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Type: Article
Published: 2016-08-19
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Lavellodrilus notosetosus sp. nov. (Annelida, Crassiclitellata, Acanthodrilidae): a new Mexican earthworm with uncommon characters, revealed by a preliminary revision of subfamily Acanthodrilinae

Laboratorio de Invertebrados del Suelo. Red de Biodiversidad y Sistemática. Instituto de Ecología A. C. Carretera Antigua a Coatepec No. 351. Col. El Haya. Xalapa, Veracruz. C.P. 91070 MEXICO
Laboratorio de Invertebrados del Suelo. Red de Biodiversidad y Sistemática. Instituto de Ecología A. C. Carretera Antigua a Coatepec No. 351. Col. El Haya. Xalapa, Veracruz. C.P. 91070 MEXICO
Annelida Acanthodrilidae Clitellata Oligochaeta Chiapas Lacandon forest genital setae

Abstract

A new acanthodriline earthworm species, Lavellodrilus notosetosus sp. nov., is described from tropical rain forests of southern Mexico. The new species is placed within the genus Lavellodrilus by the presence of mesial spermathecal pores. It is separated from other species of the genus by the dorsal location of setae cd in most of the body, last hearts in segment 13, first intestinal segment in 20 and genital setae in segment 12. A preliminary morphological revision of all genera and species of Acanthodrilinae was undertaken in order to: i) evaluate if the mesial spermathecal pores justify the status of Lavellodrilus, ii) determine how common (expressed as percentages of species having the character) the diagnostic characters of the new species are in the subfamily, iii) clarify if these characters exhibit a geographical pattern, and iv) contribute towards a comprehensive analysis of the Acanthodrilinae. In this revision, species were separated in nine geographical regions: USA, northern Mexico, southern Mexico, Caribbean Islands (northern hemisphere), and South America, South Africa, Madagascar, Australia, New Zealand, New Caledonia and Antarctic Islands (southern hemisphere). As a result of the revision it was found that among the 511 recognized species of Acanthodrilinae only 11 species have a mesial location of the spermathecal pores, in two cases probably representing monophyletic groups (Lavellodrilus and a group of South African Parachilota species). It was also found that the distinguishing characters in L. notosetosus sp. nov., notably the location of last hearts, genital setae and the first intestinal segment, are uncommon characters in the acanthodriline earthworm fauna of southern Mexico and Central America, but more frequent in North America, the Caribbean, and the southern hemisphere. We conclude that the acanthodrilines from the northern hemisphere are morphologically more similar to those from Australia, New Zealand, and New Caledonia than to those of South Africa and South America.

 

References

  1. Beddard, F.E. (1890) Contributions to the anatomy of earthworms, with descriptions of some new species. Quarterly Journal of Microscopical Science, 30 (120), 421–479.

    Beddard, F.E. (1895) Preliminary account of new species of earthworms belonging to the Hamburg Museum. Proceedings of the Zoological Society of London, 1895, 210–239.
    http://dx.doi.org/10.1111/j.1469-7998.1895.tb00005.x

    Benham, W.B. (1903) On an earthworm from the Auckland Islands – Notiodrilus aucklandicus. Transactions and Proceedings of the New Zealand Institute, 35 (30), 275–277.

    Benham, W.B. (1905) On the Oligochaeta from the Southern Islands of the New Zealand Region. Transactions and Proceedings of the New Zealand Institute, 37 (19), 285–297.

    Blakemore, R.J. (2000) Tasmanian earthworms. CD-ROM Monograph with Review of World. Families. ‘VermEcology’, Canberra, 800 pp.

    Blakemore, R.J. (2005) Whither Octochaetidae? – its family status reviewed. In: Pop, V.V. & Pop, A.A.. (Eds.), Advances in Earthworm Taxonomy. II. Proceedings of the 2nd International Oligochaete Taxonomy Meeting. Cluj University Press: Cluj-Napoca, pp. 63–84.

