Search

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

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"]}

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.

It is generally accepted that populations of most species experience some level of density dependence; however, this has rarely been shown for soil-dwelling invertebrate species, in particular for earthworms. Experimental data, mostly obtained at high laboratory densities, suggest intense intraspecific competition for resources and/or living space and density-dependent dynamics in earthworm populations, irrespective of their ecological group affiliation. In the present study performed in large field microcosms, I investigated whether density-dependent responses occur within the earthworm density gradients more realistic for the natural sites. Five lumbricid species from epigeic ( Dendrobaena octaedra , Lumbricus rubellus ), endogeic ( Allolobophora chlorotica , Aporrectodea caliginosa ) and anecic ( Lumbricus terrestris ) ecological groups were tested. Soil systems populated with earthworm monocultures in large (20-L) microcosms were exposed in a beech-oak forest for 4.5 months; each species was represented by two ( L. terrestris ) or three treatments forming gradients of increasing density. In two endogeic species, manifold and generally similar density-dependent responses (a retardation of growth, maturation and reproduction rates, but higher mortality rates with density increase) were revealed, likely explained by spatial competition rather than by direct food competition. In each of epigeic species density-dependent responses were less variable and more species-specific. In contrast to endogeics, direct food competition was presumably a more important cause of density-dependence. In L. terrestris no significant density-dependent responses in adult earthworms were revealed; however, they need to be further investigated in relation to the age and territorial behaviour of individual earthworms. Importantly, in any earthworm species the density variations in the reproducing generation had significant consequences for the advancing generation, affecting either the numbers or/and the size (individual weight) of the cocoons produced. It is suggested that an underestimation of density-dependent processes may cause inaccurate estimates of the activities of local lumbricid populations.

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.

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."]}

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.

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.

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

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.

