For A. chlorotica, a weak, but significant, pattern of IBD associated with positive autocorrelation was observed in one of the plots. In the other plot, which had been recently ploughed, two genetically differentiated clusters were identified. These results suggest a spatial neighbourhood structure in A. chlorotica, with neighbour individuals that tend to be more genetically similar to one another, and also highlight that habitat perturbation as a result of human activities may deeply alter the genetic structure of earthworm species, even at a very small scale. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114, 335–347.

Afforestation can improve the delivery of ecosystem services from reclaimed landfill sites. Tree health is a key determinant of ecosystem service delivery, and is directly impacted by soil quality; which is driven by biological processes in the soil, reliant on leaf litter inputs to function. Different tree species have different litter quality, affecting the degree to which they support biological processes in soils and the development of abundant and diverse soil faunal communities. In recognition of their key role in improving soil structure and fertility—key attributes of soil quality, earthworms have often been the subject of research as a part of land reclamation, and these organisms have displayed preferences for specific types of leaf litter. This work utilised a choice chamber design to measure the foliar material palatability of two tree species used in land restoration ( Alnus cordata and Acer platanoides ) as a food source for two common European earthworm species ( Aporrectodea longa and Allolobophora chlorotica ), and the effect of a reclaimed soil quality on earthworm growth, survival and feeding preferences. The research revealed that both earthworm species initially preferred the foliar material of A. cordata over A. platanoides, with the leaves of the latter requiring higher degradation to become palatable to earthworms. The consumption of fresh leaves showed these are a suitable food source for earthworms in choice chamber experiments, which historically have instead relied on senescent leaf litter. Finally, high survival rates of both A. longa and A. chlorotica in the reclaimed soil treatment, in addition to consumption of leaf material of two tree species now widely used on reclaimed landfill sites, demonstrated that these earthworm species are suitable candidates for inoculation to reclaimed land.

International audience; Aims : The passage of seeds through the earthworm gut potentially damages seeds, altering seed and seedling performances depending on seed traits. This work was conducted to study to what extent chemical and morphological seed traits determine the seed attractiveness for earthworms. Methods : We tested seed selection via the ingestion and digestion of 23 grassland plant species spanning a range of 14 morphological and chemical traits by two common earthworm species: the anecic Lumbricus terrestris and the endogeic Allolobophora chlorotica. Results : Both earthworm species ingested seeds from all plant species. A. chlorotica digested almost all ingested seeds (out of the 15 % ingested), whereas L. terrestris excreted them in varying quantities (out of the 86 % ingested), depending on plant species identity. Seed ingestion rate by L. terrestris was driven by seed oil content and earthworm initial weight. The apparent effect of seed length was explained via seed oil content. Seed digestion rate by L. terrestris was negatively impacted by seed size. Seed ingestion rate by A. chlorotica tended to be impacted by seed protein content and seed length. Conclusion : Earthworms–seed interactions depend on a variety of seed traits and earthworm identity. Thus, earthworms, via their specific feeding behavior, might facilitate or impede the regeneration of certain plant species and drive plant communities.

International audience; Landscape features are known to alter the spatial genetic variation of aboveground organisms. Here, we tested the hypothesis that the genetic structure of belowground organisms also responds to landscape structure. Microsatellite markers were used to carry out a landscape genetic study of two endogeic earthworm species, Allolobophora chlorotica (N = 440, eight microsatellites) and Aporrectodea icterica (N = 519, seven microsatellites), in an agricultural landscape in the North of France, where landscape features were characterized with high accuracy. We found that habitat fragmentation impacted genetic variation of earthworm populations at the local scale. A significant relationship was observed between genetic diversity (He, Ar) and several landscape features in A. icterica populations and A. chlorotica. Moreover, a strong genetic differentiation between sites was observed in both species, with a low degree of genetic admixture and high Fst values. The landscape connectivity analysis at the regional scale, including isolation by distance, least-cost path and cost-weighted distance approaches, showed that genetic distances were linked to landscape connectivity in A. chlorotica. This indicates that the fragmentation of natural habitats has shaped their dispersal patterns and local effective population sizes. Landscape connectivity analysis confirmed that a priori favourable habitats such as grasslands may constitute dispersal corridors for these species.

Earthworms can be troublesome pests on golf courses when their soil-rich casts become abundant enough to disrupt the maintenance, aesthetics, and playability of putting greens and fairways. Management of the problem is hindered by lack of knowledge of earthworm community structure on North American golf courses. We surveyed communities of endogeic earthworms inhabiting golf course fairways and putting greens in central Kentucky and tracked the seasonal population structure and casting activity of the predominant species. Seven earthworm species, six of them non-native, were identified in varying proportions from fairways of the different golf courses. Aporrectodea trapezoides (Duges) dominated, although Allolobophora chlorotica (Savigny), Diplocardia singularis (Ude), and Amynthas sp. also were abundant on some courses. Soil characteristics (pH, percentages of sand, silt, clay, and organic matter) at fairway sample sites were not good predictors of overall earthworm density or proportionate abundance of particular species. Ap. trapezoides was found in the upper 21 cm of topsoil throughout the year except when frozen ground precluded sampling. The Ap. trapezoides population consisted mainly of adults and cocoons in late autumn and winter, and juveniles in summer. Casting by Ap. trapezoides, which also occurs on soil-based greens, was greatest in late autumn and early winter, with a secondary peak in early spring. Amynthas hupeiensis, an east Asian megascolecid earthworm, was the only species found damaging sand-based greens. Prolific casting by Am. hupeiensis continued during summer after other species’ casting activity on fairways had waned. Am. hupeiensis is established along river banks in Kentucky and neighboring states, and is used as fish bait. We hypothesize that it may be introduced onto golf courses as cocoons in river sand used for course renovations, or from bait discarded by anglers fishing in golf course ponds.

