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Allolobophora chlorotica (Savigny) An introduced species, found in a few antropogenic locations. Endogeic. Distribution (Fig. 1e): Gomel Reg.: Gomel, 52°25’23.15”N, 31°01’0.27”E, park, 21.10.2020, 14 ex. Vitebsk Reg.: Vitebsk, 55°11’9.86”N, 30°12’2.04”E, park, 05.10.2019, 4 ex. Note: A. chlorotica individuals from Gomel deviated from the typical morphology by the absence of pigmentation., Published as part of Shekhovtsov, Sergei V., Derzhinsky, Yevgeniy A., Ermolov, Sergei A., Kim-Kashmenskaya, Maria N., Poluboyarova, Tatiana V., Kotsur, Vladimir M. & Golovanova, Elena V., 2023, Earthworms of Belarus (Annelida: Megadrili), pp. 38-61 in Zootaxa 5255 (1) on page 46, DOI: 10.11646/zootaxa.5255.1.9, http://zenodo.org/record/7745192

Earthworms enhance plant growth but the precise mechanism by which this occurs is not known. An understanding of the mechanism could potentially support changes in agricultural management reducing fertiliser usage and therefore costs and the carbon footprint of agriculture. We conducted a factorial experiment in which 5 strains of wheat were grown in the presence and absence of earthworms under regular watering and droughted conditions. The different wheat strains all responded in a similar fashion. Plant biomass was greater in the presence of earthworms and under regular watering. The presence of earthworms reduced the impact of drought on plant biomass and also slowed down the rate of drying of the droughted soils. Plant nutrient content (N, P, Si) showed no consistent pattern with treatments but plant total N, P and Si mirrored plant biomass and decreased in the order earthworm-present watered > earthworm-present droughted > earthworm-absent watered > earthworm-absent droughted. Nutrient availability in the soil, as assessed by chemical extractions showed no consistent pattern with treatments. Differential gene expression of plants was greater between watering treatments than between earthworm treatments. Genes that were differentially expressed between the earthworm treatments predominantly related to plant defences, abiotic stress and control of plant growth though a couple were linked to both nitrogen cycling and stress responses. The soil microbiome of the earthworm-present treatments was more associated with nutrient-rich environments, the promotion of plant growth and the suppression of plant pathogens whilst that of the earthworm-absent treatments included a variety of plant pathogens. Our data are consistent with enhanced plant growth being due to changes in the microbiome brought about by earthworm processing of the soil rather than changes in nutrient availability directly due to earthworm activity.