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 (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. 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. Vol. 5. Paris, pp. 176 - 184.","Hoffmeister, W. (1843) Beitrage zur Kenntnis deutscher Landnanneliden. Archiv fur Naturgeschichte, 9 (1), 183 - 198.","Hoffmeister, W. (1845) Ubersicht aller bis jetzt bekannten Arten aus der Familie der Regenwurmer. Friedrich Vieweg & Sohn, Braunschweig, 43 pp.","Vejdovsky, F. (1884) System und Morphologie der Oligochaeten. Franz Rivnac, Prag, 166 pp.","Michaelsen, W. (1890) Die Lumbriciden Norddeutschlands. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, 7, 1 - 19.","Wilcke, D. E. (1949) Bestimmungstabelle fur einheimische Lumbriciden. Senckenbergiana, 30, 171 - 181.","Zuck, W. (1951) Untersuchungen uber das Vorkommen und die Biotope einheimischer Lumbriciden. Jahreshefte des Vereins fur vaterlandische Naturkunde in Wurttemberg (Stuttgart), 107, 95 - 132.","Graff, O. (1953 b) Die Regenwurmer Deutschlands. Schriften der Forschungsanstalt fur Landwirtschaft Braunschweig. Verlag M. & H. Schaper, Hannover, 81 pp.","Bouche, M. B. (1972) Lombriciens de France. - Ecologie et Systematique. Institut National de la Recherche Agronomique, Paris, 671 pp. http: // dx. doi. org / 10.2307 / 2402311","Easton, E. G. (1983) A guide to the valid names of Lumbricidae (Oligochaeta). In: Satchell, J. E. (Ed.), Earthworm ecology from Darwin to vermiculture. Chapman and Hall, London, pp. 475 - 785. http: // dx. doi. org / 10.1007 / 978 - 94 - 009 - 5965 - 1 _ 41","Sims, R. W. & Gerard, B. M. (1999) Earthworms. In: Barnes, R. S. K. & Crothers, J. H. (Eds.), Synopses of the British Fauna (New Series) No. 31 (Revised). London: E. J., 167 pp.","Csuzdi, C. & Zicsi, A. (2003) Earthworms of Hungary (Annelida: Oligochaeta, Lumbricidae). Pedozoologica Hungarica, 1, 1 - 272.","Blakemore, R. J. (2008) An updated list of valid, invalid and synonymous names of Criodriloidea (Criodrilidae) and Lumbricoidea (Annelida: Oligochaeta: Sparganophilidae, Ailoscolecidae, Hormogastridae, Lumbricidae, and Lutodrilidae), Robert J. Blakemore PhD, C / - Soil Ecology Group, Graduate School of Enviro and Info Sciences, Yokohama, National University, 79 - 7 Tokiwadai, Hodogaya, Yokohama 240 - 8501. Japan, 80 pp. Available from: www. annelida. net / earthworm / Lumbricidae. pdf (accessed 16 December 2013)","Michaelsen, W. (1900 a) Oligochaeta. Das Tierreich X. Friedlander & Sohn, Berlin, 575 pp.","Michaelsen, W. (1903) Die geographische Verbreitung der Oligochaeten. R. Friedlander & Sohn, Berlin, 186 pp. http: // dx. doi. org / 10.5962 / bhl. title. 11667","Jansch, S., Steffens, L., Hofer, H., Horak, F., Roß-Nickoll, M., Russell, D., Toschki, A. & Rombke, J. (2013) State of knowledge of earthworm communities in German soils as a basis for biological soil quality assessment. Soil Organisms, 85 (3), 215 - 233.","Graff, O. (1954) Die Regenwurmfauna im ostlichen Niedersachsen und in Schleswig-Holstein. Beitrage zur Naturkunde Niedersachsens, 7 (2), 48 - 56.","Eggert, U. J. (1982) Vorkommen und Verbreitung der Regenwurmer (Lumbricidae) des Naturparks \" Hoher Vogelsberg \". Beitrage zur Naturkunde in Osthessen, 18, 61 - 103.","Westernacher-Dotzler, E. (1988) Abundanz von Regenwurmern (Lumbricidae, Oligochaeta) unter verschiedenen Kulturpflanzen. PhD-thesis, 149 pp.","Neubert, E. (2011) Lumbriciden 2011 - Regenwurmer auf Boden-Dauerbeobachtungsflachen in Sachsen-Anhalt. Bericht des Landesamtes fur Umweltschutz Sachsen-Anhalt, 30 pp.","Graff, O. (1950) Die Regenwurmer der Umgebung von Braunschweig und ihre Bedeutung fur die Landwirtschaft. PhD-thesis, 87 pp.","Graefe, U. (1993) Veranderungen der Zersetzergesellschaften im Immissionsbereich eines Zementwerkes. Mitteilungen der Deutschen Bodenkundlichen Gesellschaft, 72, 531 - 534.","Kula, H. (1994) Auswirkungen von Pflanzenschutzmitteln auf Regenwurmer (Oligochaeta: Lumbricidae) - Zur Problematik der Bewertung letaler und subletaler Effekte in Labor- und Feldversuchen. PhD-thesis, 151 pp.","Irmler, U. (1999) Die standortlichen Bedingungen der Regenwurmer (Lumbricidae) in Schleswig-Holstein. Faunistisch- Okologische