Through their manipulation of soil physical and chemical properties, earthworms have the potential to increase nutrient mineralization and benefit crop production. However, this effect is dependent on numerous factors including earthworm densities and ecological strategies. In addition to impacting density, soil environmental conditions may limit earthworm activity to specific times during the growing season, moderating their effects on nutrient availability and crop growth. The purpose of this study was to determine the active period of earthworms to better assess their potential impact on crop productivity in dryland wheat production systems in the Palouse region of northern Idaho and eastern Washington, U.S.A. Earthworm community density, age structure, diversity, and soil moisture and soil temperature were monitored over a 14-month (April 2014–June 2015) period in six dryland wheat fields. All earthworms collected were exotic species, with the endogeic species, Aporrectodea trapezoides being predominant at all sites (87% of all adults identified). Allolobophora chlorotica, Lumbricus terrestris and Aporrectodea tuberculata were collected at lower frequencies (1.8–6.4% of adults). Mean densities ranged from 14 to 75 individuals m−2, and populations transitioned to aestivation beginning in mid-June of both years. All earthworms were active (no aestivating individuals) on 56.9% of the sampling dates in 2014 and 71% of the days in 2015. A short active period and average densities at most sites of less than 100 individuals per m2 suggest that the impact of earthworms on crop growth may be limited in conventional, dryland agricultural fields of this region.

Short Rotation Forestry (SRF) practice which includes use of rapidly growing native and non-native tree species has been introduced to the UK as a method to increase woody biomass production. A largely unknown aspect of SRF is the quality of leaf litter, and its palatability to the soil decomposer community of which earthworms are a major component. The aims of the present study were to investigate and compare the preference of selected native British earthworms for selected SRF species litter. These were addressed through a series of controlled laboratory experiments. Choice chambers were used to quantify litter removal by Allolobophora chlorotica, Aporrectodea caliginosa, Aporrectodea longa and Lumbricus terrestris over a period up to five weeks. In addition, an infrared webcam recording technique was used to directly observe litter selection behaviour of L. terrestris under cover of darkness. Choice chamber experiments revealed that earthworms significantly preferred (p

We tested experimentally if photoautotrophic microorganisms are a carbon source for invertebrates in temperate soils. We exposed forest or arable soils to a 13 CO 2 -enriched atmosphere and quantified 13 C assimilation by three common animal groups: earthworms (Oligochaeta), springtails (Hexapoda) and slugs (Gastropoda). Endogeic earthworms ( Allolobophora chlorotica ) and hemiedaphic springtails ( Ceratophysella denticulata ) were highly 13 C enriched when incubated under light, deriving up to 3.0 and 17.0%, respectively, of their body carbon from the microbial source in 7 days. Earthworms assimilated more 13 C in undisturbed soil than when the microbial material was mixed into the soil, presumably reflecting selective surface grazing. By contrast, neither adult nor newly hatched terrestrial slugs ( Deroceras reticulatum ) grazed on algal mats. Non-photosynthetic 13 CO 2 fixation in the dark was negligible. We conclude from these preliminary laboratory experiments that, in addition to litter and root-derived carbon from vascular plants, photoautotrophic soil surface microorganisms (cyanobacteria, algae) may be an ecologically important carbon input route for temperate soil animals that are traditionally assigned to the decomposer channel in soil food web models and carbon cycling studies.

Within the regulatory framework for authorisation of plant protection products (PPPs) (EU Directive 91/414/1991/EEC replaced by Regulation (EC) 1107/2009), higher tier risk assessments and earthworm field tests are conducted in different countries across Europe. This paper describes dominant earthworm species for regulatory and biogeographical regions in agricultural landscapes across Europe and examines regional differences in earthworm communities and densities and their respective response to a toxic reference. For the assessment of earthworm abundance and species distribution, data of untreated control plots from 30 earthworm field studies were analysed; each conducted according to the ISO 11268–3 (1999) guideline by European Crop Protection Association member companies in the context of registration of PPPs. For the evaluation of the response to PPPs under different regional and climatic conditions, the effect on earthworm abundance was assessed by comparing plots treated with toxic references with untreated control plots. Additionally, a comparative literature review was included providing an overview of earthworm species composition and densities in agricultural crops from 14 European countries. The assessment of earthworm field studies from six different European countries revealed that common earthworm species of anecic and endogeic ecological groups are present at most field sites. Dominant species groups of endogeic and anecic earthworms can be defined that are abundant in all assessed countries. These are the endogeic species Aporrectodea caliginosa, Aporrectodea rosea and Allolobophora chlorotica, and the anecic species Lumbricus terrestris (Northern and Central Europe) and Lumbricus friendi (Southern Europe). Taking into account the high variability in total earthworm abundances, it can be concluded that the variability within regions was larger than the variability between regions. Analysis of the earthworm community and data of toxic references lead to the conclusion that testing in different zones is not considered necessary.

Earthworm immune-competent cells (coelomocytes) can be adversely affected by soil metal contamination. The aim of the present paper was to perform comparative studies on the coelomocytes of four lumbricid species dermally exposed for 2 days to filter paper soaked with Ni, Cu, Zn, Cd, or Pb chlorides (1 mM). Coelomocytes were expelled by electro-stimulation, counted, and assayed ex vivo by flow cytometry and spectrofluorometry. The coelomocytes of Allolobophora chlorotica and Dendrobaena veneta contain both amoebocytes and riboflavin-storing eleocytes; consequently, they were subjected to assays based on flow cytometric quantification of autofluorescent eleocytes and spectrofluorometric measurement of riboflavin content in coelomocyte lysates. In Al. chlorotica, the number of coelomocytes, the percentage of eleocytes, and the amount of riboflavin were significantly lower in Cu-exposed worms although these cytometric parameters were less affected by Ni, Zn, Cd exposure, and entirely unaffected by Pb exposure. In D. veneta, such cellular effects were also only observed in Cu-exposed worms. The coelomic fluids of Aporrectodea caliginosa and Lumbricus rubellus are constitutionally impoverished of eleocytes; therefore, these species could not be subjected to assays measuring the riboflavin content of these granular cells. Rather, the amoebocytes of Ap. caliginosa and L. rubellus were subjected to flow cytometric measurements of in vitro neutral red uptake (NR). However, the NR uptake assay was technically demanding, requiring a strictly normalized incubation period over all samples to yield useful comparative data. In contrast, the riboflavin content in the coelomocyte lysates of eleocyte-rich species appears to be a robust and convenient immune-function biomarker of environmental stress.