Allolobophora chlorotica L7 MATERIAL COLLECTED. — France • 32 adult specimens, 1 juvenile specimen; Haute-Corse, Cagnano; 42°52’33”N, 9°26’6”E; 150 m a.s.l.; 31.III.2021; T. Decaëns, D. Fernández Marchán leg.; Eco & Sols (BOLD Sample ID: DFM-0118, DFM-0119, DFM-0120, DFM-0121, DFM-0122) • Haute-Corse, Castellare-Di-Casinca; 42°27’39”N, 9°29’27”E; 50 m a.s.l.; 29.III.2021; T. Decaëns, D. Fernández Marchán leg.; Eco & Sols (BOLD Sample ID: DFM- 0093, DFM-0094, DFM-0095, DFM-0096, DFM-0097, DFM- 0099) • Haute-Corse, Ghisonaccia, Marais de Cattolica; 42°1’15”N, 9°28’12”E; 8 m a.s.l.; 2.IV.2021; T. Decaëns, D. Fernández Marchán leg.; MNHN (BOLD Sample ID: DFM-0430, DFM-0431, DFM- 0432, DFM-0433) • Haute-Corse, Oletta, Aliso riparian forest; 42°39’14”N, 9°17’45”E; 24 m a.s.l.; 29.III.2021; T. Decaëns, D. Fernández Marchán leg.; MNHN (BOLD Sample ID: DFM- 0519) • Haute-Corse, Palasca, les Jardins de l’Ostriconi; 42°39’10”N, 9°4’44”E; 6 m a.s.l.; 30.III.2021; T. Decaëns, D. Fernández Marchán leg.; MNHN (BOLD Sample ID: DFM-0561, DFM-0562, DFM- 0563, DFM-0564, DFM-0565) • Haute-Corse, Poggio-d’Oletta, Saint Florent; 42°40’19”N, 9°20’42”E; 49 m a.s.l.; 1.IV.2021; T. Decaëns, D. Fernández Marchán leg.; MNHN (BOLD Sample ID: DFM-0421, DFM-0422, DFM-0423) • Haute-Corse, SantoPietro-di-Tenda, Casta; 42°39’57”N, 9°12’46”E; 251 m a.s.l.; 30.III.2021; T. Decaëns, D. Fernández Marchán leg.; Eco & Sols (BOLD Sample ID: DFM-0291, DFM-0292, DFM-0293, DFM- 0294, DFM-0295, DFM-0296) • Haute-Corse, Sisco, Chapelle Saint Michel; 42°49’22”N, 9°25’58”E; 344 m a.s.l.; 31.III.2021; T. Decaëns, D. Fernández Marchán leg.; MNHN (BOLD Sample ID: DFM-0381, DFM-0382, DFM-0383). REMARKS Five species-level lineages were previously described within A. chlorotica (Dupont et al. 2011) with the geographically restricted L5 (Switzerland and Southeastern France) suggested to be divided into three mitochondrial lineages (L5, L6 and L7). Marchán et al. (2020) attributed the A. chlorotica individuals found in the Hyères Archipelago to L5 (without considering the further subdivision). Specimens collected in Corsica cluster together with this same lineage, except for one specimen more closely related to specimens from Provence and the Alps. Upon closer inspection, both groups appear to be separate cryptic lineages which would correspond to L5 and L7 (as defined by Dupont et al. 2011)., Published as part of Marchán, Daniel Fernández, Gérard, Sylvain, Hedde, Mickaël, Rougerie, Rodolphe & Decaëns, Thibaud, 2022, An updated checklist and a DNA barcode library for the earthworms (Crassiclitellata, Oligochaeta) of Corsica, France, pp. 439-461 in Zoosystema 44 (17) on page 446, DOI: 10.5252/zoosystema2022v44a17, http://zenodo.org/record/7145097, {"references":["DUPONT L., LAZREK F., PORCO D., KING R. A., ROUGERIE R., SYMONDSON W. O. C., LIVET A., RICHARD B., DECAENS T., BUTT K. R. & MATHIEU J. 2011. - New insight into the genetic structure of the Allolobophora chlorotica aggregate in Europe using microsatellite and mitochondrial data. Pedobiologia 54 (4): 217 - 224. https: // doi. org / 10.1016 / j. pedobi. 2011.03.004","MARCHAN D. F., FERNANDEZ R., DOMINGUEZ J., DIAZ COSIN D. J. & NOVO M. 2020. - Genome-informed integrative taxonomic description of three cryptic species in the earthworm genus Carpetania (Oligochaeta, Hormogastridae). Systematics and biodiversity 18 (3): 203 - 215. https: // doi-org. inee. bib. cnrs. fr / 10. 1080 / 14772000.2020.1730474"]}

Allolobophora chlorotica (Savigny, 1826) Enterion chloroticum Savigny 1826: 182. Allolobophora chlorotica Černosvitov 1935: 54; ��apkarev 1972: 76; ��apkarev 1977: 50; Mr��ić 1991: 211; Csuzdi 2012: 97-99. E. Valley meadow, anthropogenic biotopes (Stojanović 1996); forest soils, cultivated soils, meadows (Stojanović et al. 2008; Milutinović et al. 2010). D. Babino selo (Černosvitov 1935); Mostar (��apkarev 1972); Jekovac, Sarajevo, Visoko (��apkarev 1977); Lisina (��ibovi) (Mr��ić 1991). Z. Peregrine species native in the Palearctic., Published as part of Sekuli��, Jovana, Stojanovi��, Mirjana, Popovi��, Filip & Traki��, Tanja, 2022, Checklist of the earthworm fauna of Bosnia & Herzegovina (Oligochaeta: Lumbricidae), pp. 501-518 in Zootaxa 5093 (5) on page 503, DOI: 10.11646/zootaxa.5093.5.1, http://zenodo.org/record/5964425, {"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.","Cernosvitov, L. (1935) Monographie des tschechoslovakischen Lumbriciden. Archiv pro Prirodovedecky Vyzkum Cech, 19, 1 - 86.","Sapkarev, J. (1972) Beitrage zur Kenntnis der Lumbricidenfauna Jugoslawiens. Archives of Biological Sciences, Belgrade, 24 (1 - 2), 73 - 86.","Sapkarev, J. (1977) Contribution to the knowledge of earthworms (Oligochaeta: Lumbricidae) from Slovenia, Yugoslavia. Godisen zbornik PMF, 30, 47 - 56.","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.","Csuzdi, C. (2012) Earthworm species, a searchable database. Opuscula Zoologica Budapest, 43, 97. [http: // earthworm. uw. hu]","Stojanovic, M. (1996) Faunisticko-ekoloska studija Lumbricida (Oligochaeta) Srbije. PhD, Faculty of Science, University of Kragujevac, Serbia. [unknown pagination, in Serbian]","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","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"]}