Mitteilungen, 7, 509 - 518.","Unger, K. (1999) Untersuchungen zur Lumbricidenfauna der Stadt Greifswald. Zeitschrift fur Okologie und Naturschutz, 8, 125 - 134.","Groth, J. & Irmler, U. (2012) Spatial distribution of earthworms (Oligochaeta: Lumbricidae) and the relationship to environmental parameters in northern German wet grassland. Faunistisch-Okologische Mitteilungen, 9, 294 - 303.","Bauchhenß, J. (1982) Artenspektrum, Biomasse, Diversitat und Umsatzleistung von Lumbriciden (Regenwurmer) auf unterschiedlich bewirtschafteten Grunlandflachen verschiedener Standorte Bayerns. Bayrisches Landwirtschaftliches Jahrbuch, 59, 119 - 125.","Babel, U., Ehrmann, O. & Krebs, M. (1992) Relationships between earthworms and some plant species in a meadow. Soil Biology and Biochemistry, 24, 1477 - 1481. http: // dx. doi. org / 10.1016 / 0038 - 0717 (92) 90136 - l","Ehrmann, O. (1996) Regenwurmer in einigen sudwestdeutschen Agrarlandschaften: Vorkommen, Entwicklung bei Nutzungsanderungen und Auswirkungen auf das Bodengefuge. Hohenheimer Bodenkundliche Hefte, 35, 1 - 135.","Beck, L., Rombke, J., Paulus, R., Ruf, A., Scheurig, M., Spelda, J. & Woas, S. (2001) Bodenfauna und Umwelt - Bodenokologische Inventur und Beurteilung von ausgewahlten Standorten in Baden-Wurttemberg. PAOE-Report (Baden- Wurttemberg), 131 pp.","Sommer, M., Ehrmann, O., Friedel, J. K., Martin, K., Vollmer, T. & Turian, G. (2002) Boden als Lebensraum fur Organismen - Regenwurmer, Gehauselandschnecken und Bodenmikoorganismen in Waldern Baden-Wurttembergs. Hohenheimer Bodenkundliche Hefte, 63, 1 - 163.","Auerswald, K., Weigand, S., Kainz, M. & Philipp, C. (2009) Influence of soil properties on the population and activity of geophagous earthworms after five years of bare fallow. Biology and Fertility of Soils, 23, 382 - 387.","Wilcke, D. E. (1939) Der gegenwartige Stand unserer Kenntnis der markischen Lumbricidenfauna. Markische Tierwelt, 4, 34 - 50.","Fuller, H. (1953) Tiergeographisch-okologische Untersuchungen uber die Lumbriciden des mittleren Saaletales. Wissenschaftl. Zeitschrift der Friedrich-Schiller-Universitat Jena, 2, 51 - 60.","Bosener, R. (1965) Untersuchungen uber das Vorkommen und die forstliche Bedeutung von Lumbriciden (Oligochaeta) in verschiedenen Waldbestanden des Osterzgebirges. Wissenschaftliche Zeitschrift der Technischen Universitat Dresden, 14 (3), 741 - 746.","Kruck, S. (1999) Einfluß der Nutzung auf Bodenfruchtbarkeitsparameter, Humushaushalt und Regenwurmaktivitat, und deren Beziehung zur Ertragsfahigkeit sandiger Boden in Brandenburg. Berichte aus der Agrarwissenschaft, Shaker Verlag, Aachen, 154 pp.","Kruck, S., Nitzsche, O. & Schmidt, W. (2001) Regenwurmer vermindern Erosionsgefahr. Landwirtschaft ohne Pflug, 1, 18 - 21.","Beylich, A. & Graefe, U. (2002) Annelid coenoses of wetlands representing different decomposer communities. In: Broll, G., Merbach, W. & Pfeiffer, E. - M. (Eds.), Wetlands in Central Europe. Soil organisms, soil ecological processes and trace gas emissions. Springer, Berlin, pp. 1 - 10. http: // dx. doi. org / 10.1007 / 978 - 3 - 662 - 05054 - 5 _ 1","Muskolus, A. (2008) Anthropogenic Plant Nutrients as Fertiliser. PhD-thesis Humboldt-Universitat zu Berlin, 138 pp.","Baltzer, R. (1956) Die Regenwurmer Westfalens. Zoologisches Jahrbuch, 84, 355 - 413.","Schulte, W., Frund, H. - C., Sontgen, M., Graefe, U., Ruszkowski, B., Voggenreiter, V. & Weritz, N. (1989) Zur Biologie stadtischer Boden - Beispielraum: Bonn-Bad Godesberg. KILDA-Verlag, 184 pp.","Keplin, B. & Broll, G. (1997) Earthworms and dehydrogenase activity of urban biotopes. Soil Biology and Biochemistry, 29 (3 / 4), 533 - 536. http: // dx. doi. org / 10.1016 / s 0038 - 0717 (96) 00177 - 0","Emmerling, C. (2001) Response of earthworm communities to different types of soil tillage. Applied Soil Ecology, 17, 91 - 96. http: // dx. doi. org / 10.1016 / s 0929 - 1393 (00) 00132 - 3","Frund, H. - C., Egbert, E. & Dumbeck, G. (2004) Spatial distribution of earthworms (Lumbricidae) in recultivated soils of the Rhenish lignite-mining area. Journal of Plant Nutrition and Soil Science, 167, 494 - 502. http: // dx. doi. org / 10.1002 / jpln. 200421372"]}