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.

The introduction of earthworms into soils contaminated with metals and metalloids has been suggested\ud to aid restoration practices. Eisenia veneta (epigeic), Lumbricus terrestris (anecic) and Allolobophora\ud chlorotica (endogeic) earthworms were cultivated in columns containing 900 g soil with 1130, 345, 113\ud and 131 mg kg1 of As, Cu, Pb and Zn, respectively, for up to 112 days, in parallel with earthworm-free\ud columns. Leachate was produced by pouring water on the soil surface to saturate the soil and generate\ud downflow. Ryegrass was grown on the top of columns to assess metal uptake into biota. Different\ud ecological groups affected metals in the same way by increasing concentrations and free ion activities in\ud leachate, but anecic L. terrestris had the greatest effect by increasing leachate concentrations of As by\ud 267%, Cu by 393%, Pb by 190%, and Zn by 429% compared to earthworm-free columns. Ryegrass\ud grown in earthworm-bearing soil accumulated more metal and the soil microbial community exhibited\ud greater stress. Results are consistent with earthworm enhanced degradation of organic matter leading\ud to release of organically bound elements. The degradation of organic matter also releases organic acids\ud which decrease the soil pH. The earthworms do not appear to carry out a unique process, but increase\ud the rate of a process that is already occurring. The impact of earthworms on metal mobility and\ud availability should therefore be considered when inoculating earthworms into contaminated soils as\ud new pathways to receptors may be created or the flow of metals and metalloids to receptors may be\ud elevated.

Metal pollution affects earthworm coelomocytes, including their differential counts, riboflavin content and metallothioneins (MT) involved in metal homoeostasis and detoxification. The present work shows effects of Ni, Cu, Zn, Cd, and Pb at the same molarity (1 mM) on coelomocytes of Allolobophora chlorotica after 2-day worm dermal exposure to metal chlorides. Numbers of coelomocytes/eleocytes extruded by electric shock and amounts of riboflavin in coelomocyte lysates were significantly decreased in Cu-exposed worms, less diminished in response to Ni, Zn, Cd, and unaffected by Pb. In sharp contrast, real-time PCR revealed a very strong (272 fold) MT-mRNA induction in response to Cd only. The induction was very low in response to Zn, Cu, Pb, and Ni ions (2.6, 2.1, 1.4, and 1.3-fold, respectively). In conclusion, decreased cell counts and riboflavin content are molecular biomarkers of Cu exposure while induction of MT-mRNA is a molecular biomarker of worm Cd exposure.

A neotype is designated for the large and ecologically interesting species of Japanese earthworm, Drawida hattamimizu Hatai, 1930. Its morphological redescription is unambiguously combined with the neotype’s sequence of the Cytochrome Oxidase I (COI) locus of the mitochondrial DNA, the fi rst time an earthworm species’ type has been thus profi led. Probably it is an introduced exotic rather than a translocated native, with a patchy distribution that appears only partly defi ned in Japan where it is both a restricted and an endangered listed species. Brief comparison of sympatric Drawida japonica (Michaelsen, 1892) to the type-species Drawida barwelli (Beddard, 1886) – and this latter from Shiga appears as a new record for Japan – allows the diagnosis of Drawida Michaelsen, 1900 to be amended slightly. Th e contentious issue of molecular ‘cryptic species’ is queried in relation to the lack of molecular data from type-specimens, the unique name-bearing references employed in zoological nomenclature. Without such reference, neither eco-taxonomic nor genomic studies of earthworm taxa can progress. In this regard, questions are raised concerning the molecular identities and provisional divergences of cosmopolitan generotypes Allolobophora chlorotica chlorotica (Savigny, 1826), the Aporrectodea caliginosa (Savigny, 1826) species-complex sensu Blakemore (2002), and of ecotoxicological standard test-species icon Eisenia fetida fetida (Savigny, 1826). Resurrection of their respective synonyms is mooted. Resolution of relationships within and between earthworm genera and families without DNA testing of the representative type-species and type-genera is fl agged as another crucial concern. ZooKeys 41: 1–29 (2010) doi: 10.3897/zookeys.41.374 www.pensoftonline.net/zookeys Copyright R.J. Blakemore et al.. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. RESEARCH ARTICLE Launched to accelerate biodiversity research A peer-reviewed open-access journal

In Canada, the influence of earthworms on soil is considered particularly important in the Southern regions where agriculture has developed. A field experiment was duplicated on a clay loam and a sandy loam soil in the Saint-Lawrence Valley. Both sites were enriched with urban compost or mineral amendment. The effects of amendments on soil properties and the earthworm community were evaluated by uni- and multivariate analyses at two soil depths (0–10 and 10–20 cm) and after two growth seasons. Compost-amended plots had higher pH, wet aggregates stability, total N and C, OM, K and Ca in the 0–10 cm soil layer than plots receiving mineral fertilizer. Two endogeic species, Aporrectodea caliginosa and Allolobophora chlorotica were dominant in both sites. Other species were all epigeic and were rare. Abundance and biomass of earthworms (0–10 cm) were significantly higher in clay loam, with a similar proportion in A. caliginosa (263 ind m −2 ) and A. chlorotica (228 ind m −2 ). In sandy loam, A. caliginosa was the most abundant (18 ind m −2 ), A. chlorotica was scarce (1.3 ind m −2 ) and epigeic species were absent. The density and biomass of earthworms were significantly higher with compost amendment, up to 12 times in clay loam: A. chlorotica reached a mean density of 420 ind m −2 vs. 36 ind m −2 with mineral amendments, A. caliginosa 344 ind m −2 vs. 183 ind m −2 and epigeic earthworms 19 ind m −2 vs. 12 ind m −2 . In co-inertia analysis, the main factors explaining data heterogeneity were soil texture (axis 1) and organic matter content (axis 2). A. chlorotica was associated with OM, while A. caliginosa was associated with silt and clay contents. Additionally, soil wet aggregate stability was the variable that best correlated with endogeic species. The overall earthworm population was low, which is common for tilled agricultural soils, but clearly benefited from the OM input from the urban compost, providing further evidence that fertilization based on organic residues, as implemented in organic agriculture, is highly beneficial for earthworms and soil quality.

