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Type: Article
Published: 2009-04-06
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Species diversity and distributions of microdrile earthworms (Annelida, Clitellata, Enchytraeidae) from South America

Universidade Federal da Paraíba, Departamento de Sistemática e Ecologia, 58059-900, João Pessoa, Paraíba. Brazil
Annelida biodiversity Crassiclitellata Neotropical region Oligochaeta sister group of megadrile earthworms terrestrial taxon

Abstract

A comprehensive biodiversity database of Enchytraeidae produced 66 nominal species reported to date from South America (Achaeta, 6 species; Buchholzia, 1; Cognettia, 1; Enchytraeus, 3; Fridericia, 5; Grania, 1; Guaranidrilus, 10; Hemienchytraeus, 12; Henlea, 2; Lumbricillus, 7; Marionina, 12; Stephensoniella, 1; Timmodrilus, 1; Tupidrilus, 4). Almost 76% of this fauna (50 species) is endemic to the South American continent. The remaining 16 species are more or less largely distributed. Detailed South American occurrences are provided. This is the first reassessment of South American enchytraeids in 27 years. The group is conspicuoulsy absent from the northeast region of Brazil. Enchytraeidae are microdrile earthworms, representing the sister group of the Crassiclitellata, or megadrile earthworms. This relationship is supported by molecular phylogenies, ontogenetic transformations, ecological evidence, and several morphological synapomorphies (dorsal pores, contractil dorsal vessel, lateral position of dorsal chaetal bundles, and shape of chaetae).

References

  1. Baylis, H.A. (1916) Some Nemertinea, free-living Nematoda and Oligochaeta from the Falklands. Annals and Magazine of Natural History Series 8, 17, 288−289.

    Beddard, F.E. (1891) On the anatomy of Ocnerodrilus (Eisen). Transactions of the Royal Society of Edinburgh, 36, 563−583.

    Beddard, F.E. (1895) A monograph of the order Oligochaeta. Clarendon Press, Oxford, 769 pp.

    Bell, A.W. (1962) Enchytraeids (Oligochaeta) from various parts of the world. Transactions of the American Microscopical Society, 81, 158−178.

    Benham, W.B. (1915) Oligochaeta from the Kermadec Islands. Transactions and Proceedings of the New Zealand Institute, 47, 174−185.

    Bittencourt, E.C.R. (1974) Algumas Enchytraeidae (Oligochaeta) de São Paulo. Revista Brasileira de Biologia, 34, 369−378.

    Botea, F. (1987) Oligochaeta from Venezuela. In: Decu, V., Orghidan, T., Dancau, D., Bordon, C., Urbani, O., Linares, F., Tronchoni, J. & Bosque, C. (Eds.), Fauna hipogea y humiedafica de Venezuela y de otros paises de America del Sur. Editura Academiei Republicii Socialiste Romania, Bucarest, pp. 67−72.

    Bretscher, K. (1899) Beitrag zur Kenntniss der Oligochaetenfauna der Schweiz. Revue Suisse de Zoologie, 6, 369−426.

    Buchholz, R. (1863) Beiträge zur Anatomie der Gattung Enchytraeus. Schrift der Physischen und Ökonomischen Gesellschaft im Köningsberg, 3, 93−132, pls. 4−6.

    Černosvitov, L. (1930) Eine neue Enchytraeiden-Art aus den Ameisenhaufen Argentiniens. Zoologischer Anzeiger, 88, 85−89.

    Černosvitov, L. (1934a) Oligochètes. Expedition Antarctique Belge Ressultats du Voyage de la Belgica 1897-99 Rapports Scientifiques Zoologie, 1934, 1−11.

    Černosvitov, L. (1934b) Zur Kenntnis der Enchytraeiden II. Zoologischer Anzeiger, 105, 295–305.

    Černosvitov, L. (1934c) Zur Kenntnis der Enchytraeiden. I. Über Enchytraeus marinus Moore, 1902; Über die Beziehung zwischen den Gattung Enchytraeus und Pachydrilus. Zoologischer Anzeiger, 105, 233−247.

    Černosvitov, L. (1935) Oligochäten aus dem Tropischen Süd-Amerika. Capita Zoologica, 6, 1−36.

    Černosvitov, L. (1937a) System der Enchytraeiden. Bulletin de la Associacion Russe de Recherche Scientifique in Prague, 5, 263−295.