We carried out mesocosm experiments using either the anecic earthworm Lumbricus terrestris or the endogeic earthworm Allolobophora chlorotica and loam, silt loam and sandy loam soils to investigate the differing impact of these earthworm of different ecotypes on aggregate formation (percentage water stable aggregates, %WSA) and soil water holding capacity (WHC), two soil properties that underpin many of the ecosystem services provided by soils. Earthworms significantly increased %WSA (by 16–56% and 19–63% relative to earthworm-free controls for L. terrestris and A. chlorotica, respectively). For L. terrestris, this increase was significantly greater in the upper 6.5 cm of the soil where their casts were more obviously present. Allobophora chlorotica treatments significantly increased WHC by 7–16%. L. terrestris only caused a significant increase in WHC (of 11%) in the upper 6.5 cm of the sandy loam soil. Linear regression indicated a consistent relationship between increases in %WSA and WHC for both earthworm species. However, for a given %WSA, WHC was higher for A. chlorotica than L. terrestris likely due to the known differences in their burrow structure. Overall, earthworms increased soil %WSA and WHC but the significant species/ecotype differences need to be considered in discussions of the beneficial impacts of earthworms to soil properties.

Earthworms are an integral part of soil ecosystems, especially for their role in soil functions such as organic matter (OM) decomposition and nutrient cycling. Earthworms and microorganisms are interdependent, and a considerable portion of the contribution earthworms make to influencing OM fate is through interactions with microorganisms. However, the importance of the earthworm-associated microbiome is not fully understood, because it is difficult to separate the direct influence of the earthworms from the indirect influence of their microbiome. Here, we evaluated an antibiotic-based procedure to suppress the microbiome of individuals of ecologically-contrasting earthworm species (Eisenia fetida, Lumbricus terrestris, Allolobophora chlorotica) as the first step towards soil studies aimed at understanding the importance of the earthworm microbiome for host health and function. Individual earthworms were exposed to antibiotics: cycloheximide (150 μg ml−1), ampicillin (100 μg ml−1), ciprofloxacin (50 μg ml−1), nalidixic acid (50 μg ml−1), and gentamicin (50 μg ml−1) either singly or in a cocktail via culture (96 h) in a semi-solid agar carrier. Compared to the non-antibiotic treated control, the cocktail (for all three species) and ciprofloxacin (for E. fetida and A. chlorotica) treatments significantly reduced (P < 0.05) culturable microbial abundance on nutrient agar and potato dextrose agar. The microbial counts were reduced to below detection (

The current study deals with earthworm biodiversity of Adana Province. Identification was made by examining the earthworm specimens collected in 6 different localities. At the end of the study, 5 species belonging to 3 genus were found: Aporrectodea caliginosa (Savigny, 1826), Aporrectodea trapezoides (Dugès, 1828), Allolobophora chlorotica (Savigny, 1826), Aporrectodea rosea (Savigny, 1826) and Octodrilus transpadanus (Rosa, 1884).

The functioning of soil ecosystems greatly depends on the interactions occurring between soil biota communities. It is well known that earthworms are an important soil component that substantially affects its function, including their meaningful impact on the development of different phytopathogenic soil fungi. Phytopathogenic fungi are responsible for crop disease and cause great economic damage. It has previously been established that earthworms’ coelomic fluid can suppress the growth of phytopathogenic fungi, but the exact molecular mechanism is unknown. The present study aimed at broadening the proof of this observed phenomenon by investigating the effects of the coelomic fluid extract of three different earthworm species (Eisenia andrei, Dendrobaena veneta and Allolobophora chlorotica) on the growth of six different phytopathogenic fungi species (Berkeleyomyces basicola, Fusarium culmorum, Globisporangium irregulare, Rhizoctonia solani, Macrophomina phaseolina, and Sclerotinia sclerotiorum). Coelomic fluid extract was obtained by electrostimulation or usage of extraction buffer (only in case of A. chlorotica) and prepared in three different concentrations by diluting the obtained coelomic fluid with physiological saline. The coelomic fluid extract of the three investigated earthworm species had an inhibitory effect on the growth of all six phytopathogenic fungi species. The greatest inhibitory effect was achieved with the E. andrei coelomic fluid extract reducing the growth of R. solani fungi. The findings of this research confirm the antifungal activity of coelomic fluid obtained from earthworm species belonging to different ecological categories and may be of potential use in crop protection against phytopathogenic fungi.