Summary Soil biota such as earthworms, arbuscular mycorrhizal fungi (AMF) and plant roots are known to play a major role in engineering the belowground part of the terrestrial ecosystems, thus strongly influencing the water budget and quality on earth. However, the effect of soil organisms and their interactions on the numerous soil physical properties to be considered are still poorly understood. Shrinkage analysis allows quantifying a large spectrum of soil properties in a single experiment, with small standard errors. The objectives of the present study were, therefore, to assess the ability of the method to quantify changes in soil properties as induced by single or combined effects of leek roots ( Allium porrum ), AMF ( Glomus intraradices ) and earthworms ( Allolobophora chlorotica ). The study was performed on homogenised soil microcosms and the experiments lasted 35 weeks. The volume of the root network and the external fungal hyphae was measured at the end, and undisturbed soil cores were collected. Shrinkage analysis allowed calculating the changes in soil hydro-structural stability, soil plasma and structural pore volumes, soil bulk density and plant available water, and structural pore size distributions. Data analysis revealed different impacts of the experimented soil biota on the soil physical properties. At any water content, the presence of A. chlorotica resulted in a decrease of the specific bulk volume and the hydro-structural stability around 25%, and in a significant increase in the bulk soil density. These changes went with a decrease of the structural pore volumes at any pore size, a disappearing of the thinnest structural pores, a decrease in plant available water, and a hardening of the plasma. On the contrary, leek roots decreased the bulk soil density up to 1.23 g cm −3 despite an initial bulk density of 1.15 g cm −3 . This increase in volume was accompanied with a enhanced hydro-structural stability, a larger structural pore volume at any pore size, smaller structural pore radii and an increase in plant available water. Interestingly, a synergistic effect of leek roots and AMF in the absence of the earthworms was highlighted, and this synergistic effect was not observed in presence of earthworms. The structural pore volume generated by root and AMF growth was several orders of magnitude larger than the volume of the organisms. Root exudates as well as other AMF secretion have served as carbon source for bacteria that in turn would enhance soil aggregation and porosity, thus supporting the idea of a self-organization of the soil–plant–microbe complex previously described.

Summary The soil seed bank functions to escape unfavourable environmental conditions and seed predation. Anecic earthworms are increasingly recognised as important dispersers and predators of plant seeds. In contrast, the role of the usually more abundant endogeic earthworms which live and feed in the soil on plant seeds is largely unknown. We tested whether (A) endogeic earthworms ( Aporrectodea rosea , Allolobophora chlorotica , Octolasion tyrtaeum , Aporrectodea caliginosa ) ingest and digest grassland plant seeds ( Phleum pratense , Bellis perennis , Trifolium repens , Poa trivialis , Plantago lanceolata , Medicago varia ), (B) the passage of seeds through the gut of endogeic earthworms modifies seed germination, and (C) excreta (mucus and casts) of endogeic earthworms modify seed germination. As a reference effects of the well-studied anecic species Lumbricus terrestris were determined. Endogeic earthworms ingested and digested all of the studied plant seeds; however, both ingestion and digestion were earthworm and plant species specific. Moreover, passage through the gut of endogeic earthworms and their excreta modified plant seed germination (gut passage: B. perennis ; excreta: Ph. pratense and Pl. lanceolata ). The results indicate that endogeic earthworms may strongly impact the composition of the soil seed bank and, consequently, plant community assembly via direct and indirect effects on plant seeds. Since post-dispersal seed predation is a key factor for the structure of plant communities, with the effect on seed survival potentially exceeding that of pre-dispersal predation, seed predation and changes in germination of seeds by endogeic earthworm species is likely to drive plant community composition.

Bioturbation of metal contaminated soils contributes considerably to redistribution and surfacing of contaminated soil from deeper layers. To experimentally measure the contribution of Allolobophora chlorotica, Aporrectodea caliginosa, Lumbricus rubellus and L. terrestris to soil surface casting, a time-course experiment was performed under laboratory conditions. Earthworms were incubated in perspex columns filled with sandy soil (2% organic matter, 2.9% clay) or loamy clay soil (15% organic matter, 20% clay), and surface casts were collected after up to 80 days. On the sandy soil, A. caliginosa and L. rubellus brought approximately 7.1–16 g dry wt. casts/g fresh wt. earthworm to the surface, which is significantly more than A. chlorotica and L. terrestris (2.5–5.0 g dry wt./g fresh wt.). A. caliginosa was the only species that produced significantly more surface casts in the sandy soil than in the loamy clay soil. In the loamy clay soil, no differences in biomass-corrected casting rates were found among the species. Surface casting rates tended to decrease after 20 days. Considering the densities of the different species in a Dutch floodplain area Afferdensche and Deestsche Waarden, surface cast production is estimated to amount to 2.0 kg dry soil/m2 after 80 days, which could be extrapolated to 2.7–9.1 kg/m2 per year. These amounts correspond to a surface deposition of a layer of approximately 1.9–6.5 mm/year, which is of the same order or even slightly higher than the sedimentation rate and much higher than the amount of soil brought to the soil surface by bioturbating small mammals.