    Blakemore, R.J. (2007) Checklist and phylogeny of Exxidae Blakemore, 2000 (Oligochaeta). European Journal of Soil Biology, 43, 9–11.

    Blakemore, R.J. (2013) The major megadrile families of the world reviewed again on their taxonomic types (Annelida: Oligochaeta: Megadrilacea). Opuscula Zoologica Budapest, 44 (2), 107–127.

    Borges, S. & Moreno, A. (1991) Nuevas especies del género Trigaster Benham, 1886 (Oligochaeta: Octochaetidae) para Puerto Rico. Bolletino del Museo Regionale di Scienze Naturali- Torino, 9 (1), 39–54.

    Borges, S. & Moreno, A. (1992) Redescripción de Trigaster rufa Gates 1962 (Oligochaeta: Octochaetidae). Caribbean Journal of Science, 28 (1–2), 47–50.

    Buckley, T.R., James, S., Allwood, J., Bartlett, S., Howitt, R. & Prada, D. (2011) Phylogenetic analysis of New Zealand earthworms (Oligochaeta: Megascolecidae) reveals ancient clades and cryptic taxonomic diversity. Molecular Phylogenetics and Evolution, 58, 85–96.
    http://dx.doi.org/10.1016/j.ympev.2010.09.024

    Černosvitov, L. (1935) Oligochètes. Résultats du voyage du S.Y. Belgica en 1897 – 1899. Expédition Antarctique Belge. Rapports Scientifiques Zoologie, 5, 1–11.

    Černosvitov, L. (1939) The Percy Sladen Expedition to Lake Titicaca.VI. Oligochaeta. Transactions of the Linnean Society of London, 1, 81–116.
    http://dx.doi.org/10.1111/j.1096-3642.1939.tb00006.x

    Csuzdi, C.S. (2010) A monograph of the Paleotropical Benhamiinae earthworms. Hungarian Natural History Museum & Hungarian Academy of Sciences, Budapest, 348 pp.

    Csuzdi, C.S. & Szlávecz, K. (2002) Diplocardia patuxtensis sp. n., a new earthworm species from Maryland, North America (Oligochaeta: Acanthodrilidae). Annales Historico-Naturales Musei Nationalis Hungarici, 94, 45–51.

    Csuzdi, C.S. & Mischis, C. (2010) Earthworms from Argentinean Patagonia with description of two remarkable new species (Oligochaeta: Acanthodrilidae, Lumbricidae and Megascolecidae). Journal of Natural History, 44 (1–2), 31–40.

    Dyne, G.R. (1980) Rhododrilus sexpapillatus n.sp. (Oligochaete: Megascolecidae) from New Zealand. New Zealand Journal of Zoology, 7, 505–508.

    Dyne, G.R. & Jamieson, B.G.M. (2004) Native Earthworms of Australia II (Megascolecidae, Acanthodrilinae). CD monograph. Department of the Environment and Heritage, Australian Government, NSW, 200 pp.

    Eisen, G. (1896) Pacific Coast Oligochaeta II. Memoirs of the California Academy of Sciences, 2 (5), 123–198.

    Eisen, G. (1899) Notes on North-American Earthworms of the Genus Diplocardia. Zoological Bulletin, 2, 161–172.
    http://dx.doi.org/10.2307/1535422

    Eisen, G. (1900) Researches in the American Oligochaeta, with special reference to those of the Pacific coast and adjacent islands. Proceedings of the California Academy of Sciences, 2 (3), 85–276.

    Fragoso, C. (1988) Sistemática y ecología de un género nuevo de lombriz de tierra (Acanthodrilini, Oligochaeta) de la Selva Lacandona, Chiapas. México. Acta Zoológica Mexicana, New Series, 25, 1–34.