    Černosvitov, L. (1937b) Notes sur les Oligochaeta (naïdidées et enchytraeidées) de l´Argentine. Anales del Museo Argentino de Ciencias Naturales Bernardino Rivadavia Buenos Aires, 39, 135−157.

    Černosvitov, L. (1939) Oligochaeta. The Percy Sladen Trust Expedition to Lake Titicaca in 1937. Transactions of the Linnean Society of London Zoology 3, 1, 81−116.

    Černosvitov, L. (1942) Oligochaeta from various parts of the world. Proceedings of the Zoological Society of London Series B, 111, 197−236.

    Christensen, B. & Dózsa-Farkas, K. (1999). The enchytraeid fauna of the Palearctic tundra (Oligochaeta, Enchytraeidae). Biologiske Skrifter, 52, 1−37.

    Christensen, B. & Dózsa-Farkas, K. (2007) Achaeta pigmentosa sp. n. from Borneo - with dorsal pores (Enchytraeidae, Oligochaeta). Newsletter on Enchytraeidae. No. 10. In: Schlaghamersky, J. (Ed.), Proceedings of the 7th International Symposium on Enchytraeidae, May 25–28, 2006, Bruno, Czech Republic. Folia Facultatis Scientiarum Naturalium Universitatis Masarykianae Brunensis Biologia, 110, 41−52.

    Christoffersen, M.L. (1973) Sistema excretor de Fridericia agricola Moore, 1895 (Oligochaeta, Enchytraeidae). Ciência e Cultura São Paulo, 25, 340.

    Christoffersen, M.L. (1976) Two species of Fridericia Mich., 1889 (Oligochaeta, Enchytraeidae) from Brazil. Boletim de Zoologia da Universidade de São Paulo, 1, 239−256.

    Christoffersen, M.L. (1977) New species of Guaranidrilus Černosv. (Enchytraeidae, Oligochaeta) from Serra do Mar, Săo Paulo, Brasil. Studies of Neotropical Fauna and Environment, 12, 187−206.

    Christoffersen, M.L. (1979a) Achaeta neotropica Černosvitov and A. iridescens sp.n. (Oligochaeta, Enchytraeidae) from Serra do Mar, Săo Paulo, Brasil. Zoologica Scripta, 8, 153−158.

    Christoffersen, M.L. (1979b) Some Enchytraeidae (Oligochaeta) from Serra do Mar, Săo Paulo, Brasil. Boletim de Zoologia da Universidade de São Paulo, 4, 39−52.

    Claparède, E. (1861) Recherches anatomiques sur les annelides, turbellariès, opalines et grégarines observées dans les Hébrides. Mémoires de la Societé de Physique et d´Histoire Naturelle de Geneve, 16, 71−164, pls. 217−291.

    Claus, C. (1880) Grundzüge der Zoologie. Fourth edition. Volume 1. N. G. Elwert´sche Universität, Marburg, 482 pp.

    Coates, K.A. (1989) Phylogeny and origins of Enchytraeidae (Annelida: Oligochaeta). Hydrobiologia, 180, 17−33.

    Coates, K.A. & Diaz, R.J. (1988) Description of Guaranidrilus oregonensis, new species (Oligochaeta: Enchytraeidae) from North America, with additional comments on the genus. Proceedings of the Biological Society of Washington, 101, 773−783.

    Coates, K.A. & Erséus, C. (1985) Marine enchytraeids (Oligochaeta) of the coastal northwest Atlantic (Northern and Mid U.S.A.). Zoological Scripta, 14, 103−116.

    Coates, K.A. & Healy, B. (2008) List of marine species in the family Enchytraeidae (Annelida: Clitellata: Enchytraeidae). Available from: http://www.ihhs.uiuc.edu/~mjwetzel/ENCH/MarEnchList.html (accessed 9 September, 2008)

    Coates, K.A., Locke, J.M., Healy, B. & Wetzel, M.J. (2008) The aphanoneuran and clitellate Annelida occurring in the United States and Canada. Families Enchytraeidae and Propappidae. Available from: http://www.ihhs.uiuc.edu/~mjwetzel/EnchytraeidaeNA.html (accessed 9 September, 2008)

    Coates, K.A. & Stacey, D.F. (1994) Oligochaetes (Naididae, Tubificidae, Enchytraeidae and Alluroididae) of Guyana, Peru and Ecuador. Hydrobiologia, 278, 79−84.