An investigation of earthworms across a sand dune system in NW England examined species distribution and abundance with respect to soil physico-chemical conditions and management. Replicated 0.1 m2 quadrats were examined every 50 m along 700 m transects across areas of vegetation succession and samples were then taken every 10 m where earthworms were first encountered. A translocation of Aporrectodea longa assessed the ability of this species to live in soils with a high sand content and laboratory work examined growth, maturation and survival of this species in soils with increasing proportions of sand (0–100%). Nine earthworm species were found on the dunes, but none in yellow dunes where organic matter (OM) content was

Earthworms are known to influence water infiltration in soils, but most of the existing knowledge relates to grasslands and arable systems; little is known on the role of earthworms for water infiltration in forests. We studied earthworm populations and water infiltration rates in woodland flooding sites used for groundwater recharge and the production of drinking water. Intensive flooding may detrimentally affect earthworm populations and simultaneously result in clogging of the topsoil, with the latter being a common problem in groundwater recharge systems. However, clogging does not occur at our study site, the “Lange Erlen” (Basel, Switzerland) and total earthworm numbers and biomass in flooded sites exceeded those of non-flooded sites (+51% and +71%, respectively). Total earthworm numbers ( r = 0.85***), numbers of endogeic ( r = 0.64*) and epigeic ( r = 0.81**) earthworms and numbers of two species ( Lumbricus rubellus, r = 0.62* and Allolobophora chlorotica, r = 0.77**) significantly correlated with water infiltration rates. The results suggest that short-term flooding (max. 10 days) interrupted by longer recovery periods favor earthworm populations which likely contribute to the long-term (ca. 100 years) sustainability of the studied forest groundwater recharge system and thereby to effective and cost efficient drinking water production.

Earthworm (Lumbricidae) populations were assessed in three replicated, field experiments with different management intensities conducted on grassland research farms on medium-textured mineral soils, located at Solohead (Co. Tipperary), Grange (Co. Meath) and Johnstown Castle (Co. Wexford), Ireland. The experiment at Solohead involved four levels of mineral fertilizer application (80, 175, 225 and 350 kg N ha−1), and the plots were rotationally grazed by dairy cows. Three levels of mineral fertilizer application were compared at Johnstown Castle (0, 225 and 390 kg N ha−1) and two at Grange (100 and 225 kg N ha−1), and the plots at these sites were rotationally grazed by suckler cows and followers or dry cattle. Between 10 and 15 earthworm species were recorded per site, with Allolobophora chlorotica and Aporrectodea caliginosa being dominant. Other abundant species included Aporrectodea rosea, Aporrectodea longa, Lumbricus terrestris, Lumbricus friendi, Lumbricus festivus, Lumbricus rubellus and Satchellius mammalis. Mean earthworm population densities per treatment ranged from 203 to 324 individuals m−2 and biomass from 60 to 176 g m−2. ANCOVA revealed significant overall positive relationships between management intensity (as indicated by N application rate) and earthworm abundance (F = 4.47, p

To get a better understanding of earthworm' responses towards flooding, three laboratory experiments were performed with the species Allolobophora chlorotica, Aporrectodea caliginosa and Lumbricus rubellus. Flooding response was determined in a pot experiment, in which the earthworms were incubated for 42 days in flooded or non-flooded soil, with or without heavy metal pollution. To determine moisture preference, earthworms were incubated for 9 days in aquaria with five compartments, containing soil with a moisture gradient (%, w/w), ranging from 35%, 45% (field capacity), 55%, 65% (saturated) to 65%+ (saturated and an extra water layer). Effects on earthworm health were studied by incubating earthworms of each species for 42 days in soil with the same range of moisture contents and determining the dry/wet weight ratio and dry weight gain as an indication of earthworm health. A. chlorotica was tolerant to water, although the worms tended to escape from flooded soil. Their health was significantly lower in the flooded soils (P

Earthworms exposed dermally to heavy metals can accumulate copper, lead, and cadmium avidly, but zinc is efficiently regulated. Allolobophora chlorotica is more sensitive to Cu, and less sensitive to both Pb and Cd ions than Eisenia fetida . During dermal exposure to heavy metals, the elemental composition of E. fetida immunocompetent cells, coelomocytes, is disrupted, including time-dependent fluctuations in intracellular metal content and Ca/P and S/P balance. In coelomocytes of the both species, all investigated heavy metals induce upregulation of the protective stress proteins (HSP70, HSP72, and metallothionein w-MT2), while an upregulation of caspase-3, a key molecular component of the apoptotic programmed cell death pathway, is induced by Cu, Pb, Cd, but not Zn exposure. In the both species, the number of coelomocytes is significantly reduced in a time dependent manner in Cu, Pb, and Cd exposed animals, but not in those exposed to Zn. In conclusion, coelomocytes homeostasis is significantly disrupted, at least transiently, in coelomocytes from earthworms subjected to metal stressors under laboratory conditions. The overall similarity in the responses of the epigeic E. fetida and endogeic A. chlorotica indicates that E. fetida is a reasonable surrogate for soil-dwelling earthworm species in short-duration laboratory-based ecotoxicological tests.

Specific composition of earthworm community has indicative value for evaluating the impact of agricultural practice on soil. The occurrence of species only like Aporrectodea caliginosa, Aporrectodea rosea, Lumbricus rubellus tolerant to disturbance is the result of intensive tillage and agricultural practice or the influence of strong limiting ecological factor. A community including more sensitive species Lumbricus terrestris and Aporrectodea longa, or the most sensitive species Allolobophora chlorotica and Lumbricus castaneus, indicates more favourable conditions of habitat.

Coelomic fluid and cells (coelomocytes) were extruded by the electric shock (1 min, 4.5 V) from field-collected individuals of Allolobophora chlorotica (Sav.). Coelomocytes were subjected to bright field and fluorescence light as well as electron microscopy (transmission and scanning). Results of all methods combined, revealed two distinct populations of circulating cells in the coelomic fluid: (1) autofluorescent eleocytes/chloragocytes filled with numerous large granules (chloragosomes) containing fluorophores, mainly riboflavin; and (2) amoebocytes, which can be differentiated into: (a) hyaline amoebocytes, firmly spreading in form of crinoline-like membranes; (b) granular amoebocytes with numerous dark granules spreading in form of fine pseudopodia. Flow cytometry confirmed the presence of highly complex autofluorescent cells (eleocytes/chloragocytes) and two cohorts of not-autofluorescence much less complex amoebocytes.