International audience; Earthworms are common organisms in soil toxicity-testing framework, and endogeic species are currently recommended due to their ecological role in agroecosystem. However, little is known on their pesticide metabolic capacities. We firstly compared the baseline activity of B-esterases and glutathione-5-transferase in Allolobophora chlorotica and Aporrectodea caliginosa. Secondly, vulnerability of these species to pesticide exposure was assessed by in vitro trials using the organophosphate (OP) chlorpyrifosethyl-oxon (CPOx) and ethyl-paraoxon (POx), and by short-term (7 days) in vivo metabolic responses in soil contaminated with pesticides. Among B-esterases, acetylcholinesterase (AChE) activity was abundant in the microsomal fraction (80% and 70% of total activity for A. caliginosa and A. chlorotica, respectively). Carboxylesterase (CbE) activities were measured using three substrates to examine species differences in isoenzyme and sensitivity to both in vitro and in vivo exposure. CbEs were mainly found in the cytosolic fraction (80% and 60% for A. caliginosa and A. chlorotica respectively). GST was exclusively found in the soluble fraction for both species. Both OPs inhibited B-esterases in a concentration-dependent manner. In vitro trials revealed a pesticide-specific response, being A. chlorotica AChE more sensitive to CPOx compared to POx. CbE activity was inhibited at the same extent in both species. The 7d exposure showed A. chlorotica less sensitive to both OPs, which contrasted with outcomes from in vitro experiments. This non-related functional between both approaches for assessing pesticide toxicity suggests that other mechanisms linked with in vivo OP bioactivation and excretion could have a significant role in the OP toxicity in endogeic earthworms.

Interactions of earthworm species with their environment and in particular the soil they inhabit is poorly understood. In this study, we aimed to use mid infrared absorption (MIR) spectroscopy of earthworm bodies in combination with discriminant analysis of principal components (DAPC), in order to investigate if their biogeochemical composition is determined by their species belonging or the soil they inhabit. Our conceptual approach included the comparison of MIR spectra of four earthworm species (Lumbricus terrestris, Allolobophora chlorotica, Aporrectodea caliginosa and Aporrectodea icterica) sampled from four different agricultural fields in France. Moreover, we tested the influence of the habitat on the earthworm MIR spectra by exchanging individuals and soils in an incubation experiment. MIR spectra varied according to species and sampling site. The MIR spectra of earthworm bodies also changed during the incubation experiment, and for L. terrestris, A. chlorotica and A. icterica depended on the properties of the soil ingested. This suggested that the food source in different habitats impacted the biogeochemical composition of earthworm bodies. DAPC allowed for the identification of earthworm species ranging from 64 to 100% of accuracy (mean 78%). We thus conclude that MIR spectroscopy may be a suitable approach to identify earthworm species, but that their signature may also allow to investigate their ecology as determined by the soil they inhabit.

Accurate species identification is crucial in ecological research on sentinel species such as earthworms. Here, we developed a reliable and cost-efficient method for the rapid identification of taxa within the Allolobophora chlorotica complex of cryptic earthworm species. We combined high resolution melting (HRM) analysis with DNA barcoding (Bar-HRM) in a three-step approach: (i) selection of a mini-barcode in the 16S mitochondrial gene, (ii) test of the method on a panel of 16 reference individuals and (iii) identification at the species level of 24 unknown specimens. Thus, we proved the efficacy of Bar-HRM to discriminate between cryptic sister (or sibling) earthworm species and we provided a standardized method that can be easily adapted to other taxa.

Southeast Iran is located in the arid area, near one of the hottest areas in the word. This area includes several climates, which led to different diverse in soil flora and fauna. In this study, earthworms of southeast Iran were collected. Eight species were identified from 67 samples based on morphological characters. Allolobophora chlorotica (Savigny, 1826) of Lumbricidae family and Amynthas gracilis (Kinberg, 1867) from Megascolecidae family are new records for Iran’s fauna. Five species from Lumbricidae family include Aporrectodea rosea, Aporrectodea trapezoides, Perelia kaznakovi, Eisenia andrei (maybe Eisenia fetida) and Perelia sp. and one species of Megascolecidae, Amynthas corticis, were also identified. Ap. trapezoides, Ap. rosea, and Am. corticis were common species in our study areas. E. andrei (maybe E. fetida) and Am. gracilis, were only found in Baft (Khabr) and Jiroft respectively.