    Fragoso, C. (1991) Two new species of the earthworm genus Lavellodrilus (Oligochaeta, Acanthodrilini) from tropical mexican rain forests. Studies on Neotropical fauna and Environment, 26 (2), 83–91.
    http://dx.doi.org/10.1080/01650529109360838

    Fragoso, C. (1992) Las lombrices terrestres de la Selva Lacandona, Ecología y Potencial Práctico. In: Vásquez-Sánchez, M.A. & Ramos, M.A. (Eds.), Reserva de la Biosfera Montes Azules, Selva Lacandona: Investigación para su uso. Ecósfera, Publicaciones Especiales, 1, pp. 101–118.

    Fragoso, C. (2001) Las lombrices de Tierra de México (Annelida, Oligochaeta): Diversidad, Ecología y Manejo. Acta Zoológica Mexicana, New Series, Número Especial 1, 131–171.

    Fragoso, C. (2007) Diversidad y patrones biogeográficos de las lombrices de tierra de México (Oligochaeta, Annelida). In: Brown, G.G. & Fragoso, C. (Eds.), Minhocas na America Latina: Biodiversidade e Ecologia. EMBRAPA Soja, Londrina, pp. 107–124.

    Fragoso, C. (2011) Lombrices de Tierra (Annelida: Oligochaeta). In: Cruz, A.A., Lorea, H.F., Hernández, O.V. & Morales, J.E. (Eds.), La Biodiversidad en Veracruz: Estudio de Estado. Vol II. Diversidad de especies: Conocimiento Actual. Comisión Nacional para el Conocimiento y Uso de la Biodiversidad, México, D.F.; Gobierno del Estado de Veracruz de Ignacio de la Llavepp, Xalapa; Universidad Veracruzana, Xalapa; Instituto de Ecología, A.C., Xalapa, pp. 259–268.

    Fragoso, C. & Lavelle, P. (1987) The earthworm community of a mexican tropical rain forest (Chajul, Chiapas). In: Bonvicini Pagliai, A.M. & Omodeo, P. (Eds.), On Earthworms. Selected Symposia and Monographs U.Z.I. Vol. 1. Mucchi, Modena, pp. 281– 295.

    Fragoso, C. & Rojas, P. (1994) Earthworms from southeastern Mexico. New acanthodriline genera and species (Megascolecidae, Oligochaeta). Megadrilogica, 6 (1), 1–12.

    Fragoso, C. & Rojas, P. (2007) Two new species of the earthworm genus Balanteodrilus (Oligochaeta: Acanthodrilidae) from Eastern Mexico. Megadrilogica, 11, 107–114.

    Fragoso, C. & Rojas, P. (2009) A new ocnerodrilid earthworm genus from Southeastern Mexico (Annelida: Oligochaeta), with a key for the genera of Ocnerodrilidae. Megadrilogica, 13 (9), 141–152.

    Fragoso, C. & Rojas, P. (2014) New species and records of the earthworm genus Ramiellona (Annelida, Oligochaeta, Acanthodrilidae) from southern Mexico and Guatemala. Zootaxa, 3753 (6), 549–572.

    Fragoso, C., Brown, G., Patrón, J.C., Blanchart, E., Lavelle, P., Pashanasi, B., Senapati, B. & Kumar, T. (1997) Agricultural intensification, soil biodiversity and agroecosystem function in the tropics: the role of earthworms. Applied Soil Ecology, 6, 17–35.
    http://dx.doi.org/10.1016/S0929-1393(96)00154-0

    Fragoso, C., Kanyonyo, J., Moreno, A., Senapati, B., Blanchart, E., Rodríguez, C. & Rodríguez, C. (1999) A Survey of Tropical Earthworms: Taxonomy, Biogeography and Environmental Plasticity. In: Lavelle, P., Brussaard, L. & Hendrix, P. (Eds.), Earthworm Management in Tropical Agroecosystems. CAB International, New York, pp. 1–26.

    Gates, G. (1943) On some American and Oriental earthworms. Part II. Family Megascolecidae. The Ohio Journal of Science, 43 (3), 99–116.

    Gates, G. (1954) Exotic earthworms of the United States. Bulletin of the Museum of Comparative Zoology, 111 (6), 219–258.

