Enchytraeidae

Fridericia ratzeli, copyright Christer Erseus.

Belongs within: Tubificida.

The Enchytraeidae are a group of small, mostly pale-coloured oligochaetes, including both aquatic and terrestrial species.

Characters (from Dash 1990): Length <10–50 mm; colour usually pale white or grey. Setae usually simple; four bundles usually present (two laterally/dorsolaterally, two ventrolaterally) per segment. Clitellum usually covering segments XII–XIII. Gut with three pairs of primary septal glands present at segments IV–VII. One pair of nephridia usually present per segment from VII. Male reproductive system with paired seminal vesicles present in segment X; pair of sperm funnels present in XI, usually 2–3 times longer than wide; vas deferens opening throught penial bulb in clitellar segments; pair of spermathecae often present in IV (absent in parthenogenetic forms). Pair of ovaries present.

<==Enchytraeidae [Enchytraeina, Prosotheca]
    |--Aspidodrilus Baylis 1914D90
    |--ChirodrilusD90
    |--Pelmatodrilus Moore 1943D90
    |--Sinenchytraeus Liang & Hus 1979D90
    |--Enchylea Nielsen & christensen 1963D90
    |--Tupidrilus Righi 1974D90
    |    |--*T. wilsoni (Righi 1973) [=Hemienchytraeus wilsoni]R74a
    |    |--T. gei Righi 1974R74a
    |    |--T. lacteus Righi 1974R74a
    |    `--T. marcusae Righi 1974R74a
    |--Barbidrilus Loden & Locy 1981D90
    |--Bryohenlea Cernosvitov 1934D90
    |--Punahenlea Nurminen 1980D90
    |--Enchytronia Nielsen & Christensen 1959JD86
    |    `--E. hellenica Dumnicka 1979JD86
    |--Hemifridericia Nielsen & Christensen 1959D90
    |    `--H. khallikotosusD90
    |--Buchholzia Michaelsen 1887D90
    |    |--B. appendiculata (Buchholz 1862)D90
    |    `--B. fallaxJD12
    |--Stercutus Michaelsen 1888D90
    |    `--S. niveusD90
    |--Randidrilus Coates & Erseus 1985D90
    |    |--R. codensis (Lasserre 1971)D90
    |    `--R. quadrithecatus Coates & Erseus 1985D90
    |--Achaeta Vejdovsky 1877D90
    |    |--A. camerani (Cognetti 1889)D90
    |    |--A. eiseni (Vejdovsky 1877)D90
    |    |--A. neotropica Cernosvitov 1937R74a
    |    `--A. silvatica Nurminen 1973D90
    |--Cognettia Nielsen & Christensen 1959D90
    |    |--C. cognettiD90
    |    |--C. glandulosa (Michaelsen 1888)D90
    |    `--C. sphagnetorumD90
    |--Hemienchytraeus Cernosvitov 1935D90
    |    |--H. (Hemienchytraeus)R74a
    |    |    |--*H. (H.) stephensoni (Cognetti 1927) [=Enchytraeus stephensoni]R74a
    |    |    |--H. (H.) africanus Cernosvitov 1935R74a
    |    |    |--H. (H.) bifurcatus Nielsen & Christensen 1959R74a
    |    |    |--H. (H.) guineanus Omodeo 1958R74a
    |    |    |--H. (H.) inversus Omodeo 1958R74a
    |    |    |--H. (H.) maurilliae Righi 1981R81
    |    |    `--H. (H.) shirensis Bell 1954R74a
    |    `--H. (Cotienchytraeus Righi 1974)R74a
    |         |--H. (*C.) cipoensis Righi 1973R74a
    |         |--H. (C.) brasiliensis (Cognetti 1900) [=Mesenchytraeus brasiliensis]R74a
    |         `--H. (C.) rixae Righi 1974R74a
    |--Stephensoniella Cernosvitov 1934D90
    |    |--S. marina (Moore 1902)D90
    |    |--S. sterreri (Erseus & Lasserre 1976) [=Lumbricillus sterreri]D90