Earthworms directly and indirectly affect soil physico-hydraulic properties. We studied interactions between winter wheat and the common European earthworm Allolobophora chlorotica, testing for impacts upon soil physico-hydraulic properties. Column experiments with three different soil textures and field-relevant earthworm densities were conducted. Saturated hydraulic conductivity (Ks) increased over time in earthworm treatments, with the increase occurring most rapidly in the plant plus earthworm treatment. After 16 weeks Ks had increased in the treatment comprising both plants and earthworms by factors of 12, 34 and 39 in the loam, silt-loam and sandy-loam soils respectively relative to controls. The presence of earthworms resulted in an 88.5% increase in the contribution of pores >3 mm diameter to water flow. In the majority of treatments, soil water-holding capacity and percentage water-stable aggregates of the 5 cm topsoil in both the plant and the earthworm treatments increased significantly compared to controls with the plant plus earthworm treatment showing the greatest increase. Plant growth was greater in the presence than absence of earthworms. Our study shows synergistic and additive effects on soil physical properties in the presence of both earthworms and plants compared to when only earthworms or plants are present. It shows that it is not just vertical-burrowing anecic earthworms that can have a significant effect on soil hydraulic properties but that field-relevant densities of endogeic earthworms such as A. chlorotica can also contribute to soil water flow regulation.

Anaerobic digestion (AD) is used to produce biogas and can offer a solution in waste management. Digestate, the AD by-product, can be applied to soil to improve fertility. However, the response of soil biological communities is not fully understood. There are mixed reports on its impact on earthworm survival. This study aimed to investigate digestate effects on earthworm mortality, and to elucidate potential mechanisms underlying it, if observed, after digestate application to soil. Juvenile and adult Allolobophora chlorotica were used as model organisms and added to microcosms prepared in a glasshouse trial. Five replicated treatments were: liquid Digestate; Osmotic-Stress (i.e. same salt concentration as digestate); Labile-C (i.e. same Biological Oxygen Demand as Digestate); Synthetic-Digestate a mixture of Osmotic-Stress and Labile-C (i.e. same salt concentration and BOD as digestate); and Water as the control. Treatments were applied at two different standardised rates equivalent to the digestate’s N content (i.e. 150 kg N ha−1 eq. or 300 kg N ha−1 eq.). The two development stages of A. chlorotica had different responses to treatments. Adult biomass was significantly greater in the Water control R150 treatment than in Digestate. Significantly lower juvenile biomass was observed in the Digestate R300 treatment than in the Labile-C and Water control treatments. The biomass of adults in the Labile-C R300 treatment was significantly greater than in the Digestate, Osmotic-Stress, Synthetic-Digestate and Water control treatments. Both life-stages exhibited a decline in biomass across all treatments, but the adults had higher mortality rates. The biomass of adults and juveniles declined, respectively, by 90% and 62% for Digestate applied at the lower rate, and by 96% and 90% at the higher rate. Whereas the abundance of adults and juveniles exhibited 80% and 24% drop at the lower rate, and a 90% and 84% drop at the higher rate. This study demonstrates that digestate can have negative impact on earthworm morbidity and mortality when applied to soil at 60% water filed pore space, with most of the total weight loss per pot due to reduced earthworm abundance. A likely hypothesis could be the osmotic stress induced by salts present in the digestate. However, there are other factors that interact with this effect, including possibly anaerobic impacts caused by high water content soils, as well as other mechanisms that have not been fully elucidated through this experimental design. Nevertheless, this work provides the basis for further ecotoxicology studies on the impact of digestate applied to soil. Further, while this works has shown that digestate can negatively impact A. chlorotica survival, whether the same is true for other earthworm species, ecotypes and life-cycle stages warrants further investigation. Considering the important role that worms play in soil health, field scale studies are also required to monitor the impacts of repeated digestate application on earthworm communities.