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.

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.

Nightcrawler (Lumbricus terrestris L. [Oligochaeta: Lumbricidae]) is an anecic earthworm typically lacking from Persian walnut (Juglans regia L.) orchards in northern California, yet this earthworm may be established by inoculative release. Our purpose of was to determine the effect of L. terrestris inoculation on green manure incorporation, in this case woolypod vetch (Vicia villosa Roth ssp. varia [Host] Cabriere), and on the decomposer community. An experiment using controlled-environment microcosms showed that disappearance of surface-applied vetch residue was increased by the addition of L. terrestris compared to a resident earthworm complex consisting of the endogeic Lumbricidae Aporrectodea sp. prob. trapezoides (Duges) and Allolobophora chlorotica (Savigny) as well as the epigeic Eisenia foetida (Savigny). An experiment conducted in an organic Persian walnut orchard during the spring of 1996 also showed that disappearance of surface-applied vetch residue was increased in plots that had been inoculated with L. terrestris. However, an additional study conducted in the same orchard during the summer of 1998 showed similar rates of vetch biomass disappearance in plots with or without inoculated L. terrestris. Colonization by L. terrestris did not lead to significant differences in densities or live biomass for resident earthworms in this orchard. However, night observations showed significantly greater densities of isopod Crustacea (Isopoda: Oniscidae) in plots without L. terrestris, suggesting possible interference between L. terrestris and the isopods.

Burrow systems of earthworms contained in artificial cores were analysed through X-ray computed tomography and 3D skeleton reconstructions. Gas diffusion experiments were carried out on these cores to characterize soil transfer properties associated with the different burrow systems. Three types of cores were studied: cores in which Aporrectodea nocturna, an anecic earthworm was introduced (treatment 1), cores in which Allolobophora chlorotica, an endogeic earthworm was introduced (treatment 2) and cores that contained both species (treatment 3). Comparisons of the characteristics of the burrow systems of treatment 1 and 2 show important differences: the burrow system of A. nocturna comprises fewer burrows, which are longer, less branched, more vertical and have a lower sinuosity. The burrow system of A. chlorotica is characterized by lower continuity, which however did not result in a lower soil diffusivity. To study the burrow systems made by the two species in the same core (treatment 3), a separation that was based on differences in pore diameter between the two species and that takes into account the burrow orientation was designed. This separation was proven to be efficient since it resulted in low percentages of errors (around 10%) when applied to the burrow systems of treatments 1 and 2. Comparison of the burrow systems from treatments 1 and 3 demonstrated that the burrow system of A. nocturna was influenced by the presence of A. chlorotica: in treatment 3, A. nocturna made burrows that were smaller, more vertical and less branched. However, these interactions have to be confirmed under natural conditions. © 2001 Elsevier Science B.V. All rights reserved.

1. Earthworms (Annelida: Oligochaeta) were sampled on four occasions (spring, summer, autumn and winter) at 14 sites along two transects from a primary lead/ zinc/cadmium smelting works at Avonmouth, UK. 2. Total abundance and biomass of earthworms decreased with proximity to the smelter. No worms were collected from the two sites closest to the factory (

The natural abundances of the stable isotopes of nitrogen (δ 15 N) and carbon (δ 13 C) were measured in plant shoots and in seven earthworm (Lumbricidae) species from a wheat and a wheat-clover cropping system. Variations in earthworm δ 13 C were generally small in these systems containing only C 3 plants. Plant shoot δ 15 N ranged from −2.2‰ to −0.7‰ in white clover and from +0.9‰ to +6.6‰ in winter wheat. Intraspecific variability in earthworm δ 15 N was small. Earthworm δ 15 N was significantly related to the cropping system and to the ecological grouping of the earthworms. Mean δ 15 N values of all earthworm groups were significantly lower (by 3.5‰ to 4.4‰) in the wheat-clover field than in the wheat field. Within each of the two cropping systems, earthworm nitrogen isotope ratios differed significantly between ecological earthworm groups, with δ 15 N values decreasing in the order Allolobophora chlorotica and Aporrectodea caliginosa > Aporrectodea longa > Lumbricus spp. These results demonstrate for the first time that the δ 15 N signature of legumes can influence those of soil invertebrates. Because isotope measurements reflect assimilated tissue nitrogen, they offer novel insights into the feeding ecology and trophic positions of earthworms.

Allolobophora chlorotica (Savigny, 1826) Enterion chloroticum Savigny, 1826: 182. Allolobophora chlorotica: Zicsi & Michalis 1981: 248; 1993: 302. Michalis 1982: 349. Csuzdi & Zicsi 2003: 50 (for complete synonymy). Material examined. HNHM / 15721 four adult ex., No. 17, Trikala County, Athamon Mts., Athamania, small river and mixed rocky forest S of the village, 935m, N 39 ° 30.944 ’ E 21 ° 14.997 ’, leg. J. Kontschán, D. Murányi, T. Szederjesi, Zs. Ujvári, 0 9.0 5.2011.