    Cognetti, L. (1900) Viaggio del Dr. A. Borelli nel Mato Grosso e nel Paraguay, II. Contributo alla conoscenza degli Oligocheti Neotropicali. Bolletino du Musei di Zoologia ed Anatomia Comparata de la Real Università di Torino, 15, 1−15.

    Cognetti, L. (1905) Gli oligochaeti della regione neotropicale. I. Memories de la Accademia Reale de Scienze di Torino Serie 2, 55, 1−72.

    Cognetti, L. (1927) Lumbricidi dei Carpazi. Bollettino dell Museum et Laboratoire di Zoologia et Anatomia Comparata della Reale Università di Genova Serie 3, 7 (10), 1−8.

    Dash, M.C. & Senapati, B.K. (1986) Vermitechnology, an option for organic waste management in India. Proceedings of the National Seminar of Organic Waste Utilizing Vermicomposting B, 1986, 157−172.

    De Blainville, H. (1830) Vers. In: Dictionnaire des sciences naturelles, dans lequel on traite méthodiquement des différens êtres de la nature, considérés soit en eaux-memes, d´après l´état actuel de nos connoissances, soit relativement a l´utilité qu´en peuvent retirer la médicine, l´agriculture, le commerce et les arts. Suivi d´une bibliographie des plus célèbres naturalistes. Volume 57. Levrault, Strasbourg, 628 pp.

    Didden, W.A.M. (1993) Ecology of terrestrial Enchytraeidae. Pedobiologia, 37, 2−29.

    Didden, W.A.M. & Römbke, J. (2001) Enchytraeids as indicator organisms for chemical stress in terrestrial ecosystems. Ecotoxicology and Environmental Safety, 50, 25−43.

    Di Pérsia, D. (1980) Aportes a la oligoquetofauna acuática y terrestre de la Provincia de Entre Ríos. Historia Naturalis 1, 7−83.

    Dózsa-Farkas, K. (1989) Neue Enchytraeiden-Arten (Oligochaeta) aus Ekuador. Acta Zoologica Hungarica, 35, 191−203.

    Dózsa-Farkas, K. (1996) Self-fertilization: an adaptive strategy in widespread enchytraeids. European Journal of Soil Biology, 31, 207−215.

    Dózsa-Farkas, K. (1997) Timmodrilus gen.n., a genus from the family Enchytraeidae (Oligochaeta). Opuscula Zoologica Budapest, 29–30, 49−52.

    D´Udekem, J. (1854) Description d´une nouvelle espèce d´Enchytraeus. Bulletin de la Academie Royal des Sciences Lettres et Beaux-Arts de Belgique Series 1, 21, 853−864, 1 pl.

    Ehrenberg, C.G. (1828) Symbolae physicae, seu Icones et descriptiones Corporum Naturalium novorum aut minus cognitorum, quae ex itineribus per Libyam, Aegyptum, Nubiam, Dongalam, Syriam, Arabiam et Habessiniam. Pars Zoologia; Animalia evertebrata, exclusis insectis. Berlin.

    Eisen, G. (1878) Redogörelse för Oligochaeter, samlade under de Svnska expeditionerna till Arktiska trakter. Förhandlinga und Öfversight der Kongliga Svenska Vetenskapps Akademie, 35, 63−79.

    Eisen, G. (1904) Enchytraeidae. Harriman Alaska Expedition Series 12, 1−166, 20 pls.

    Erséus, C. (1980) Specific and generic criteria in marine Oligochaeta, with special emphasis on Tubificidae. In: Brinkhurst, R.O. & Cook, D.G. (Eds.), Aquatic oligochaete biology. Plenum Press, New York, pp. 9−24.

    Erséus, C. (1988) Oligochaeta. In: Higgins, R. & Thiel, H. (Eds.), Introduction to the study of meiofauna. Smithsonian Intitution Press, Washington, D.C., pp. 349−354.

    Erséus, C. (2005) Phylogeny of oligochaetous Clitellata. Hydrobiologia, 535, 357−372.

    Erséus, C. & Källersjö, M. (2004) 18S rDNA phylogeny of Clitellata (Annelida). Zoologica Scripta, 33, 187−196.