The levels of the enzymes, glutathione S -transferase, catalase, NAD(P)H-cytochrome c reductases, and DT-diaphorase were determined and compared in the tissues of three invertebrates commonly used in monitoring environmental quality: a freshwater mussel, Dreissena polymorpha , the earthworm Allolobophora chlorotica and the fourth instar of Chironomus riparius . It was found that the activities of GST, catalase, and NAD(P)-cytochrome c reductases were comparable in A. chlorotica and C. riparius , whereas comparatively a higher GST and a lower catalase activity was determined in the mussel tissues. DT-diaphorase was not detectable in A. chlorotica and the C. riparius larvae tissues, whereas this enzyme is present in the gills and the rest of soft mussel tissues (soft mussel tissues minus gills). It is suggested that the relatively low catalase activity observed in the tissues of the latter organism might be compensated by the presence of the antixidant role of DT-diaphorase. In addition, the inducibility of DT-diaphorase in D. polymorpha , by butylated hydroxyanisole (BHA) and lead (Pb) was investigated. Despite the bioaccumulation of both BHA (5.2±0.14 μg g −1 wet weight) and Pb (233.7±0.95 mg kg −1 dry weight) in the soft mussel tissues, the mussel DT-diaphorase was not induced. Although the activity of NADPH-cytochrome c (P-450) reductase was also not affected by these reagents, its activity was 2-fold higher in the gills than the rest of soft mussel tissues.

Earthworms are crucial for production and maintenance of soil structure and their activities can strongly impact soil functioning ( e.g. water regulation, nutrient dynamics). This laboratory study investigated the bioturbation activity of three endogeic species, Allolobophora chlorotica , Allolobophora icterica and Aporrectodea caliginosa , as affected by different locations of organic matter (OM) in the soil profile: OM scattered on the soil surface (surface-OM) or homogeneously mixed into the soil (mixed-OM). Microcosms, each containing a combination of one species (three individuals) and one OM location, were subjected to controlled environmental conditions (temperature, humidity and day/night cycle) for 60 days. At the end of the experiment, microcosms were cut into multiple horizontal cross-sections every centimetre and bioturbation activities were analyzed based on the number of burrows, the burrowed area and the percentage of burrowed area totally refilled with casts. Results showed that regardless of species, there was significantly fewer burrows and a greater percentage of burrowed area refilled with casts under mixed-OM than under surface-OM. A. chlorotica and A. caliginosa had a significantly greater burrowed area under mixed-OM than under surface-OM. Regardless of OM location, as depth increased, burrow number and area decreased for A. chlorotica and generally increased for A. icterica . In contrast, burrowing activity of A. caliginosa was affected by OM location as depth increased: under mixed-OM, burrow number decreased but burrowed area remained constant, whereas under surface-OM, burrow number remained constant and burrowed area increased. These results improve understanding of effects of endogeic species on soil structure and highlight effects of OM location on earthworm bioturbation. Especially this study gives information about the burrowing activity of A. icterica which has so far been little documented, and also informs about refilled burrows which is a major parameter for soil functioning.

International audience; Earthworm burrow systems are generally described based on postulated behaviours associated with the three ecological types. In this study, we used X-ray tomography to obtain 3D information on the burrowing behaviour of six very common anecic (Aporrectodea nocturna and Lumbricus terrestris) and endogeic (Aporrectodea rosea, Allolobophora chlorotica, Aporrectodea caliginosa, Aporrectodea icterica) earthworm species, introduced into repacked soil cores for 6 weeks. A simple water infiltration test, the Beerkan method, was also used to assess some functional properties of these burrow systems. Endogeic worms make larger burrow systems, which are more highly branched, less continuous and of smaller diameter, than those of anecic worms. Among the anecic species, L. terrestris burrow systems are shorter (9.2 vs 21.2 m) with a higher number (14.5 vs 23.5) of less branched burrows (12.2 vs 20.2 branches m(-1)), which are also wider (7.78 vs 5.16 mm) than those of A. nocturna. In comparison, the burrow systems made by endogeic species appeared similar to each other. However, A. rosea burrows were short and narrow, whereas A. icterica had a longer burrow system (15.7 m), more intense bioturbation intensity (refilled macropores or soil lateral compaction around them) and thus a greater number of burrows. Regarding water infiltration, anecic burrow systems were far more efficient due to open burrows linking the top and bottom of the cores. For endogeic species, we observed a linear relationship between burrow length and the water infiltration rate (R (2) = 0.49, p < 0.01). Overall, the three main characteristics significantly influencing water infiltration were burrow length, burrow number and bioturbation volume. This last characteristic highlighted the effect of burrow refilling by casts

Allolobophora chlorotica exists as two colour morphs, pink and green. Field observations have indicated that the two morphs have ecological preferences linked to soil moisture: the green morph dominating in wet soils and the pink morph in dry soils. The aim of this laboratory-based research was to investigate the potential differences in fitness and adaptation to soil-moisture conditions of the two morphs measured in terms of growth rate, reproductive output and cocoon viability. An initial experiment maintained hatchlings of both morphs individually under standard culture conditions. On maturation, these were paired (intra-morph), and cocoon production, viability and incubation time were determined. The green morph had significantly faster (P 0.05) by soil moisture. In contrast, lower soil-moisture content significantly (P

The river floodplain ‘Afferdense and Deestsche Waarden’ (ADW) in The Netherlands is diffusely contaminated with several heavy metals. It is, however, unclear whether this mixed contamination exerts any adverse ecotoxicological effects. In November 2000 and May 2001 a field survey was conducted in two areas in the ADW to collect a wide range of data concerning contamination levels, bioavailability, enchytraeids and earthworms and abiotic factors such as lutum and organic matter content, cation exchange capacity (CEC) and soil nutrient concentrations. Earthworms and enchytraeids were also analyzed for heavy metal content. At both sites arsenic and zinc were present in soil at relatively high concentrations (above the Dutch intervention value). In the two areas, both enchytraeids and earthworms accumulated metals. Fridericia ulrikae accumulated more cadmium than Enchytraeus buchholzi and Henlea perpusilla. The earthworm Lumbricus rubellus accumulated larger concentrations of Cr, Cu and Pb than Aporrectodea caliginosa and Allolobophora chlorotica. Dietary, physiological and behavioral characteristics may have contributed to these differences.