Allolobophora chlorotica (Savigny, 1826) Enterion chloroticum Savigny, 1826: 182. Allolobophora chlorotica: Zicsi & Michalis 1981: 248; 1993: 302. Michalis 1982: 349. Csuzdi & Zicsi 2003: 50 (for complete synonymy). Material examined. HNHM / 15721 four adult ex., No. 17, Trikala County, Athamon Mts., Athamania, small river and mixed rocky forest S of the village, 935m, N 39 �� 30.944 ��� E 21 �� 14.997 ���, leg. J. Kontsch��n, D. Mur��nyi, T. Szederjesi, Zs. Ujv��ri, 0 9.0 5.2011., Published as part of Szederjesi, T��mea & Csuzdi, Csaba, 2012, New and little known earthworm species from Greece (Oligochaeta: Lumbricidae, Acanthodrilidae), pp. 25-42 in Zootaxa 3304 on page 29, DOI: 10.5281/zenodo.293430, {"references":["Savigny, J. C. (1826) In 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.","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. 271."]}

Allolobophora chlorotica chlorotica (Savigny, 1826) Enterion chloroticum Savigny, 1826: 182. Allolobophora (Allolobophora) chlorotica: Rosa 1893: 8. Allolobophora chlorotica: Omodeo 1956 a: 333; Mihailova 1966: 190; Pavli&cacute;ek et al. 2003: 456. Allolobophora chlorotica chlorotica: Csuzdi & Zicsi 2003: 50. Ecological group: endogeic (Butt & Lowe 2011). Habitat: forest soils, cultivated soils, meadows (Milutinovi&cdot; et al. 2010). Zoogeographical distribution type: peregrine (Csuzdi et al. 2011). Distribution in Bulgaria: rare, but found all over the country in wet, highly organic sites (Mihailova 1966)., Published as part of Valchovski, Hristo Ivailov, 2012, Checklist of earthworms (Oligochaeta: Lumbricidae) from Bulgaria ��� a review, pp. 86-102 in Zootaxa 3458 on page 87, DOI: 10.5281/zenodo.211011, {"references":["Savigny, J. C. (1826) In G. Cuvier (Ed.): 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.","Rosa, D. (1893) Viaggio del Dr. E. Festa in Palestina, nel Libano e regioni vicin .. II. Lumbricidi. Bolletino dei Musei di Zoologia ed Anatomia Comparata della Reale Universita di Torino, 8 (160), 1 - 14.","Omodeo, P. (1956 a) Oligocheti dell'Indochina e del Mediterraneo Orientale. Memorie del Museo Civico di Storia Naturale di Verona, 5, 321 - 336.","Mihailova, P. (1966) Dzdovni cervi Lumbricidae (Oligohaeta) v Trakija. Fauna na Trakja, Bulgarian Academy of Science, Sofia, 3, 181 - 200.","Pavlicek, T., Csuzdi, C. S. & Nevo, E. (2003) Species richness and zoogeographic affinities of earthworms in the Levant. Pedobiologia, 47, 452 - 457.","Butt, K. R. & Lowe, C. N. (2011) Controlled cultivation of endogeic and anecic earthworms. In: A. Karaca (Ed.), Biology of Earthworms, Springer, Berlin, pp. 107 - 121.","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). Second Balkan conference on biology Plovdiv 50 years university of Plovdiv, pp. 628 - 635."]}

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

Earthworm cocoons are mostly found in the uppermost soil layers and are therefore often exposed to low temperatures during winter. In the present study, cocoons of five taxa of earthworms were investigated for their tolerance to freezing, melting points of cocoon fluids and dehydration of cocoons when exposed to a frozen environment. Embryos of the taxa investigated were freeze intolerant. The melting points of fully hydrated cocoon fluids were high (above −0.3°C) and thermal hysteresis factors were absent. Exposure to a frozen environment caused the cocoons to dehydrate drastically and dehydrated cocoons showed significantly lower super-cooling points than fully hydrated cocoons, reducing the risk of freezing for dehydrated cocoons. It is proposed therefore that the cold-hardiness strategy of the earthworm cocoons is based on dehydration upon exposure to subzero temperatures in the frozen environment. Cocoons of three surface-dwelling taxa, Dendrobaena octaedra, Dendrodrilus rubidus tenuis and Dendrodrilus rubidus norvegicus had lower supercooling points and survived frost exposure better than cocoons of two deeper-dwelling taxa, Aporrectodea caliginosa and Allolobophora chlorotica. One of the investigated taxa, D. r. norvegicus, was collected from a cold alpine habitat. However, it was not more cold hardy than the closely related D. r. tenuis collected from a lowland temperate habitat. D. octaedra was the most cold hardy taxon, its cocoons being able to withstand −8°C for 3 months and −13.5°C for 2 weeks in frozen soil.