    Erséus, C. & Lassèrre, P. (1976) Taxonomic status and geographic variation of the marine enchytraeid genus Grania Southern (Oligochaeta). Zoologica Scripta, 5, 121−132.

    Friend, H. (1911) New British Henleas. Zoologist London Series 4, 15, 464−468.

    Gavrilov, K. (1979) Oligochaeta. In: Hurlbert, S.H. (Ed.), Biota acuatica de Sudamerica Austral. San Diego State University, San Diego, pp. 99−121.

    Gavrilov, K. (1981) Oligochaeta. In: Hurlbert, S.H., Rodriguez, G. & Dos Santos, N.D. (Eds.), Aquatic biota of tropical South America. Being a compilation of taxonomic bibliographies for the fauna and flora of inland waters of the tropical portion of South America. Part 2. Anarthropoda. San Diego University, San Diego, pp. 170−190.

    Graefe, U. (1993a) Die Gliederung von Zersetzergesellschaften für die standortsökologische Ansprache. Mitteilungen der Deutsche Bodenkunde Geselschaft, 69, 95−98.

    Graefe, U. (1993b) Verhänderungen der Zersetzegesellschaften im Immissionsbereich eines Zementwerkes. Mitteilungen der Deutsche Bodenkunde Geselschaft, 72, 531−534.

    Grube, A.E. (1850) Die Familien der Anneliden. Archiv für Naturgeschicht, 16, 249−364.

    Healy, B. (1979) Review of the genus Guaranidrilus (Oligochaeta, Enchytraeidae) with the description of two new species. Bulletin of the British Museum of Natural History Zoology, 37, 7−15.

    Henle, F. (1837) Über Enchytraeus, eine neue Anneliden Gattung. Archiv für Anatomie und Physiologie Wissenschaftliche Medizin, 1837, 74−90, pl. 6.

    Jackson, R.M. & Raw, F. (1974) La vida en el suelo. Editora Omega, Barcelona, 70 pp.

    Jänsch, S., Römbke, J. & Didden, W.A.M. (2005) The use of enchytraeids in ecological soil classification and assessment concepts. Ecotoxicology and Environmental Safety, 62, 266−277.

    Jamieson, B.G.M. (1981) The ultrastructure of the Oligochaeta. Academic Press, London, 462 pp.

    Jamieson, B.G.M. (1988) On the phylogeny and higher classification of the Oligochaeta. Cladistics, 4, 367−410.

    Kasprzak, K. (1984a) Generic criteria in Enchytraeidae (Oligochaeta) family. Biologia Bratislava, 39, 163−172.

    Kasprzak, K. (1984b) The previous and contemporary conceptions on phylogeny and systematic classifications of Oligochaeta (Annelida). Annales Zoologici Warszawa, 38, 205−223.

    Lavelle, P. & Lapied, E. (2003) Endangered earthworms of Amazonia: an homage to Gilberto Righi. Pedobiologia, 47, 419−427.

    Levinsen, G.M.R. (1884) Systematisk-geografisk öfverssigt över de norddiske Annulata, Gephyrea, Chaetognathi og Balanoglossi, II. Videnskabelige Meddelser fra den Naturhistorische Forening in Kjobenhavn, 9-11, 9−350, pls 1−7.

    Ljungström, P.O. (1970) Introduction to the study of earthworm taxonomy. Pedobiologia, 10, 265−285.

    Lüderwaldt, H. (1927) A colleção de minhocas (Oligochaeta) do Museu Paulista com notas biológicas. Revista do Museu Paulista, 15, 545−556.

    Marcus, E. (1965) Naidomorpha aus brasilianischem Brackwasser. Beiträge zür Neotropische Fauna, 4, 61−83.

    Martin, P. (2001) On the origin of the Hirudinea and the demise of the Oligochaeta. Proceedings of the Royal Society of London Series B Biological Science, 268, 1089−1098.

    Martínez-Ansemil, E. & Giani, N. (1986) Algunos oligoquetos acuáticos de Bolivia. Oecologia Aquatica, 8, 107−115.

    Michaelsen, W. (1886a) Über Chylusgefäss systeme bei Enchytraeiden. Archiv für Mikroscopische Anatomie, 28, 292−304, 21 pls.

    Michaelsen, W. (1886b) Untersuchungen über Enchytraeus Moebii Mich. und andere Enchytraeiden. Lipsius und Tischer, Kiel, 50 pp.