    Gates, G. (1955) Notes on several species of the earthworm genus Diplocardia Garman 1888. Bulletin of the Museum of Comparative Zoology, 113 (3), 227–260.

    Gates, G. (1959) On a taxonomic puzzle and the classification of the earthworms. Bulletin of the Museum of Comparative Zoology, 121 (6), 229–261.

    Gates, G. (1962) On a new species of the earthworm genus Trigaster Benham 1886 (Octochaetidae). Breviora, 178, 1–4.

    Gates, G. (1967) On a new species of earthworm from a Mexican cave. International Journal of Speleology, 3, 63–70.
    http://dx.doi.org/10.5038/1827-806X.3.1.5

    Gates, G. (1970) On a new species of earthworm from another Mexican cave. The Southwestern Naturalist, 15 (2), 261–273.
    http://dx.doi.org/10.2307/3670359

    Gates, G. (1971) On some earthworms from Mexican caves. Association for Mexican Cave Studies Bulletin, 4, 3–8.

    Gates, G. (1973) On more earthworms from Mexican caves. Association for Mexican Cave Studies Bulletin, 5, 21–24.

    Gates, G. (1977) More on the earthworm genus Diplocardia. Megadrilogica, 3 (1), 1–48.

    Graff, O. (1957) Regenwürmer aus El Salvador (Oligochaeta). Senckenbergiana Biologica, 38, 115–143.

    Iturralde-Vinent, M. (2006) El origen paleogeográfico de la biota de Guatemala. In: Cano, E.B. (Ed.), Biodiversidad de Guatemala. Vol. I. Universidad del Valle de Guatemala, Guatemala City, pp. 1–6.

    James, S.W. (1988) Diplocardia hulberti and D. rugosa, new earthworms (Annelida: Oligochaeta: Megascolecidae) from Kansas. Proceedings of the Biological Society of Washington, 101 (2), 300–307.

    James, S.W. (1990a) Diplocardia kansensis, a new earthworm from Kansas, with redescriptions of D. riparia Smith and D. fuscula Gates (Annelida: Oligochaeta: Megascolecidae). Proceedings of the Biological Society of Washington, 103 (1), 179–186.

    James, S.W. (1990b) Diplotrema murchiei and D. papillata new earthworms (Oligochaeta: Megascolecidae) from Mexico. Acta Zoologica Mexicana, New Series, 38, 18–27.

    James, S.W. (1991) New species of earthworms from Puerto Rico, with a redefinition of the earthworm genus Trigaster (Oligochaeta: Megascolecidae). Transactions of the American Microscopical Society, 110 (4), 337–353.
    http://dx.doi.org/10.2307/3226771

    James, S.W. (1993) New acanthodriline earthworms from Mexico (Oligochaeta: Megascolecidae). Acta Zoologica Mexicana, New Series, 60, 1–21.

    James, S.W. (1994) New species of Diplocardia and Argilophilus (Annelida: Oligochaeta: Megascolecidae) from Southern California. Proceedings of the Biological Society of Washington, 107 (4), 622–633.

    James, S.W. & Davidson, S. (2012) Molecular phylogeny of earthworms (Annelida: Crassiclitellata) based on 28S, 18S and 16S gene sequences. Invertebrate Systematics, 26, 213–229.
    http://dx.doi.org/10.1071/IS11012

    Jamieson, B.G.M. (1971) A review of the megascolecoid earthworm genera (Oligochaeta) of Australia. Part II — The subfamilies Ocnerodrilinae and Acanthodrilinae. Proceedings of the Royal Society of Queensland, 82, 95–108.

    Jamieson, B.G.M. (1974) Generic type-species and other Megascolecidae (Annelida, Oligochaeta) in the Museum of Comparative Zoology, University of Turin. Bolletino del Museo di Zoologia dell'Università di Torino, 8, 57–88.

    Jamieson, B.G.M. & Bennett, J.D. (1979) New species of Acanthodrilinae and a new genus of Perionychini (Megascolecidae: Oligochaeta) from New Caledonia, their phylogeny and zoogeography. Bulletin du Muséum National d'Histoire Naturelle, 2, 335–403.

