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 1914 D90
    |--Chirodrilus D90
    |--Pelmatodrilus Moore 1943 D90
    |--Sinenchytraeus Liang & Hus 1979 D90
    |--Enchylea Nielsen & christensen 1963 D90
    |--Tupidrilus Righi 1974 D90
    |--Barbidrilus Loden & Locy 1981 D90
    |--Bryohenlea Cernosvitov 1934 D90
    |--Punahenlea Nurminen 1980 D90
    |--Enchytronia Nielsen & Christensen 1959 JD86
    |    `--E. hellenica Dumnicka 1979 JD86
    |--Hemifridericia Nielsen & Christensen 1959 D90
    |    `--H. khallikotosus D90
    |--Buchholzia Michaelsen 1887 D90
    |    `--B. appendiculata (Buchholz 1862) D90
    |--Stercutus Michaelsen 1888 D90
    |    `--S. niveus D90
    |--Randidrilus Coates & Erseus 1985 D90
    |    |--R. codensis (Lasserre 1971) D90
    |    `--R. quadrithecatus Coates & Erseus 1985 D90
    |--Achaeta Vejdovsky 1877 D90
    |    |--A. camerani (Cognetti 1889) D90
    |    |--A. eiseni (Vejdovsky 1877) D90
    |    `--A. silvatica Nurminen 1973 D90
    |--Cognettia Nielsen & Christensen 1959 D90
    |    |--C. cognetti D90
    |    |--C. glandulosa (Michaelsen 1888) D90
    |    `--C. sphagnetorum D90
    |--Hemienchytraeus Cernosvitov 1935 D90
    |    |--H. brasiliensis (Cognetti 1900) R73
    |    |--H. cipoensis Righi 1973 R73
    |    `--H. wilsoni Righi 1973 R73
    |--Stephensoniella Cernosvitov 1934 D90
    |    |--S. marina (Moore 1902) D90
    |    |--S. sterreri (Erseus & Lasserre 1976) [=Lumbricillus sterreri] D90
    |    `--S. trevori (Coates 1980) [=Marionina trevori] D90
    |--Guaranidrilus Cernosvitov 1937 D90
    |    |--‘Henlea’ columbiana Michaelsen 1913 R73
    |    |--G. glandulosus Cernosvitov 1937 R73
    |    |--G. lamottei Omodeo 1958 R73
    |    |--G. rarus Cernosvitov 1937 R73
    |    `--G. sawayai Righi 1973 R73
    |--Bryodrilus Ude 1892 D90
    |    |--B. arctica (Bell 1962) D90
    |    |--B. ehlersi Ude 1892 D90
    |    |--B. novaescotiac (Bell 1962) D90
    |    `--B. parvus Nurminen 1970 D90
    |         |--B. p. parvus D90
    |         `--B. p. kananaskis Dash 1970 D90
    |--Cernosvitoviella Nielsen & Christensen 1959 P86
    |    |--C. aggtelekiensis Dozsa-Farkas 1970 JD86
    |    |--C. atrata (Bretscher 1903) D90
    |    |--C. celere Nurminen 1973 D90
    |    |--C. christenseni Dash 1970 D90
    |    |--C. immota (Knöllner 1935) P86
    |    |--C. parviseta Gadzinka 1974 JD86
    |    `--C. pusilla Nurminen 1973 D90
    |--Grania Southern 1913 [incl. Hemigrania] D90
    |    |--G. americana Kennedy 1966 D90
    |    |--G. atlantica Coates & Erseus 1985 D90
    |    |--G. incerta Coates & Erseus 1980 D90
    |    |--G. levis Coates & Erseus 1985 D90
    |    |--G. macrochaeta (Pierantoni 1901) P86
    |    |--G. maricola Southern 1913 P86
    |    |--G. paucispina Eisen 1904 D90
    |    |--G. postclitellochaeta (Knöllner 1935) [=Marionina postclitellochaeta, Hemigrania postclitellochaeta] D90
    |    |--G. reducta Coates & Erseus 1985 D90
    |    `--G. variochaeta P86
    |--Enchytraeus Henle 1837 JD86
    |    |--*E. albidus Henle 1837 H62 [incl. E. mobii J31]
    |    |--E. alaskae Eisen 1904 D90
    |    |--E. berhampurosus D90
    |    |--E. buchholzi Vejdovsky 1879 D90
    |    |--E. capitatus Bulow 1957 D90
    |    |--E. cryptosetosus Tynen 1969 D90
    |    |--E. dominicae Dumnicka 1976 JD86
    |    |--E. fragmentosus Bell 1959 D90
    |    |--E. gillettensis Welch 1914 D90
    |    |--E. kincaidi Eisen 1904 D90
    |    |--E. minutus Nielsen & Christensen 1961 D90