Concentrations of heavy metals (Pb, Cd, Zn and Cu) were determined in five species of earthworms, representing different ecophysiological groups, sampled from soils associated with an abandoned Pb and Zn mine site. Surface soil-dwelling species (Aporrectodea caliginosa and Allolobophora chlorotica) accumulated significantly greater concentrations of Cd than deep-burrowing (Lumbricus terrestris and Aporrectodea longa) or litter-dwelling (L. rubellus) species. Aporrectodea caliginosa generally accumulated greater concentrations of Pb than other species. The greatest concentrations of Zn were generally found in the deep-burrowing species, whilst lowest concentrations were found in Allolobophora chlorotica. No consistent inter-species pattern was evident for Cu. The composition (metal and organic carbon) of the ingesta of earthworms was different to that of the soil. Concentrations of Cd in the ingesta of Aporrectodea caliginosa were significantly greater than in the ingesta of Aporrectodea longa and L. terrestris; concentrations ofPb in the ingesta of Aporrectodea caliginosa and Aporrectodea longa were greater than for L. terrestris. The metal concentrations in the faeces were similar for all species. A significant correlation exists between ingesta and tissue Cd concentrations, but was not evident for Pb or Zn. This suggests that ingested total Cd concentration is a primary factor in determining tissue Cd concentrations of earthworms. For the other metals other interactive factors appear to markedly influence accumulated tissue metal concentrations.

During the past 25 yr, research by the authors at Rothamsted Experimental Station investigated many aspects of the utilization of earthworms in land improvement and environmental management. Results of some of these investigations are summarized in this paper with the aim of illustrating the general principles of how earthworm populations can be manipulated and managed for environmental improvement. The use of earthworms in land improvement and reclamation: we investigated the effects of inoculating earthworms of the species Lumbricus terrestris L., Aporrectodea longa (Ude), Aporrectodea caliginosa (Sav.) and Allolobophora chlorotica (Sav.) into intact soil profiles in the laboratory, plots on direct-drilled, arable land in the field and newly-capped waste disposal sites that had few or no earthworms. In all these studies the earthworms increased significantly in number and rate of growth and yield of plants growing on the inoculated sites. Earthworms for inoculation were obtained by field collection after watering soil with dilute formaldehyde solution. The use of earthworms in organic waste management: the life cycles and productivity of Eisenia fetida (Sav.), Eudrilus eugeniae (Kinberg), Perionyx excavatus (Michaelsen) and Dendrobaena veneta (Rosa), and their potential in processing animal and plant wastes, from sewage, agricultural, domestic, urban and industrial sources are summarized. The preprocessing of wastes, their population ecology, optimum stocking rates, the mechanization of processing and utilization of the product are discussed. Results of experiments on the effects of temperatures of 10, 15, 20 and 25°C and a range of soil moisture contents of 70, 75, 80, 85 and 90% on the growth, cocoon production and cocoon hatching of the four species are summarized. The use of earthworms in assessment of the environmental effects of chemicals: earthworms can be used as key indicators to predict the effects of chemicals on other soil invertebrates. Methods of testing chemicals against earthworms in field and laboratory are reviewed. Two standardized laboratory test methods, one exposing earthworms to chemicals on filter paper and one to chemicals in artificial soils are described, and the median lethal concentration (LC 50 ) for chloracetamide, pentachlorophenol, chlordane, carbaryl, potassium bromide, copper sulfate and trichloracetic acid calculated, based on assays done in 34 laboratories. The relevancy of the two tests in environmental toxicity testing is reviewed.