    Jamieson, B.G.M., MacDonald, K.R. & James, S. (2015) A new species of Diplotrema (Acanthodrilinae, Metagynophora, Crassiclitellata, Oligochaeta) from the Einasleigh Uplands Bioregion of Queensland. Memoirs of the Queensland Museum, 59 (1), 171–176.
    http://dx.doi.org/10.17082/j.2204-1478.59.2015.2014-12

    Jamieson, B.G.M., Tillier, S., Tillier, A., Justine, J.-L., Ling, E., James, S.W., McDonald, K. & Hugall, A.F. (2002) Phylogeny of the Megascolecidae and Crassiclitellata (Annelida, Oligochaeta): combined versus partitioned analysis using nuclear (28S) and mitochondrial (12S, 16S) rDNA. Zoosystema, 24 (4), 707–734.

    Julka, J.M. (1988) Fauna of India. Megadrile Oligochaeta (Earthworms). Vol. 1. Family Octochaetidae. Zoological Survey of India, Calcutta, 400 pp.

    Lee, K.E. (1959) The Earthworm Fauna of New Zealand. New Zealand Department of Scientific and Industrial Research Bulletin, 130, 1–486.

    Lee, K.E. (1968) Oligochaeta from Subantarctic Islands. In: Thomas P.M. B.A.N.Z. Antarctic Research Expedition 1929–1931. Reports - Series B (Zoology and Botany) 8 (8) (Mawson Institute for Antarctic Research, University of Adelaide, Adelaide), pp. 149–165.

    Ljungström, P.O. (1969) On the earthworm genus Udeina (Acanthodrilidae, Oligochaeta) in South Africa. Zoologischer Anzeiger, 182, 370–379.

    Ljungström, P.O. (1972) Taxonomical and ecological notes on the earthworm genus Udeina and a requiem for the South African acanthodrilines. Pedobiologia, 12 (2), 100–110.

    López-Gamund, O.R. & Buatois, L.A. (2010) Introduction: Late Paleozoic glacial events and postglacial transgressions in Gondwana. In: López-Gamund, O.R. & Buatois, L.A. (Eds.), Late Paleozoic Glacial Events and Postglacial Transgressions in Gondwana. Geological Society of America Special Paper, 468, pp. v–viii.
    http://dx.doi.org/10.1130/2010.2468(00)

    Macnab, J.A. & Mckey-Fender, D. (1955) Studies in the genus Diplocardia (Oligochaeta: Megascolecidae). The Wasmann Journal of Biology, 13 (1), 113–143.

    Michaelsen, W. (1900) Oligochaeta. In: Das Tierreich. Lief. 10. R. Friedländer & Sohn, Berlin, pp. 1–575.

    Michaelsen, W. (1903) Neue Oligochaeten und neue Fundorte altbekannter. Mitteilungen aus dem Naturhistorischen Museum in Hamburg 19, 1–55.

    Michaelsen, W. (1904) Catálogo de los Oligoquetos del territorio chileno-magallánico y descripción de especies nuevas. Revista Chilena de Historia Natural, 8, 262–292.

    Michaelsen, W. (1905a) Die Oligochaeten der schwedischen Südpolar–Expedition. Wissenschaftliche Ergebnisse der Schwedischen Südpolar-Expedition 1901–1903, V, 1– 12.

    Michaelsen, W. (1905b) Die Oligochaeten der deutschen Südpolar-Expedition 1901–1903. Deutsche Südpolar-Expedition 1901–1903, IX, 1–58.

    Michaelsen, W. (1907) Oligochaeten von Madagaskar, den Comoren und anderen Inseln des westlichen Indischen Ozeans. In: Voeltzkow, A. Reise in Ostafrika in den Jahren 1903–05, Wissenschaftliche Ergebnisse, 2, 41–50.

    Michaelsen, W. (1910) Oligochäten von verschiedenen Gebieten. Mitteilungen aus dem Naturhistorischen Museum in Hamburg, 27 (2), 47–169.