    |    `--E. nodosus J31
    |--Henlea Michaelsen 1889 D90
    |    |--H. californica Eisen 1904 D90
    |    |    |--H. c. californica D90
    |    |    `--H. c. monticola Eisen 1904 D90
    |    |--H. eiseni Bell 1942 D90
    |    |--H. erhorni Eisen 1904 D90
    |    |--H. glabra Altman 1936 D90
    |    |--H. helenae Cernosvitov 1937 D90
    |    |--H. heleotropha Stephenson 1922 D90
    |    |--H. moderatoidea Altman 1936 D90
    |    |--H. nasuta Eisen 1878 D90
    |    |--H. ochracea (Eisen 1878) (see below for synonymy) D90
    |    |--H. parva Friend 1912 D90
    |    |--H. perpusilla Friend 1911 D90
    |    |--H. scharffi Southern 1910 D90
    |    |--H. udei (Eisen 1904) [=Bryohenlea udei] D90
    |    |--H. urbanensis Welch 1914 D90
    |    |--H. ventriculosa Udekem 1854 D90
    |    `--H. welchi Bell 1942 D90
    |--Lumbricillus Orsted 1844 D90
    |    |--L. annulatus Eisen 1904 D90
    |    |--L. arenarius (Michaelsen 1889) D90
    |    |--L. belli Tynen 1969 D90
    |    |--L. charae (Tynen 1970) D90
    |    |--L. codensis Lasserre 1971 D90
    |    |--L. curtus Coastes 1981 D90
    |    |--L. franciscanus Eisen 1904 D90
    |    |    |--L. f. franciscanus D90
    |    |    |--L. f. borealis Eisen 1904 D90
    |    |    `--L. f. unalaskae Eisen 1904 D90
    |    |--L. georgiensis Tynen 1969 D90
    |    |--L. kalatdlitus Nurminen 1970 D90
    |    |--L. lineatus (Müller 1774) [incl. L. agilis Moore 1905] D90
    |    |--L. merriami Eisen 1904 D90
    |    |    |--L. m. merriami D90
    |    |    `--L. m. elongatus Eisen 1904 D90
    |    |--L. mirabilis Tynen 1969 D90
    |    |--L. pagenstecheri (Ratzel 1869) D90
    |    |--L. qualicumensis Tynen 1969 D90
    |    |--L. ritteri Eisen 1904 D90
    |    |--L. rivalis D90
    |    |--L. rupertensis Coates 1981 D90
    |    |--L. rutilus Welch 1914 D90
    |    |--L. santaeclarae Eisen 1904 D90
    |    |--L. tsimpseanis Coates 1981 D90
    |    |--L. tuba Stephenson 1911 D90
    |    `--L. vancouverensis Tynen 1969 D90
    |--Marionina Michaelsen 1889 [incl. Michaelsena] D90
    |    |--M. achaeta Lasserre 1964 P86
    |    |--M. alaskae Eisen 1904 D90
    |    |--M. americana Eisen 1904 [=Lumbicillus americanus] D90
    |    |--M. appendiculata Nielsen & Christensen 1959 P86
    |    |--M. argentea (Michaelsen 1889) D90
    |    |--M. canadensis Dash 1970 D90
    |    |--M. charlottensis Coates 1980 D90
    |    |--M. citrina (Eisen 1904) D90
    |    |--M. craggi Dash 1970 D90
    |    |--M. diazi Coates & Erseus 1985 D90
    |    |--M. dirksi Bell 1942 D90
    |    |--M. elongata P86
    |    |--M. forbesae Smith & Welch 1913 D90
    |    |--M. klaskisharum Coates 1983 D90
    |    |--M. mesopsamma P86
    |    |--M. modesta Eisen 1904 D90
    |    |--M. neroutsensis Coates 1980 D90
    |    |--M. nevisensis (Right & Kanner 1979) D90
    |    |--‘Michaelsena’ normani J31
    |    |--M. paucispina (Eisen 1904) D90
    |    |--M. preclitellochaeta Nielsen & Christensen 1963 P86
    |    |--‘Michaelsena’ principissae J31
    |    |--M. pugetensis (Altman 1931) D90
    |    |--M. saxicola (Eisen 1904) [=Enchytraeus saxicola] D90
    |    |--M. sjaelandica Nielsen & Christensen 1959 D90
    |    |--M. southerni (Cernosvitov 1937) P86
    |    |--M. spicula (Leuckart 1847) P86
    |    |--M. subterranea (Knollner 1935) D90
    |    |--M. vancouverensis Coates 1980 D90
    |    |--M. weilli P86
    |    `--M. welchi Lasserre 1971 D90
    |--Mesenchytraeus Eisen 1878 D90