Increasing commercial application of silver nanoparticles (Ag NP) and subsequent presence in wastewater and sewage sludge has raised concerns regarding their effects in the aquatic and terrestrial environment. Several studies have employed standardised acute and chronic earthworm-based tests to establish the toxicological effects of Ag NP within soil. These studies have relied heavily on the use of epigiec earthworm species which may have limited ecological relevance in mineral soil. This study assessed the influence of Ag NP (uncoated 80 nm powder) and AgNO3 on survival, change in biomass and avoidance behaviour in a soil dwelling (endogiec) species, Allolobophora chlorotica. Earthworms were exposed for 14 days to soils spiked with Ag NP or AgNO3 at 0, 12.5, 25, 50 and 100 mg kg-1 either separately for survival and biomass measurement, or combined within a linear gradient to assess avoidance. Avoidance behaviour was shown to provide the most sensitive endpoint with an observable effect at an Ag NP / AgNO3 concentration of 12.5 mg kg-1 compared with 50 mg kg-1 for biomass change and 100 mg kg-1 for survival. Greater mortality was observed in AgNO3 (66.7%) compared with Ag NP-spiked soils (12.5%) at 100 mg kg-1, attributed to increased presence of silver ions. Although comparison of results with studies employing Eisenia fetida and Eisenia andrei suggest that the A. chlorotica response to Ag NP is more sensitive, further research employing both epigeic and endogeic earthworms under similar experimental conditions is required to confirm this observation.

Soil dwelling earthworms are now adopted more widely in ecotoxicology, so it is vital to establish if standardised test parameters remain applicable. The main aim of this study was to determine the influence of OECD artificial soil on selected soil-dwelling, endogeic earthworm species. In an initial experiment, biomass change in mature Allolobophora chlorotica was recorded in Standard OECD Artificial Soil (AS) and also in Kettering Loam (KL). In a second experiment, avoidance behaviour was recorded in a linear gradient with varying proportions of AS and KL (100% AS, 75% AS + 25% KL, 50% KS + 50% KL, 25% AS + 75% KL, 100% KL) with either A. chlorotica or Octolasion cyaneum. Results showed a significant decrease in A. chlorotica biomass in AS relative to KL, and in the linear gradient, both earthworm species preferentially occupied sections containing higher proportions of KL over AS. Soil texture and specifically % composition and particle size of sand are proposed as key factors that influenced observed results. This research suggests that more suitable substrates are required for ecotoxicology tests with soil dwelling earthworms.

Phosphorus (P) is an important nutrient for plant growth. However, P is often poorly available for uptake by roots because it strongly adsorbs to the soil mineral phase. Recent research shows that earthworms can temporally and locally increase P availability to plants. However, the pathways through which they do so are not fully understood, and it remains unclear to what extent this capacity varies among earthworm species. Here we study the variation among earthworm species with respect to readily-available P in casts as well as other physico-chemical cast properties, in a greenhouse pot experiment using a soil with a low P status. The earthworms belong to eight commonly occurring earthworm species in the Netherlands: two epigeic species (a mixture of the compost earthworms Dendrobaena veneta/Eisenia fetida; Lumbricus rubellus); four endogeic species (Allolobophora chlorotica, Aporrectodea caliginosa, Aporrectodea rosea, Octolasion lacteum); and two anecic species (Aporrectodea longa; Lumbricus terrestris). For all species, the pH in water extracts of earthworm cast (pH = 7.4–8.2) was significantly higher (p < 0.001) than for the control bulk soil (pH = 6.6) and differed significantly (p = 0.003) among earthworm species. Similarly, the dissolved organic carbon (DOC) concentration in the same water extracts was an order of magnitude higher in earthworm cast compared to the control bulk soil and varied among species (p < 0.001). The size of the total pool of reversibly adsorbed P in earthworm cast was greater than in the control bulk soil, but no significant differences were found among earthworm species. Differences among species were present for the readily-available P pools extracted from casts, including P-Olsen and water-extractable ortho-P. Water-extractable ortho-P concentrations were much higher in the casts of all species as compared to the control bulk soil (0.9–6.8 vs 0.06 mg l−1 or to 9.0–68 vs 0.6 mg kg−1). Highest ortho-P levels were measured in L. rubellus casts and the lowest in casts of A. chlorotica. A positive correlation between the concentrations of DOC and ortho-P was observed (R2 = 0.72, p < 0.001). The observed variation in all measured physico-chemical cast properties could not be explained by conventional ecological earthworm classifications. Our results show that the nature and magnitude of earthworm-induced increased P availability differs dramatically among earthworm species. This strongly suggests that, apart from its size, species composition of the earthworm community is key to optimizing P availability to plants.