Diplôme : Master Recherche

Oil spills are one of the most common types of soil pollution. Bioremediation has become an attractive alternative to physicochemical methods of remediation, where feasible. Earthworms have been shown to stimulate the degradation of petroleum hydrocarbons in soil, and it was hypothesized that the role of earthworms in remediation lies in the enhancement of an oil degrading microbial community. The aim of this study was to characterize microbial activity and community dynamics in oil-contaminated soil incubated with or without earthworms. Three earthworm species (Eisenia fetida, Allolobophora chlorotica and Lumbricus terrestris) were incubated in crude oil polluted soil (ca. 10,000 mg/kg total petroleum hydrocarbons (TPH)) and a reference soil for 28 d. Control treatments with manual mixing and/or cattle dung amendment were also included. In the oil-contaminated soil, respiration and concentration of microbial biomass was significantly enhanced by earthworm amendment, and TPH concentrations decreased significantly. These effects were less evident in treatments with A. chlorotica, possibly due to a difference in behavior, since individuals of this endogeic species were found in a state of inactivity (aestivation). Microbial community dynamics were described by phospholipid fatty acid (PLFA) analyses. After 28 d, similar shifts in the soil PLFA composition were observed in the oil-contaminated soil irrespective of worm species. Fungal:bacterial ratios were increased in the presence of worms, but also by addition of dung as a food source, indicating a non-specific effect of metabolizable substrates. In contrast, the fatty acids 17:1ω8 (=Δ9-heptadecenoic acid) and 20:4ω6c (arachidonic acid) were specifically stimulated by the presence of earthworms in the oil-contaminated soil. The results showed that earthworms can contribute positively to bioremediation of oil-contaminated soil, but that the effect may be species-dependent.

The aim of this work was to exemplify the specific contribution of both two- and three-dimensional (3D) X-ray computed tomography to characterise earthworm burrow systems. To achieve this purpose we used 3D mathematical morphology operators to characterise burrow systems resulting from the activity of an anecic ( Aporrectodea nocturna), and an endogeic species ( Allolobophora chlorotica), when both species were introduced either separately or together into artificial soil cores. Images of these soil cores were obtained using a medical X-ray tomography scanner. Three-dimensional reconstructions of burrow systems were obtained using a specifically developed segmentation algorithm. To study the differences between burrow systems, a set of classical tools of mathematical morphology (granulometries) were used. So-called granulometries based on different structuring elements clearly separated the different burrow systems. They enabled us to show that burrows made by the anecic species were fatter, longer, more vertical, more continuous but less sinuous than burrows of the endogeic species. The granulometry transform of the soil matrix showed that burrows made by A. nocturna were more evenly distributed than those of A. chlorotica. Although a good discrimination was possible when only one species was introduced into the soil cores, it was not possible to separate burrows of the two species from each other in cases where species were introduced into the same soil core. This limitation, partly due to the insufficient spatial resolution of the medical scanner, precluded the use of the morphological operators to study putative interactions between the two species

An experimental study of the consequences of intensive cattle trampling was carried out pure white clover and on a perennial rye-grass-white clover association. Direct and indirect effects of trampling on earthworm and plant communities and on soil structure were found. Trampling led to destruction of a large portion of the aerial system, stolons and roots, with removal of some of the vegetation cover (soil at least 50% bare). From a general view, trampling in experimental plots induced a large decrease (from 70 to 86%) of earthworm density and biomass. The functional structure was modified for the benefit of large size species. Small size species were more sensitive: some species, such as Lumbricus castaneus and Allolobophora chlorotica chlorotica typica disappeared. An increase of the proportion of adults could be related to larger sensitiveness of juvenile individuals in the same population and explained a significant increase of the individual mean weight. Different responses existed in function of the grassland type related to depth.

SUMMARYDilute formalin was used to sample earthworm populations after the second (1969) and sixth (1973) harvests from a continuous barley experiment in which the cultivation treatments include deep (30 cm), normal ploughing (20 cm) and tined (three passes to 20 cm) cultivations and no tillage (direct drilling), at four nitrogen rates: 0, 50, 100 and 150 kgN/ha. Eight species of Lumbricidae were found, with Allolobophora chlorotica and A. longa co-dominant in both years, joined by A. caliginosa in 1973.In 1969, Lumbricus spp., particularly large Lumbricus terreslris, tended to be more numerous in the untilled plots.In 1973, the total populations had increased by 170%, with 50–100% more earthworms in untilled than ploughed plots, one third more with 50 and 100 kg N/ha fertilizer than with no nitrogen, and one-third more in the slightly heavier sandy clay loam soil than the lighter loam.Most earthworm species reacted in the same way, and the reasons for any unusual reactions are discussed.