    |    `--S. trevori (Coates 1980) [=Marionina trevori]D90
    |--Guaranidrilus Cernosvitov 1937D90
    |    |--‘Henlea’ columbiana Michaelsen 1913R73
    |    |--G. glandulosus Cernosvitov 1937R73
    |    |--G. lamottei Omodeo 1958R73
    |    |--G. oiepe Righi 1974R74a
    |    |--G. rarus Cernosvitov 1937R73
    |    `--G. sawayai Righi 1973R73
    |--Bryodrilus Ude 1892D90
    |    |--B. arctica (Bell 1962)D90
    |    |--B. ehlersi Ude 1892D90
    |    |--B. novaescotiac (Bell 1962)D90
    |    `--B. parvus Nurminen 1970D90
    |         |--B. p. parvusD90
    |         `--B. p. kananaskis Dash 1970D90
    |--Cernosvitoviella Nielsen & Christensen 1959P86
    |    |--C. aggtelekiensis Dozsa-Farkas 1970JD86
    |    |--C. atrata (Bretscher 1903)D90
    |    |--C. celere Nurminen 1973D90
    |    |--C. christenseni Dash 1970D90
    |    |--C. immota (Knöllner 1935)P86
    |    |--C. parviseta Gadzinka 1974JD86
    |    `--C. pusilla Nurminen 1973D90
    |--Grania Southern 1913 [incl. Hemigrania]D90
    |    |--G. americana Kennedy 1966D90
    |    |--G. atlantica Coates & Erseus 1985D90
    |    |--G. incerta Coates & Erseus 1980D90
    |    |--G. levis Coates & Erseus 1985D90
    |    |--G. macrochaeta (Pierantoni 1901)P86
    |    |--G. maricola Southern 1913P86
    |    |--G. paucispina Eisen 1904D90
    |    |--G. postclitellochaeta (Knöllner 1935) [=Marionina postclitellochaeta, Hemigrania postclitellochaeta]D90
    |    |--G. reducta Coates & Erseus 1985D90
    |    `--G. variochaetaP86
    |--Enchytraeus Henle 1837JD86
    |    |--*E. albidus Henle 1837H62 [incl. E. mobiiJ31]
    |    |--E. alaskae Eisen 1904D90
    |    |--E. berhampurosusD90
    |    |--E. buchholzi Vejdovsky 1879D90
    |    |--E. capitatus Bulow 1957D90
    |    |--E. cryptosetosus Tynen 1969D90
    |    |--E. dominicae Dumnicka 1976JD86
    |    |--E. fragmentosus Bell 1959D90
    |    |--E. gillettensis Welch 1914D90
    |    |--E. kincaidi Eisen 1904D90
    |    |--E. minutus Nielsen & Christensen 1961D90
    |    |--E. nodosusJ31
    |    `--E. sepultus Menge 1866W93
    |--Henlea Michaelsen 1889D90
    |    |--H. californica Eisen 1904D90
    |    |    |--H. c. californicaD90
    |    |    `--H. c. monticola Eisen 1904D90
    |    |--H. eiseni Bell 1942D90
    |    |--H. erhorni Eisen 1904D90
    |    |--H. glabra Altman 1936D90
    |    |--H. helenae Cernosvitov 1937D90
    |    |--H. heleotropha Stephenson 1922D90
    |    |--H. leptodera (Vejdovsky 1879)R74b
    |    |--H. moderatoidea Altman 1936D90
    |    |--H. nasuta Eisen 1878D90
    |    |--H. ochracea (Eisen 1878) (see below for synonymy)D90
    |    |--H. parva Friend 1912D90
    |    |--H. perpusilla Friend 1911D90
    |    |--H. scharffi Southern 1910D90
    |    |--H. udei (Eisen 1904) [=Bryohenlea udei]D90
    |    |--H. urbanensis Welch 1914D90
    |    |--H. ventriculosa Udekem 1854D90
    |    `--H. welchi Bell 1942D90
    |--Lumbricillus Orsted 1844D90
    |    |--L. annulatus Eisen 1904D90
    |    |--L. arenarius (Michaelsen 1889)D90
    |    |--L. belli Tynen 1969D90
    |    |--L. charae (Tynen 1970)D90
    |    |--L. codensis Lasserre 1971D90