    Michaelsen, W. (1911) Zur Kenntnis der Eodrilaceen un ihrer Verbreitungsverhältnisse. Zoologischen Jahrbücher Abteiling für Systematik, Geographie und Biologie der Tiere, 30, 527–572.

    Michaelsen, W. (1921) Neue und wenig bekannte Oligochäten aus skandinavischen Sammlungen. Arkiv för Zoologi, 13 (19), 1–25.
    http://dx.doi.org/10.5962/bhl.part.20150

    Michaelsen, W. (1913) Die Oligochäten von Neu-Caledonien und den benachbarten Inselgruppen. In: Sarasin, F. & Roux, J. (Eds.), Nova Caledonia, Zoologie, 1 (5), pp. 173–280.

    Michaelsen, W. (1923) Oligochäten von den wärmeren Gebieten Amerikas und des Atlantischen Ozeans sowie ihre faunistischen Beziehungen. Mitteilungen aus dem Naturhistorischen Museum in Hamburg, 41,71–83.

    Moore, J.P. (1904) Description of a new species of earthworm (Diplocardia longa) from Georgia. Proceedings of the Academy of Natural Sciences of Philadelphia, 56, 803–808.

    Murchie, W.R. (1961) A new diplocardian earthworm from Illinois (Oligochaeta: Megascolecidae). The Ohio Journal of Science, 61 (6), 367–371.

    Murchie, W.R. (1963) Description of a new diplocardian earthworm, Diplocardia longiseta (Oligochaeta: Megascolecidae). The Ohio Journal of Science, 63 (1), 15–18.

    Murchie, W.R. (1965) Diplocardia gatesi, a new earthworm from North Carolina (Oligochaeta: Megascolecidae). The Ohio Journal of Science, 65 (4), 208–211.

    Murchie, W.R. (1967) Description of Diplocardia macdowelli a new Megascolecid earthworm from Mississippi (Oligochaeta). The Ohio Journal of Science, 67 (1), 50–53.

    Omodeo, P. (2000) Evolution and biogeography of megadriles (Annelida, Clitellata). Italian Journal of Zoology, 67 (2), 179–201.
    http://dx.doi.org/10.1080/11250000009356313

    Pickford, G.E. (1932) Oligochaeta. Part II. Discovery Reports, 4, 265–290.
    http://dx.doi.org/10.5962/bhl.part.17060

    Pickford, G.E. (1937) A monograph of the Acanthodriline earthworms of South Africa. W. Heffer & Sons Ltd., Cambridge, 612 pp.

    Pickford, G.E. (1938) Earthworms in Yucatan Caves. In: Pearse, A.S. (Ed.), Fauna of the Caves of Yucatan. Carnegie Institution of Washington Publication, No. 49, pp. 71–100.

    Plisko, J.D. (2004) Review of the balantine genus Udeina Michaelsen, 1910 with descriptions of six new species in South Africa (Oligochaeta: Acanthodrilidae, Acanthodrilinae). African Invertebrates, 45, 287–313.

    Plisko, J.D. (2007) New species of South African acanthodriline earthworms of the genera Eodriloides and Chilota, with a redescription of Chilota quindecimus (Oligochaeta: Acanthodrilidae). African Invertebrates, 48 (2), 33–40.

    Plisko, J.D. (2008) New South African Acanthodrilinae earthworm species, with new data for some earlier known members of the genus Parachilota (Oligochaeta: Acanthodrilidae). African Invertebrates, 49 (2), 21–36.
    http://dx.doi.org/10.5733/afin.049.0203

    Plisko, J.D. (2012) An annotated checklist of the South African Acanthodrilidae (Oligochaeta: Acanthodrilidae: Acanthodrilinae, Benhamiinae). Zootaxa, 3458, 4–58.

    Redfern, R. (2000) Origins. The evolution of continents, oceans and life. Cassell & Company, London, 360 pp.