Allolobophora chlorotica (Savigny, 1826) Enterion chloroticum Savigny, 1826: 182. Enterion virescens Savigny, 1826: 182. Lumbricus riparius Hoffmeister, 1843: 189. Lumbricus communis v. luteus Hoffmeister, 1845: 29. Allolobophora chlorotica: Vejdovsk�� 1884: 60; Michaelsen 1890: 13; Wilcke 1949: 177; Zuck 1951: 127; Graff 1953 b: 26; Bouch�� 1972: 263; Easton 1983: 475; Sims & Gerard 1999: 50; Csuzdi & Zicsi 2003: 50; Blakemore 2008: 15. Helodrilus (Allolobophora) chloroticus: Michaelsen 1900 a: 486. Helodrilus chloroticus: Michaelsen 1903: 178. Ecology. Endogeic species occurring mostly on arable and grassland sites with low soil organic matter content (��� 4 %) (J��nsch et al. 2013), preference for moist habitats (Wilcke 1949, Graff 1954). Zoogeographical distribution type. A peregrine species, native to the Palaearctic, but introduced into New Zealand, North- and South America (Csuzdi & Zicsi 2003). Distribution in Germany. Central Germany (Eggert 1982; Westernacher-Dotzler 1988; PSMS in Saxony- Anhalt: Neubert 2011), northern Germany (Michaelsen 1890; Graff 1950, 1954; Graefe 1993; Kula 1994; Irmler 1999; Unger 1999; Groth & Irmler 2012); southern Germany (Zuck 1951; Bauchhen�� 1982; Babel et al. 1992; Kula 1994; Ehrmann 1996; Beck et al. 2001; Sommer et al. 2002; Auerswald et al. 2009; PSMS in Bavaria: www.edaphobase.org), eastern Germany (Wilcke 1939; F��ller 1953; B��sener 1965: Kr��ck 1999; Kr��ck et al. 2001; Beylich & Graefe 2002; Muskolus 2008; PSMS in Brandenburg: www.edaphobase.org); western Germany (Baltzer 1956; Schulte et al. 1989; Keplin & Broll 1997; Emmerling 2001; Fr��nd et al. 2004)., Published as part of Lehmitz, Ricarda, R��mbke, J��rg, J��nsch, Stephan, Kr��ck, Stefanie, Beylich, Anneke & Graefe, Ulfert, 2014, Checklist of earthworms (Oligochaeta: Lumbricidae) from Germany, pp. 221-245 in Zootaxa 3866 (2) on page 223, DOI: 10.11646/zootaxa.3866.2.3, http://zenodo.org/record/229518, {"references":["Savigny, J. 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Summary The population dynamics of earthworms and potworms (Oligochaeta: Lumbricidae, Enchytraeidae) were recorded in three floodplain meadows differing in soil type in Northern Germany during 2 years of contrasting hydrological extremes: 2002 with high precipitation, extended winter inundations and summer floods in two of the sites, and 2003 with an extraordinarily dry summer. In all sites, earthworms dominated the biomass of soil fauna, the dominant species being Octolasion tyrtaeum in marsh soil, Octolasion cyaneum in peat soil, Allolobophora chlorotica in gley soil, and Lumbricus rubellus in all three sites. The enchytraeid communities composed of 7–10 species, differed widely between sites. While the summer flood of 2002 reduced annelid populations in peat soil to zero, in gley soil only enchytraeids were sharply reduced. The earthworms there probably were less affected because of shorter duration of the flood, the lower air temperatures and SOM contents and the diapause of the dominant A. chlorotica . During summer drought in 2003, earthworms and enchytraeids were absent from gley soil, while they had maximum abundances in peat soil. The marsh soil, structured by a microrelief of shallow ditches and ridges, had highest densities of annelids. As it was not subjected to summer flooding in 2002, populations of both earthworms and potworms were stable. The drought in 2003 led to a sharp decrease in earthworm abundance in July, while the enchytraeids reached maximum values. Dynamics and survival strategies of different annelid species are discussed. The water holding capacity as well as the organic matter content of the soil are important factors for annelids in sites subjected to such hydrological extremes. To promote terrestrial annelids, controlled flooding should be kept short, especially in winters following natural summer floods. If possible, recovery times for annelids should be guarenteed, that is 2–3 months without inundations for enchytraeids and about half a year for earthworms. A microrelief with drier refuge sites can be judged as favourable for earthworms. An artificially raised groundwater table will favour annelids during drought periods.

Summary Developments in ecotoxicology and soil restoration have led to an increase in earthworm research over the past 20 years. Rearing of earthworms under laboratory conditions has therefore become desirable, but perceived difficulties have beset this practice. A review of recent research on the development of earthworm culture techniques for temperate, soil dwelling (anecic and endogeic) species is presented. Critical abiotic and biotic factors including soil moisture, organic matter, temperature, pH, earthworm density and species composition are considered, and optimal values provided for four species ( Allolobophora chlorotica, Aporrectodea caliginosa, Aporrectodea longa and Lumbricus terrestris ). Published figures relating to growth to maturity, cocoon production and cocoon viability, for these frequently studied species are collated and advice offered for culture of other soil dwelling species.

Earthworms dominate the animal biomass in moist floodplain soils. They are known to survive long periods in aerated water, but little is known about earthworm population dynamics in floodplain systems with changing inundation frequencies. This study determined earthworm population dynamics in a floodplain system, in relation to frequency and duration of flooding events. From October 2000 to May 2003 earthworms were hand sorted in the 'Afferdensche en Deestsche Waarden', a floodplain on the south bank of the river Rhine, near Druten, The Netherlands. Earthworm numbers and biomasses per age class (adult, subadult, juvenile) were recorded. Numbers and biomasses tend to decrease during flooding. Lumbricus terrestris was found in high numbers (>10/m2) only at the end of a flooding period. Allolobophora chlorotica was hardly affected by flooding; their biomass remained stable during the year. Aporrectodea caliginosa showed fluctuating numbers and biomasses during the sampling period that did not correlate with flooding frequency. Numbers and biomasses of Lumbricus rubellus were strongly reduced at the end of each flooding event, but their population densities fully recovered until next flooding event. Earthworm populations in floodplains fluctuate in time, depending on the season and on the time, duration and frequency of flooding. Different earthworm species react differently towards these flooding dynamics. © 2004 Elsevier GmbH. All rights reserved.