    Michaelsen, W. (1888) Oligochaeten des Naturhistorischen Museums in Hamburg, II. Jahrbüch der Hamburgischen Wissenschaftlichen Anstalten, 6, 57−69, 1 pl.

    Michaelsen, W. (1889) Synopsis der Enchytraeiden. Abhandlungen der Gesellschaft in Hamburg, 11, 1−60, 1 pl.

    Michaelsen, W. (1900) Oligochaeta. Das Tierreich, 10. Frieländer und Sohn, Berlin, 575 pp.

    Michaelsen, W. (1905) Oligochaeten der Deutschen Südpolar-Expedition, 1901–1903. Nebst Erörterung der Hypothese über einen früheren grossen, die Südspitzen der Kontinente verbindenden antarktischen Kontinent. Deutsche Südpolar Expedition 1901-1903 Zoology 1 9, 1−58.

    Michaelsen, W. (1913) The Oligochaeta of Natal and Zululand. Annals of the Natal Government Museum, 13, 397−458, 23 pls.

    Michaelsen, W. (1914) Die Oligochaeten Columbias. Voyage d´exploratión scientifique em Colombie. Mémoires de la Societé des Sciences Naturelles de Neuchätel Nouvelle Série, 5, 202−252, 8 pls.

    Michaelsen, W. (1927) Die Oligochätenfauna Brasiliens. Abhandlungen von der Senckenbergischen Naturforschenden Gesellschaft, 40, 369−374.

    Michaelsen, W. (1928) Ordnung der Clitellata: Oligochaeta = Regenwürmer und Verwandte. In: Kükenthal, W. & Krumbach, T. (Eds.), Handbuch der Zoologie, Band 2 (2) Lief 8 und Nachtrag. Walter de Gruyter, Berlin, pp. 1–116 .

    Moore, J.P. (1895) Notes on American Enchytraeidae I. New species of Fridericia from the vicinity of Philadelphia. Proceedings of the Academy of Natural Sciences of Philadelphia, 1895, 341−345, pl. 13.

    Moore, J.P. (1902) Some Bermuda Oligochaeta, with a description of a new species. Proceedings of the Academy of Natural Sciences of Philadelphia, 54, 80−84.

    Müller, O.F. (1774) Vermium terrestrium et fluviatilium, seu animalium infusoriorum, helminthicorum, et testaceorum, non marionorum, succinta historia. Pars altera. Helminthica. Heineck and Faber, Havniae et Lipsiae, 72 pp.

    Nakamura, Y. (2000) Checklist of enchytraeids (Oligochaeta: Enchytraeidae) of the world. Miscellaneous Publications of the Tohoku National Agricultural Experimental Station, 24, 29−104.

    Nielsen, C.O. & Christensen, B. (1959) The Enchytraeidae. Critical revision and taxonomy of European species (Studies on Enchytraeidae VII). Natura Jutlandica, 8-9, 1−160.

    Nielsen, C.O. & Christensen, B. (1961) The Enchytraeidae. Critical revision and taxonomy of European species Supplement 1. Natura Jutlandica 10 (1), 1−23.

    Nielsen, C.O. & Christensen, B. (1963) The Enchytraeidae. Critical revision and taxonomy of European species Supplement 2. Natura Jutlandica, 10 (3), 1−19.

    Ørsted, A.S. (1844) De regionibus marinis. Elementa topographiae historico-naturalis freti eresund. Inaugural Thesis, Kobenhavn, Havniae, 88 pp., 2 pls.

    Omodeo, P. (2000) Evolution and biogeography of megadriles (Annelida, Clitellata). Italian Journal of Zoology, 67, 179−201.

    Rafinesque, C.S. (1815) Analyse de la nature ou tableau de l´univers et des corps organisés. Palerme, 224 pp.

    Reynolds, J.W. (1994) Earthworms of the world. Global Biodiversity, 4, 11−16.

    Reynolds, J.W. & Eggen, A.B. (1993) Earthworm biology and vermi-composting I. Sir Sandford Fleming College, Lindsay, 72 pp.

    Richards, K.S. (1977) Structure and function in the oligochaete epidermis (Annelida). In: Spearman, R.C. (Ed.), Comparative biology of the skin. Academic Press, New York, pp. 171−193.