    Reynolds, J.W. & Righi, G. (1994) On some earthworms from Belize, C.A. with the description of a new species (Oligochaeta: Acanthodrilidae, Glossoscolecidae and Octochaetidae). Megadrilogica, 5 (9), 97–106.

    Reynolds, J.W. & Wetzel, M.J. (2015) Nomenclatura Oligochaetologica – A catalogue of names, descriptions and type specimens. Editio Secunda. Available from: http://wwx.inhs.illinois.edu/people/mjwetzel/nomenoligo (accessed 31 November 2015)

    Righi, G. (1972) On some earthworms from Central America (Oligochaeta). S,tudies on the Neotropical Fauna 7, 207–228.
    http://dx.doi.org/10.1080/01650527209360445

    Rodríguez, C. & Fragoso, C. (1995) Especies nuevas de Zapatadrilus James, 1991 (Oligochaeta: Megascolecidae) registradas en Cuba. Acta Zoologica Mexicana, New Series, 64, 21–33.

    Rodríguez, C. & Fragoso, C. (2002) Filogenia y biogeografía de Cubadrilus (Oligochaeta: Octochaetidae), un género nuevo de lombriz de tierra de Cuba. Acta Zoologica Mexicana, New Series, 87, 125–146.

    Rosa, D. (1901) Gli Oligocheti raccolti in Patagonia. Atti della Societá dei Naturalisti e Matematici di Modena, 4 (35), 7–10.

    Sherlock, E. & Csuzdi, C. (2013) Two new earthworm species from Belize (Oligochaeta: Acanthodrilidae). Journal of Natural History, 47 (29–30), 1911–1919.
    http://dx.doi.org/10.1080/00222933.2013.770931

    Szlavecz, K., Chang, C.-H. Burgess, J.L. & Csuzdi, C. (2014) Earthworms (Annelida: Clitellata) of Plummers Island, Maryland, U.S.A., with description of a new species. Proceedings of the Biological Society of Washington, 126 (4), 312–322.
    http://dx.doi.org/10.2988/0006-324X-126.4.312

    Smith, F. (1895) A preliminary account fo two new Oligochaeta from Illinois. Bulletin of the Illinois State Laboratory of Natural History, 4, 138–148.

    Smith, F. (1924) A new earthworm from Texas belonging to the genus Diplocardia. Proceedings of the United States National Museum, 66, 1–6.
    http://dx.doi.org/10.5479/si.00963801.2549

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

    Zicsi, A. (1989) Revision der Gattung Yagansia Michaelsen, 1899 (Oligochaeta, Acanthodrilidae). Regenwürmer aus Südamerika 11. Acta Zoologica Hungarica, 35 (3–4), 413–430.

    Zicsi, A. (1993a) Revision der Gattung Chilota Michaelsen sowie weitere neue Angaben zur Regenwurmfauna Chiles (Oligochaeta: Acanathodrilidae, Ocnerodrilidae). Regenwürmer aus Südamerika 20. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 90, 151–173.

    Zicsi, A. (1993b) Neue und bekannter Regenwürmer aus Chile (Oligochaeta). Regenwürmer aus Südamerika 19. Revue Suisse de Zoologie, 100 (3), 627–640.
    http://dx.doi.org/10.5962/bhl.part.79877

    Zicsi, A. (1998) Neue und seltene terrestrische Oligochaeten aus Südafrika. Mitteilungen aus dem Naturhistorischen Museum in Hamburg, 95, 59–77.

    Zicsi, A. & Pajor, I. (1992) Über die Regenwürmer (Oligochaeta) in den Natal-Drakensbergen Südafrikas. Opuscula Zoologica Budapest, 25, 125–136.

    Zicsi, A. & Csuzdi, C. (2001) Weitere Angaben zur Regenwurmfauna Chiles (Oligochaeta: Acanthodrilidae, Lumbricidae) Regenwürmer aus Südamerika 33. Berichte des Naturwissenschaftlich-Medizinischen Vereins in Innsbruck, 88, 129–140.