    |    |--L. curtus Coastes 1981D90
    |    |--L. franciscanus Eisen 1904D90
    |    |    |--L. f. franciscanusD90
    |    |    |--L. f. borealis Eisen 1904D90
    |    |    `--L. f. unalaskae Eisen 1904D90
    |    |--L. georgiensis Tynen 1969D90
    |    |--L. kalatdlitus Nurminen 1970D90
    |    |--L. lineatus (Müller 1774) [incl. L. agilis Moore 1905]D90
    |    |--L. merriami Eisen 1904D90
    |    |    |--L. m. merriamiD90
    |    |    `--L. m. elongatus Eisen 1904D90
    |    |--L. mirabilis Tynen 1969D90
    |    |--L. pagenstecheri (Ratzel 1869)D90
    |    |--L. qualicumensis Tynen 1969D90
    |    |--L. ritteri Eisen 1904D90
    |    |--L. rivalisD90
    |    |--L. rupertensis Coates 1981D90
    |    |--L. rutilus Welch 1914D90
    |    |--L. santaeclarae Eisen 1904D90
    |    |--L. semifuscus (Claparede 1861)R74b
    |    |--L. tsimpseanis Coates 1981D90
    |    |--L. tuba Stephenson 1911D90
    |    `--L. vancouverensis Tynen 1969D90
    |--Marionina Michaelsen 1889 [incl. Michaelsena]D90
    |    |--M. achaeta Lasserre 1964P86
    |    |--M. alaskae Eisen 1904D90
    |    |--M. americana Eisen 1904 [=Lumbicillus americanus]D90
    |    |--M. appendiculata Nielsen & Christensen 1959P86
    |    |--M. argentea (Michaelsen 1889)D90
    |    |--M. canadensis Dash 1970D90
    |    |--M. charlottensis Coates 1980D90
    |    |--M. citrina (Eisen 1904)D90
    |    |--M. craggi Dash 1970D90
    |    |--M. diazi Coates & Erseus 1985D90
    |    |--M. dirksi Bell 1942D90
    |    |--M. ecuadoriensis Righi 1981R81
    |    |--M. elgonensis (Cernosvitov 1938R81
    |    |--M. elongataP86
    |    |--M. forbesae Smith & Welch 1913D90
    |    |--M. istriae Giere 1974R81
    |    |--M. klaskisharum Coates 1983D90
    |    |--‘Michaelsena’ macrochaeta Pierantoni 1903M07
    |    |--M. mesopsamma Lassere 1964R81
    |    |--M. modesta Eisen 1904D90
    |    |--‘Michaelsena’ monochaetaM07
    |    |--M. neroutsensis Coates 1980D90
    |    |--M. nevisensis (Right & Kanner 1979)D90
    |    |--‘Michaelsena’ normani Michaelsen 1907M07
    |    |--M. patua Righi et al. 1978R81
    |    |--M. paucispina (Eisen 1904)D90 [=Michaelsena paucispinaM07]
    |    |--M. preclitellochaeta Nielsen & Christensen 1963P86
    |    |--‘Michaelsena’ principissae Michaelsen 1907M07
    |    |--M. pugetensis (Altman 1931)D90
    |    |--M. saxicola (Eisen 1904) [=Enchytraeus saxicola]D90
    |    |--M. sjaelandica Nielsen & Christensen 1959D90
    |    |--M. southerni (Cernosvitov 1937)P86
    |    |--M. spicula (Leuckart 1847)P86
    |    |--M. sublitoralisJD12
    |    |--M. subterranea (Knollner 1935)D90
    |    |--*Michaelsena’ subtilis Ude 1896M07
    |    |--M. ticha Righi 1980R81
    |    |--‘Enchytraeoides’ unisetosus [=Michaelsena unisetosa]M07
    |    |--M. vancouverensis Coates 1980D90
    |    |--M. weilliP86
    |    `--M. welchi Lasserre 1971D90
    |--Mesenchytraeus Eisen 1878D90
    |    |--M. altus Welch 1917D90
    |    |--M. americanus Bell 1942D90
    |    |--M. arcticus Bell 1942D90
    |    |--M. argentatus Nurminen 1973D90
    |    |--M. armatus (Levinsen 1884)D90
    |    |    |--M. a. armatusD90
    |    |    `--M. a. kananaskis Dash 1970D90