    Righi, G. (1973) Sobre três espécies brasileiras de Enchytraeidae (Oligochaeta). Boletim de Zoologia e Biologia Marinha Nova Série, 30, 469−482.

    Righi, G. (1974a) Notas sobre os Oligochaeta, Enchytraeidae do Brazil. Papéis Avulsos de Zoologia São Paulo, 28, 127−145.

    Righi, G. (1974b) Marionina pituca, sp.n. (Oligochaeta, Enchytraeidae) from Brazilian soil and its interstitial parasite Buetschliella marioninae, sp.n. (Ciliata, Astomata). Zoologischer Anzeiger, 192, 414−419.

    Righi, G. (1975) Algumas Enchytraeidae (Oligochaeta) brasileiras. Ciência e Cultura São Paulo, 27, 143−150.

    Righi, G. (1978) Notas sobre os Oligochaeta da Amazônia. Acta Amazônica, 8, 485−488.

    Righi, G. (1981a) Notas sobre Enchytraeidae (Oligochaeta) brasileiras. Revista Brasileira de Biologia, 41, 427−430.

    Righi, G. (1981b) Algumas Oligochaeta cavernícolas do Equador. Papéis Avulsos de Zoologia São Paulo, 34, 235−249.

    Righi, G. (1988) Dois novos Microdrili, Oligochaeta, terrestres da Amazonia. Papéis Avulsos de Zoologia São Paulo, 36, 315−321.

    Righi, G. (1995) Colombian earthworms. Studies on Tropical Andean Ecosystems, Volume 4. Cramer, Berlin, 485−607 pp.

    Righi, G., Ayres, I. & Bittencourt, E.C.R. (1978) Oligochaeta (Annelida) do Instituto Nacional de Pesquisas da Amazônia. Acta Amazônica Suplemento, 1, 1−49.

    Römbke, J. (1991) Estimates of the Enchytraeidae (Oligochaeta, Annelida) contribution to energy-flow in the soil system of an acid beech wood forest. Biology and Fertility of Soils, 11, 255−260.

    Römbke, J. (2003) The role of Gilberto Righi in the development of tropical microdrile taxonomy. Pedobiologia, 47, 405−412.

    Römbke, J. (2007) Enchytraeidae of tropical soils: State of the art, with special emphasis on Latin America. Newsletter on Enchytraeidae. No. 10. In: Schlaghamersky, J. (Ed.) Proceedings of the 7th International Symposium on Enchytraeidae, May 25-28, 2006, Bruno, Czech Republic. Folia Facultatis Scientiarum Naturalium Universitatis Masarykianae Brunensis Biologia, 110, 157−181.

    Römbke, J. & Hanagarth, W. (1994) The present faunistic knowledge on terrestrial Oligochaeta from Bolivia. Andrias, 13, 7−16.

    Römbke, J., Meller, M. & Garcia, M. (1999) Earthworm densities in central Amazonian primary and secondary forests and a polyculture forestry plantation. Pedobiologia, 43, 518−522.

    Rosa, D. (1887) Il Neoenchytraeus bulbosus n. sp. Bollettino dell Museum di Zoologia et Anatomia Comparata dell Reale Università di Torino, 2(29), 1−3.

    Rosa, D. (1891) Die exotischen terricolen des K. K. naturhistorischen Hofmuseums. Annalen des K. K. Naturhistorischen Hofmuseums Wien, 6, 379−406, pls 13−14.

    Rota, E. & Brinkhurst, R.O. (2000) Mesenchytraeus antaeus, a new giant enchytraeid (Annelida, Clitellata) from the temperate rainforest of British Columbia, Canada, with a revised diagnosis of the genus Mesenchytraeus. Journal of Zoology London, 252, 27−40.

    Rota, E. & Healy, B. (1994) The enchytraeid fauna of North Africa. Hydrobiologia, 278, 53−66.

    Rota, E., Matamoros, L. & Erséus, C. (2008) In search of Marionina (Clitellata, Enchytraeidae): A taxonomic history of the genus and re-description of the type species Pachydrilus georgianus Michaelsen, 1888. Italian Journal of Zoology, 75, 1−20 Available from: http://dx.doi.org/10.1080/11250000801930433 (accessed 9 September, 2008)

    Schmelz, R.M. (1996) Species separation and identification in the Enchytraeidae (Oligochaeta, Annelida): Combining morphology with general protein data. Hydrobiologia, 334, 31−36.