    |    |--M. altus Welch 1917 D90
    |    |--M. americanus Bell 1942 D90
    |    |--M. arcticus Bell 1942 D90
    |    |--M. argentatus Nurminen 1973 D90
    |    |--M. armatus (Levinsen 1884) D90
    |    |    |--M. a. armatus D90
    |    |    `--M. a. kananaskis Dash 1970 D90
    |    |--M. atriophorus Altman 1936 D90
    |    |--M. beringensis Eisen 1904 D90
    |    |--M. chromophorus Altman 1936 D90
    |    |--M. diplobulbosus Bell 1949 D90
    |    |--M. eastwoodi (Eisen 1904) D90
    |    |--M. fontinalis Eisen 1904 D90
    |    |--M. franciscanus Eisen 1904 D90
    |    |--M. fuscus Eisen 1904 D90
    |    |    |--M. f. fuscus D90
    |    |    `--M. f. inermis Eisen 1904
    |    |--M. gelidus Welch 1916 D90
    |    |--M. glandulosus FFB88
    |    |--M. grandis Eisen 1904 D90
    |    |--M. harrimani Eisen 1904 D90
    |    |--M. hydrius Welch 1919 D90
    |    |--M. johanseni Welch 1919 D90
    |    |--M. kincaidi Eisen 1904 D90
    |    |--M. macnabi Bell 1942 D90
    |    |--M. maculatus Eisen 1904 D90
    |    |--M. magnus Altman 1936 D90
    |    |--M. minimus Altman 1936 D90
    |    |--M. monothecatus Bell 1945 D90
    |    |--M. nanus Eisen 1904 D90
    |    |--M. obscurus Eisen 1904 D90
    |    |--M. orcae Eisen 1904 D90
    |    |--M. pedatus Eisen 1904 D90
    |    |--M. penicillus Eisen 1904 D90
    |    |--M. setchelli Eisen 1904 D90
    |    |--M. solifugus (Emery 1898) D90
    |    |    |--M. s. solifugus D90
    |    |    `--M. s. rainierensis Welch 1916 D90
    |    `--M. unalaskae Eisen 1904 D90
    `--Fridericia Michaelsen 1889 [incl. Distichopus Leidy 1882] D90
         |--F. agaricola Moore 1895 D90
         |--F. agilis Smith 1895 D90
         |--F. alba Moore 1895 D90
         |--F. anglica Friend 1912 D90
         |--F. berklyensis Bell 1936 D90
         |--F. bulboides Nielsen & Christensen 1959 D90
         |--F. bulbosa Rosa 1887 D90
         |--F. californica Eisen 1904 D90
         |--F. canadensis Dash 1972 D90
         |--F. caprensis Bell 1947 D90
         |--F. firma Smith & Welch 1913 D90
         |--F. fuchsi Eisen 1904 D90
         |--F. galba (Hoffmeister 1845) D90
         |--F. harrimani Eisen 1904 D90
         |--F. hegemon D90
         |--F. johnsoni Eisen 1904 D90
         |--F. kalffi Nurminen 1973 D90
         |--F. kalinga D90
         |--F. longa Moore 1895 D90
         |--F. losangelensis Bell 1936 D90
         |--F. macgregori Eisen 1904 D90
         |--F. magna JD86
         |    |--F. m. magna JD86
         |    `--F. m. carpathica Botea 1973 JD86
         |--F. nielseni Dash 1972 D90
         |--F. oconeensis Welch 1914 D90
         |--F. papofiana Eisen 1904 D90
         |--F. paroniana Issel 1904 D90
         |--F. parva Moore 1895 D90
         |--F. ratzeli (Eisen 1872) D90
         |--F. sacculata Bell 1936 D90
         |--F. santaebarbarae Eisen 1904 D90
         |--F. sima Welch 1914 D90
         |--F. sonorae Eisen 1904 D90
         |--F. sylvestris (Leidy 1882) D90
         |--F. syraccusa Dash & Mitchell 1981 D90
         |--F. tenera Smith & Welch 1913 D90
         |--F. tuberosa Rota 1995 RP07
         `--F. whatcomae Altman 1936 D90

Nomen nudum: Guaranidrilus fridericoides Cernosvitov 1937 R73

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.

[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.

[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.

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

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