    |    |--M. atriophorus Altman 1936D90
    |    |--M. beringensis Eisen 1904D90
    |    |--M. chromophorus Altman 1936D90
    |    |--M. diplobulbosus Bell 1949D90
    |    |--M. eastwoodi (Eisen 1904)D90
    |    |--M. flavidus Michaelsen 1887R74b
    |    |--M. flavus (Levinsen 1884)R74b
    |    |--M. fontinalis Eisen 1904D90
    |    |--M. franciscanus Eisen 1904D90
    |    |--M. fuscus Eisen 1904D90
    |    |    |--M. f. fuscusD90
    |    |    `--M. f. inermis Eisen 1904D90
    |    |--M. gelidus Welch 1916D90
    |    |--M. glandulosusFFB88
    |    |--M. grandis Eisen 1904D90
    |    |--M. harrimani Eisen 1904D90
    |    |--M. hydrius Welch 1919D90
    |    |--M. johanseni Welch 1919D90
    |    |--M. kincaidi Eisen 1904D90
    |    |--M. macnabi Bell 1942D90
    |    |--M. maculatus Eisen 1904D90
    |    |--M. magnus Altman 1936D90
    |    |--M. minimus Altman 1936D90
    |    |--M. monothecatus Bell 1945D90
    |    |--M. nanus Eisen 1904D90
    |    |--M. obscurus Eisen 1904D90
    |    |--M. orcae Eisen 1904D90
    |    |--M. pedatus Eisen 1904D90
    |    |--M. penicillus Eisen 1904D90
    |    |--M. setchelli Eisen 1904D90
    |    |--M. solifugus (Emery 1898)D90
    |    |    |--M. s. solifugusD90
    |    |    `--M. s. rainierensis Welch 1916D90
    |    `--M. unalaskae Eisen 1904D90
    `--Fridericia Michaelsen 1889 [incl. Distichopus Leidy 1882]D90
         |--F. agaricola Moore 1895D90
         |--F. agilis Smith 1895D90
         |--F. alba Moore 1895D90
         |--F. anglica Friend 1912D90
         |--F. berklyensis Bell 1936D90
         |--F. bisetosaM07
         |--F. bulboides Nielsen & Christensen 1959D90
         |--F. bulbosa Rosa 1887D90
         |--F. californica Eisen 1904D90
         |--F. canadensis Dash 1972D90
         |--F. caprensis Bell 1947D90
         |--F. firma Smith & Welch 1913D90
         |--F. fuchsi Eisen 1904D90
         |--F. galba (Hoffmeister 1845)D90
         |--F. harrimani Eisen 1904D90
         |--F. hegemon (Vejdovsky 1877)R74b
         |--F. johnsoni Eisen 1904D90
         |--F. kalffi Nurminen 1973D90
         |--F. kalingaD90
         |--F. leydigi (Vejdovsky 1877)R74b
         |--F. longa Moore 1895D90
         |--F. losangelensis Bell 1936D90
         |--F. macgregori Eisen 1904D90
         |--F. magnaJD86
         |    |--F. m. magnaJD86
         |    `--F. m. carpathica Botea 1973JD86
         |--F. nielseni Dash 1972D90
         |--F. oconeensis Welch 1914D90
         |--F. papofiana Eisen 1904D90
         |--F. paroniana Issel 1904D90
         |--F. parva Moore 1895D90
         |--F. perrieri (Vejdovsky 1877)R74b
         |--F. polychaeta Bretscher 1900R74b
         |--F. ratzeli (Eisen 1872)D90
         |--F. sacculata Bell 1936D90
         |--F. santaebarbarae Eisen 1904D90
         |--F. sima Welch 1914D90
         |--F. sonorae Eisen 1904D90
         |--F. sylvestris (Leidy 1882)D90 [=Distichopus silvestrisM07]
         |--F. syraccusa Dash & Mitchell 1981D90
         |--F. tenera Smith & Welch 1913D90
         |--F. tuberosa Rota 1995RP07
         `--F. whatcomae Altman 1936D90
Nomen nudum: Guaranidrilus fridericoides Cernosvitov 1937R73