Fifteen species of earthworm were collected from a reclaimed fen peat soil used for rough hay in County Offaly, Ireland. The most common species were Aporrectodea tuberculata, Allolobophora chlorotica, Eisenia rosea and Octolasion tyrtaeum. A. tuberculata, A. chlorotica and E. rosea formed a significant association. The total numbers of earthworms collected by hand-sorting reached a maximum of 197.1 m−2. Compared with hand-sorting, formalin-expulsion was an inefficient sampling method for most species. Soil moisture varied within the study area. Total densities and biomasses of the earthworms were greater where the soil was drier. A. tuberculata, A. turgida, Satchellius mammale and the pink form of A. chlorotica were more common in the dry soils whilst O. tyrtaeum, Lumbricus rubellus and the green form of A. chlorotica were more common in the wet soils.

Two types of Allobophora chlorotica exist in one of which porphyrins and a biliverdin-like substance accumulate during the individuals life-time, rendering it green. Populations of the green form o...

Surface cast production was investigated for several species of earthworms, including some not normally considered to produce surface casts. In single-species culture, the amount of cast material deposited on the surface by different species varied, with Allolobophora chlorotica < Lumbricus rubellus < Aporrectodea caliginosa < Aporrectodea longa. In field communities, results indicated that a single species dominated surface casting activity. The order of species dominance was consistent with the above findings from single-species cultures, if Lumbricus terrestris was considered to be equivalent to Ap. longa. Surface casting is an essential function within earthworm communities which maintains their living space. However, it involves an energy cost and carries a risk of predation. Therefore, there are disadvantages associated with this activity. All of the species studied produced surface casts unless other, usually larger, species were present.

Glutathione S-transferase activity (EC 2.5.1.18) was demonstrated in six species of earthworms of the family Lumbricidae: Eisenia foetida, Lumbricus terrestris, Lumbricus rebellus, Allolobophora longa, Allolobophora caliginosa and Allolobophora chlorotica. Considerable activity was obtained with 1-chlorl-2,4-dinitrobenzene and low activity with 3,4-dichloro-1-nitrobenzene, but no enzymic reaction was detectable with sulphobromophthalein 1,2-epoxy-3-(p-nitrophenoxy)propane of trans-4-phenylbut-3-en-2-one as substrates. Enzyme prepartations from L. rubellus and A. longa were the most active, whereas A. chlorotica gave the lowest activity. The ratio of the activities obtained with 1-chloro-2,4-dinitrobenzene and 3,4-cichloro-1-nitrobenzene was very different in the various species, but no phylogenetic pattern was evident. Isoelectric focusing gave rise to various activity peaks as measured with 1-chloro-2,4-dinitrobenzene as a substrate, and the activity profiles of the species examined appeared to follow a taxonomic pattern. The activity of Allolobophora had the highest peak in the alkaline region, whereas that of Lumbricus had the highest peak in the acid region. Eisenia showed a very complex activity profile, with the highest peak ne pH 7. As determined by an enzymic assay, all the species contained glutathione, on an average about 0.5 mumol/g wet wt. Conjugation with glutathione catalysed by glutathione S-transferases may consequently be an important detoxification mechanism in earthworms.

The carabid beetle Pterostichus melanarius is a major natural enemy of pests, such as aphids and slugs in agricultural systems. Earthworms are a dominant non-pest component of the diet of P. melanarius which help sustain the beetles during periods when the pest population is low or absent. In this study we wanted to test whether this predator exercises prey choice among different earthworm species or ecological groups. High levels of genetic diversity within morphological species of earthworm necessitated the development of primers that were specific not just to species but lineages and sub-lineages within species as well. Gut samples from beetles were analysed using multiplex-PCR and fluorescent-labelled primers. Calibratory feeding trials were undertaken to calculate median detection times for prey DNA following ingestion. Extensive testing demonstrated that the primers were species-specific, that detection periods were negatively related to amplicon size and that meal size had a highly significant effect on detection periods. Monte Carlo simulations showed that, in general, worms were being predated in proportion to their densities in the field with little evidence of prey choice, other than probable avoidance of the larger, deep-living species. There was no evidence that epigeic species were being taken preferentially in comparison with endogeic species. There was also no evidence that defensive secretions by Allolobophora chlorotica reduced predation pressure on this species by P. melanarius. We concluded that any management system that increases earthworm densities generally, regardless of component species, is likely to be optimal for increasing numbers of this beneficial beetle predator.