While many studies have examined the effects of flooding on earthworm population distributions, few studies have investigated physiological and behavioural responses of earthworms to the low oxygen conditions caused by flooding. An earthworm's skin is its oxygen exchange organ, allowing earthworms to survive in flooded environments provided that the water contains sufficient dissolved oxygen. Individuals of three species of earthworm, the anecic Lumbricus terrestris (Linneaus, 1758), the green morph of the endogeic Allolobophora chlorotica (Savigny, 1826) and the epigeic Lumbricus castaneus (Savigny, 1826) were placed in reconstituted groundwater that was either kept aerated or kept in a sealed container so that dissolved oxygen was gradually consumed as the earthworm respired. Oxygen saturation of the water was measured over time in sacrificial triplicate replicates from each treatment at discrete time points; earthworm mortality and mass were recorded. Before treatments, oxygen levels in all treatment tubes were 9.53 (±0.64) mg O2 L−1. L. terrestris, a large species which emerges at night to forage at the soil surface died when oxygen levels reached 0.82 (±0.46) mg O2 L−1 after approximately 36 h. L. castaneus, a smaller species which lives on the soil surface, died when oxygen levels reached 3.60 (±2.01) mg O2 L−1 after approximately 168 h. A. chlorotica, which is similar in size to L. castaneus, lives in the upper 20 cm of soil and is known to aestivate during the summer, did not die, even when oxygen levels reached 1.49 (±0.40) mg O2 L−1 after 280 h. The results suggest that earthworm respiration is closely linked to both body size and to behavioural ecotype. These findings suggest that if flooding increases in frequency resulting in episodic reductions in soil oxygen levels, the species composition of earthworm communities may change, with an increased presence of endogeic earthworms which show a responsive plasticity to flooding events.

We performed a greenhouse experiment to test how the functional diversity of earthworms, the dominant group of soil macro-invertebrates in many terrestrial ecosystems, affects nitrogen cycling and plant growth. Three species were chosen to represent a range of functional traits: Lumbricus terrestris (large, mainly detritivorous, makes vertical permanent burrows open at the surface), Aporrectodea longa (medium-large, feeds on both detritus and soil, makes burrows more branched than L. terrestris), and Allolobophora chlorotica (small, geophagous, makes ephemeral burrows below the soil surface). Mesocosms with ryegrass (Lolium perenne) were inoculated with none to all three species (similar total biomass), using an experimental design (Simplex) suited to partition single species and diversity effects. Two contrasting N sources, urea or mammalian dung, were labelled with 15N so that the acquisition by plants and earthworms and recovery of applied 15N could be estimated. Over 3 months, plant production was higher with urea applications, but there were also species-specific earthworm effects: A. chlorotica and, to a lesser extent, A. longa increased shoot biomass, whereas L. terrestris increased root biomass. Earthworms did not affect soil N concentrations or leaching losses, whereas more N was leached under urea. A. chlorotica tended to increase dung-15N recovery in grass shoots, but in interaction with A. longa had the opposite effect, possibly through increased N immobilization in the microbial biomass. Earthworms assimilated negligible amounts of urea-15N but a substantial proportion (17% on average) of the dung-15N, with no clear-cut differences between species. Our findings show that earthworm species may have similar trophic response to N sources and yet different effects on plant N uptake and primary production, and that inter-specific earthworm interactions can result in non-additive diversity effects.

By means of a literature survey, earthworm species of significant relevance for soil functions in different biogeographical regions of Europe (Atlantic, Boreal, Mediterranean) were identified. These focal earthworm species, defined here according to the EFSA Guidance Document on the environmental risk assessment (ERA) of genetically modified plants, are typical for arable soils under crop rotations with maize and/or potatoes within the three regions represented by Ireland, Sweden and Spain, respectively. Focal earthworm species were selected following a matrix of four steps: Identification of functional groups, categorization of non-target species, ranking species on ecological criteria, and final selection of focal species. They are recommended as appropriate non-target organisms to assess environmental risks of genetically modified (GM) crops; in this case maize and potatoes. In total, 44 literature sources on earthworms in arable cropping systems including maize or potato from Ireland, Sweden and Spain were collected, which present information on species diversity, individual density and specific relevance for soil functions. By means of condensed literature data, those species were identified which (i) play an important functional role in respective soil systems, (ii) are well adapted to the biogeographical regions, (iii) are expected to occur in high abundances under cultivation of maize or potato and (iv) fulfill the requirements for an ERA test system based on life-history traits. First, primary and secondary decomposers were identified as functional groups being exposed to the GM crops. In a second step, anecic and endogeic species were categorized as potential species. In step three, eight anecic and endogeic earthworm species belonging to the family Lumbricidae were ranked as relevant species: Aporrectodea caliginosa, Aporrectodea rosea, Aporrectodea longa, Allolobophora chlorotica, Lumbricus terrestris, Lumbricus friendi, Octodrilus complanatus and Octolasion cyaneum. Five out of these eight species are relevant for each biogeographical region with an overlap in the species. Finally, the earthworm species Ap. caliginosa (endogeic, secondary decomposer) and L. terrestris (anecic, primary decomposer) were selected as focal species. In the Mediterranean region L. terrestris may be substituted by the more relevant anecic species L. friendi. The selected focal species are recommended to be included in a standardized laboratory ERA test system based on life-history traits. © 2016 Elsevier B.V.