Henlea ochracea (Eisen 1878) [incl. H. arctica Welch 1912, H. moderata Welch 1914, H. tenella Eisen 1878, H. tubulifera Welch 1914]D90

*Type species of generic name indicated

References

[D90] Dash, M. C. 1990. Oligochaeta: Enchytraeidae. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 311–340. John Wiley & Sones: New York.

[FFB88] Franke, U., B. Friebe & L. Beck. 1988. Methodisches zur Ermittlung der Siedlungsdichte von Bodentieren aus Quadratproben und Barberfallen. Pedobiologia 32: 253–264.

[H62] Howell, B. F. 1962. Worms. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt W. Miscellanea: Conodonts, Conoidal Shells of Uncertain Affinities, Worms, Trace Fossils and Problematica pp. W144–W177. Geological Society of America, and University of Kansas Press.

[J31] Jackson, A. 1931. The Oligochaeta of south-western Australia. Journal of the Royal Society of Western Australia 17: 71–136.

[JD12] James, S. W., & S. K. Davidson. 2012. Molecular phylogeny of earthworms (Annelida: Crassiclitellata) based on 28S, 18S and 16S gene sequences. Invertebrate Systematics 26: 213–229.

[JD86] Juget, J., & E. Dumnicka. 1986. Oligochaeta (incl. Aphanoneura) des eaux souterraines continentales. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 234–244. E. J. Brill/Dr W. Backhuys: Leiden.

[M07] Michaelsen, W. 1907. Oligochaeta. Die Fauna Südwest-Australiens 1 (2): 115–232, 2 pls.

[P86] Pfannkuche, O. 1986. Oligochaeta from marine and brackish-water subterranean habitats. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 245–249. E. J. Brill/Dr W. Backhuys: Leiden.

[R73] Righi, G. 1973. Sobre três espécies brasileiras de Enchytraeidae (Oligochaeta). Bol. Zool. e Biol. Mar., N. S. 30: 469–482.

[R74a] Righi, G. 1974a. Notas sobre os Oligochaeta, Enchytraeidae do Brasil. Papéis Avulsos de Zoologia 28 (7): 127–145.

[R74b] Righi, G. 1974b. Alguns Sporozoa (Gregarinida e Haplosporida) de Oligochaeta (Naididae e Enchytraeidae) brasileiros. Papéis Avulsos de Zoologia 28 (11): 185–195.

[R81] Righi, G. 1981. Alguns Oligochaeta cavernícolas do Equador. Papéis Avulsos de Zoologia 34 (22): 235–249.

[RP07] Rousset, V., F. Pleijel, G. W. Rouse, C. Erséus & M. E. Siddall. 2007. A molecular phylogeny of annelids. Cladistics 23: 41–63.

[W93] Wills, M. A. 1993. Annelida. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 271–278. Chapman & Hall: London.

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