The objective of this study was to determine the impact of earthworm bioturbation on the distribution and availability of zinc in the soil profile. Experiments were carried out with Allolobophora chlorotica and Aporrectodea caliginosa in 24 perspex columns (0 10 cm), filled with 20-23 cm non-polluted soil (OM 2%, clay 2.9%, pH 0.01 M CaCl2 6.4), that was covered by a 3-5 cm layer of aged zinc spiked soil (500 mg Zn/kg dry soil) and another 2 cm non-polluted soil on top. After 80 and 175 days, columns were sacrificed and each cm from the top down to a depth of 15 cm was sampled. Earthworm casts, placed on top of the soil, were collected. Each sample was analyzed for total and CaCl2-exchangeable zinc concentrations. Effects of earthworm bioturbation were most pronounced after 1,75 days. For A. chlorotica, total and CaCl2-exchangeable zinc concentrations in the polluted layers were lower with than without earthworms. Total zinc concentrations in the non-polluted layers were higher in columns with earthworms. Casts of A. chlorotica collected on the soil surface showed slightly higher total zinc concentrations than non-polluted soil. Casts were found throughout the whole column. For A. caliginosa there were no differences in total zinc concentration between columns with and without earthworms. CaCl2-exchangeable zinc concentrations in the polluted layers were lower for columns with earthworms. Casts were mainly placed on top of the soil and contained total zinc concentrations intermediate between those in non-polluted and polluted soil layers. This study shows that different endogeic earthworm species have different effects on zinc distribution and availability in soils. A. chlorotica transfers soil throughout the whole column, effectively mixing it, while A. caliginosa decreases metal availability and transfers polluted soil to the soil surface. (c) 2004 Elsevier Ltd. All rights reserved.

Earthworms Allolobophora chlorotica and Aporrectodea longa were inoculated into Calvert landfill site in spring 1992, in conjunction with the planting of two tree species Alnus glutinosa and Acer pseudoplatanus. Monitoring has taken place over a period of 11 years. Sampling in 2003 revealed that earthworm distribution no longer equated to the inoculation treatments; the worms had spread extensively. The presence of A. glutinosa had a significant effect (p

Ziel der beschriebenen Untersuchungen war die praktische Uberprufung eines Methodenentwurfs zur Erfassung der Bioakkumulation von Umweltchemikalien mitEisenia fetida, der Eignung vonE. fetida als Modellorganismus und des OECD-Kunstbodens als Standardsubstrat. Als Testorganismen wurden aus eigener Zucht der KompostwurmEisenia fetida und nach FeldentnahmeAllolobophora caliginosa, Allolobophora chlorotica, Allolobophora longa undLumbricus rubellus eingesetzt. Testboden waren der OECD-Kunstboden und ein Boden vom Versuchsfeld der BBA in Berlin-Dahlem. Beiden Boden wurden die Testsubstanzen Hexachlorbenzol und Pyren in einer Konzentration von 10 mg/kg Bodentrockengewicht zugemischt. Die Expositionszeit der Regenwurmer betrug 4 Wochen bei wochentlicher Bestimmung der Konzentrationen der Testsubstanzen in den Wurmern und Boden. Die furE. fetida sowie die Wildarten errechneten Anreicherungsfaktoren (AF) beliefen sich bei HCB in Abhangigkeit des verwendeten Bodens auf Werte zwischen 10 und 17 und bei Pyren auf Werte zwischen 0,9 und 1,7. Durch Umrechnung der Bodenkonzentrationen auf Gehalte in der Wasserphase und Ableitung der entsprechenden Anreicherungsfaktoren kann ein Vergleich mit BCF-Werten, die aus QSAR’S anderer Regenwurmarten und Fischen berechnet wurden, hergestellt werden. Die Ergebnisse lassen den Schluss zu, dass der Regenwurmtest mitE. fetida (OECD 1984, ISO 1998) fur die Untersuchung der Bioakkumulation geeignet ist.

Summary Earthworms are sensitive bioindicators of soil pollution. The aim of present investigations was to study the effects of heavy metals on earthworms and on their coelomocytes involved in the defence reactions. Adult individuals of Allolobophora chlorotica collected in Krakow (K) soil were kept in the laboratory either in the K soil, or were transferred to unpolluted soil from the rural area Sierbowice (S) or to the heavily polluted (Zn>Pb>Cd>Cu) soil from the industrial area, Bukowno (B). They were kept there at 22C for up to 8 weeks. Cocoons and juveniles appeared in S and K soil samples. The number and activity of the coelomocytes of worms maintained in S and K soils were unaffected despite some accumulation of heavy metals in the earthworm tissues. In contrast, in the B soil samples, bioaccumulation of metals was strongest, high mortality of adults was recorded, body mass was reduced, and reproduction completely inhibited. Coelomocytes retrieved from the B soil survivors exhibited significant impairment of pinocytosis and plastic adherence. Perhaps impairment of immune functions contributed to the poor survival under conditions of heavily polluted B soil samples.

Earthworms are sensitive bioindicators of soil pollution. The aim of present investigations was to study the effects of heavy metals on earthworms and on their coelomocytes involved in the defence reactions. Adult individuals of Allolobophora chlorotica collected in Krakow (K) soil were kept in the laboratory either in the K soil, or were transferred to unpolluted soil from the rural area Sierbowice (S) or to the heavily polluted (Zn>Pb>Cd>Cu) soil from the industrial area, Bukowno (B). They were kept there at 22 °C for up to 8 weeks. Cocoons and juveniles appeared in S and K soil samples. The number and activity of the coelomocytes of worms maintained in S and K soils were unaffected despite some accumulation of heavy metals in the earthworm tissues. In contrast, in the B soil samples, bioaccumulation of metals was strongest, high mortality of adults was recorded, body mass was reduced, and reproduction completely inhibited. Coelomocytes retrieved from the B soil survivors exhibited significant impairment of pinocytosis and plastic adherence. Perhaps impairment of immune functions contributed to the poor survival under conditions of heavily polluted B soil samples.