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.

Allolobophora chlorotica chlorotica (Savigny, 1826) Enteri&ocy;n chl&ocy;r&ocy;ticum Savigny, 1826: 182. All&ocy;l&ocy;b&ocy;ph&ocy;ra chl&ocy;r&ocy;tica: ��apkarev 1980: 166; Stojanovi&cacute; 1996: 26; Stojanovi&cacute; & Karaman 2005a: 128; Milutinovi&cacute; et al. 2010: 629. All&ocy;l&ocy;b&ocy;ph&ocy;ra chl&ocy;r&ocy;tica chl&ocy;r&ocy;tica: Mr��i&cacute; 1991: 211; Stojanovi&cacute; et al. 2008: 60; Csuzdi 2012: 97; Stojanovi&cacute; & Milutinovi&cacute; 2014: 308. E. V&acy;ll&iecy;y m&iecy;&acy;d&ocy;w, &acy;nthr&ocy;p&ocy;g&iecy;nic bi&ocy;t&ocy;p&iecy;s (St&ocy;j&acy;n&ocy;vi&cacute; 1996); f&ocy;r&iecy;st s&ocy;ils, cultiv&acy;t&iecy;d s&ocy;ils, m&iecy;&acy;d&ocy;ws (St&ocy;j&acy;n&ocy;vi&cacute; et al. 2008; Milutin&ocy;vi&cacute; et al. 2010). D. P&acy;nn&ocy;ni&acy;n p&acy;rt���P&acy;li&cacute;, P&acy;n&ccaron;&iecy;v&ocy;, Zr&iecy;nj&acy;nin (��&acy;pk&acy;r&iecy;v 1978); B&iecy;l&acy; Crkv&acy; (��&acy;pk&acy;r&iecy;v 1978, 1980); Fru��k&acy; g&ocy;r&acy;, Pl&acy;ti&ccaron;&iecy;v&ocy;, K&acy;r&acy;vuk&ocy;v&ocy;, Tursk&iecy; liv&acy;d&iecy; (St&ocy;j&acy;n&ocy;vi&cacute; & Milutin&ocy;vi&cacute; 2014). B&acy;lk&acy;n p&acy;rt: B&iecy;&ocy;gr&acy;d (��&acy;pk&acy;r&iecy;v 1972); Gr&ocy;ck&acy;, K&acy;r&acy;burm&acy;, (��&acy;pk&acy;r&iecy;v 1980, 1989; St&ocy;j&acy;n&ocy;vi&cacute; 1996); Ml&acy;d&iecy;n&ocy;v&acy;c, K&ocy;r&acy;&cacute;ic&acy;, Kr&acy;guj&iecy;v&acy;c, L&acy;z&acy;r&iecy;v&acy;c, T&ocy;p&ocy;l&acy;, Ar&acy;n&dstrok;&iecy;l&ocy;v&acy;c (��&acy;pk&acy;r&iecy;v 1989); Knj&acy;&zcaron;&iecy;v&acy;c, N&ocy;vi P&acy;z&acy;r, Ni��, Svrljig (��&acy;pk&acy;r&iecy;v 1980; St&ocy;j&acy;n&ocy;vi&cacute; 1996); Kru��&iecy;v&acy;c, Pir&ocy;t, Klisur&iecy; n&iecy;&acy;r Vl&acy;sin&acy;, L&iecy;p&iecy;nic&acy;, P&iecy;tr&ocy;v&acy;&ccaron;k&acy; r&iecy;k&acy;, M&ocy;r&acy;v&acy;, H&ocy;m&ocy;lj&iecy;, Sj&iecy;nic&acy; (St&ocy;j&acy;n&ocy;vi&cacute; 1996); J&acy;str&iecy;b&acy;c (St&ocy;j&acy;n&ocy;vi&cacute; & K&acy;r&acy;m&acy;n 2005&acy;; Milutin&ocy;vi&cacute; et al. 2010). Z. P&iecy;r&iecy;grin&iecy; sp&iecy;ci&iecy;s n&acy;tiv&iecy; in th&iecy; P&acy;l&iecy;&acy;rctic., 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.

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.

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.

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.