    Schmelz, R.M. (2003) Taxonomy of Fridericia (Oligochaeta, Enchytraeidae). Revision of species with morphological and biochemical methods. Abhandlungen der Naturwissenchaften Vereins in Hamburg NF, 38, 1−414.

    Schmelz, R.M. & Collado, R. (2005) Achaeta becki sp. nov. (Oligochaeta: Enchytraeidae) from Amazonian forest soils. Zootaxa, 1084, 49−57.

    Schmelz, R.M. & Collado, R. (2007) Revision of Hemienchytraeus stephensoni (Cognetti, 1927) (Enchytraeidae, Oligochaeta, Annelida). Newsletter on Enchytraeidae. In: Schlaghamersky, J. (Ed.), No. 10: Proceedings of the 7th International Symposium on Enchytraeidae, May 25–28, Brno, Czech Republic. Folia Facultatis Scientiarum Naturalium Universitatis Masarykianae Brunensis Biologia, 110, 67−85.

    Schmelz, R.M. & Collado, R. (2008) A type-based redescription of Pachydrilus georgianus Michaelsen, 1888, the type species of Marionina Michaelsen, 1890, with comments on Christensenidrilus Dózsa-Farkas & Convey, 1998 (Enchytraeidae, Oligochaeta, Annelida). Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg (Neue Folge), 44, 7−22.

    Schmelz, R.M., Collado, R. & Römbke, J. (2008) Mata Atlântica enchytraeids (Paraná, Brazil): The genus Achaeta (Oligochaeta, Enchytraeidae). Zootaxa, 1809, 1−35.

    Schmelz, R.M. & Römbke, J. (2005) Three new species of Hemienchytraeus (Enchytraeidae, Oligochaeta) from Amazonian forest soil. Journal of Natural History, 39, 2967−2986.

    Siddall, M.E., Apakupakul, K., Burreson, E.M., Coates, K.A., Erséus, C., Gelder, S.R., Källerjö, M. & Trapido-Rosenthal, H. (2001) Validating Livanow: Molecular data agree that leeches, branchiobdellidans and Acanthobdella peledina form a monophyletic group of oligochaetes. Molecular Phylogenetics and Evolution, 21, 346−351.

    Sims, R.W. (1980) A classification and the distribution of earthworms, suborder Lumbricina (Haplotaxida: Oligochaeta). Bulletin of the British Museum Natural History Zoology, 39, 103−124.

    Southern, R. (1913) Clare Island Survey. Part 48. Oligochaeta. Proceedings of the Royal Irish Academy Section B, 31, 1−14.

    Stacey D.F. & Coates, K.A. (1996) Oligochaetes (Naididae, Tubificidae, Opistocystidae, Enchytraeidae, Sparganophilidae and Alluroididae) of Guyana. Hydrobiologia, 334, 17−29.

    Stephenson, J. (1930a) The Oligochaeta. Oxford University Press, Oxford, 978 pp.

    Stephenson, J. (1930b) On some African oligochaetes. Archivio Zoologico Italiano, 14, 485−511.

    Stephenson, J. (1932) Oligochaeta I. Microdrili, mainly Enchytraeidae. Discovery Reports, 4, 233−264.

    Ude, H. (1896) Enchytraeiden. Die Oligochaeten des Südlichen Süd-Amerikas und geographische Beziehung der selben. Ergebnisse der Hamburger Malgalhaensischen Sammelreise, 1. Fridrichsen, Hamburg, 43 pp.

    Ulrich, H. & Schmelz, R.M. (2001) Enchytraeidae as prey of Dolichopodidae, recent and in Baltic amber (Oligochaeta, Diptera). Bonner Zoologische Beiträge, 50, 89−101.

    Vejdovský, F. (1877) Zur Anatomie und Systematik der Enchytraeiden. Sitzungsberichte der Königlichen Böhmischen Gesellschaft Wissenschaften Prag, 1877, 294−304.

    Vejdovský, F. (1879) Beiträge zur vergleichenden Morphologie der Anneliden. I. Monographie der Enchytraeiden. Friedrich Temspky, Prag, 61 pp.

    Xie, Z.C., Wang, H.Z. & Liang, Y.L. (1999) Studies on the Enchytraeidae of China I. On new species and new records of the genus Hemienchytraeus. Acta Hydrobiologica Sinica, 23, 352−358.