Aquatic Annelids of Illinois | INHS Annelid Collection | Illinois Natural History Survey

Wetzel (1992) published the first comprehensive review of aquatic Annelida in Illinois, including 132 species representing 71 genera in 15 families. The list below has been modified to: a) include species identified from the state since publication of that review; b) reflect recent nomenclatural and systematic changes; and c) correct several actual date-of-publication errors that had been perpetuated in past literature. Recent works on the phylogeny of aquatic Annelida have been included in the classification section. References to publications noted in the text of this website are included in the ‘Literature Cited and Selected References’ section, which follows the checklist of species.

Phylogeny and Classification of Aquatic Annelida

The first global compendium on aquatic oligochaetes was published by Brinkhurst and Jamieson (1971). The section in that book discussing the phylogeny and classification of aquatic oligochaetes set the stage for numerous investigations into the phylogenetic relationships of all annelids. Most noteable are: Jamieson (1978, 1980, 1988), Timm (1981), Brinkhurst (1982, 1986, 1988, 1989, 1991a, 1991b, 1991c, 1992, 1994, 1999), Kasprzak (1982, 1984), Kathman and Brinkhurst (1998), Parker (1982), Holt (1986, 1989), Erséus (1987, 1990), Sawyer (1986a, 1986b, 1986c), Brinkhurst and Nemec (1987), Coates (1987, 1989), Nemec and Brinkhurst (1987), Brinkhurst and Gelder (1989), Gelder and Brinkhurst (1990), Purschke et al. (1993), Sima (1993), Sidall and Burreson (1995, 1996, 1998), Bartolomaeus (1997), Trapido-Rosenthal (1997), Omodeo (1998, 2000), Sidall et al. (1998), McHugh (1999, 2000), Purschke (1999), Westheide et al. (1999), Martin et al. (2000) [plus more recent publications to be added here soon]. This paper follows the division of the phylum Annelida into two classes: the Polychaeta and the Clitellata (including the Acanthobdellae, Branchiobdellae, Oligochaeta, and Hirudinida). Nomenclature follows Reynolds and Cook (1976, 1981, 1989, 1993) and Reynolds and Wetzel (2025), and Schmelz et al. (2021) for the oligochaetes; Klemm (1985) for Acanthobdellida and Polychaeta; Klemm (1985), Sawyer (1986a,1986b, 1986c), Barta and Sawyer (1990), and Jones and Woo (1990) for Hirudinida; and Holt (1986), Holt and Opell (1993), Gelder (1996), Gelder et al. (2002), and Govedich et al. (2010) for Branchiobdellida.

Checklist of Aquatic Annelida Occurring in Illinois

Species preceded by a single asterisk (*) were reported by Wetzel (1992) as new records for the state of Illinois. Species preceded by two asterisks (**) indicate new records for the state (published and unpublished) since 1992. Illinois species collected only from inshore Lake Michigan habitats are preceded by a ‘plus’ sign (+).

Phylum – Annelida

Class Polychaeta (Grube, 1850)
- Order Aeolosomatida
  - Family Aeolosomatidae
    - Genus Aeolosoma (Ehrenberg, 1828)
      - Aeolosoma hemprichi (Ehrenberg, 1828)
      - Aeolosoma tenebrarum (Vejdovský, 1880)
      - Aeolosoma variegatum (Vejdovský, 1884)
Class Clitellata
- Order Branchiobdellida
  - Family Bdellodrilidae (Holt, 1986)
    - Genus Bdellodrilus (Moore, 1895)
      - Bdellodrilus illuminatus (Moore, 1894)
  - Family Cambarincolidae (Holt, 1986)
    - Genus Cambarincola (Ellis, 1912)
      - Cambarincola chirocephalus (Ellis, 1919)
      - Cambarincola illinoisensis (Holt, 1982) **
      - Cambarincola macrodontus Ellis, 1912
      - Cambarincola philadelphicus (Leidy, 1851)
      - Cambarincola vitreus (Ellis, 1919)
    - Genus Oedipodrilus (Holt, 1967)
      - Oedipodrilus macbaini (Holt, 1955)
    - Genus Pterodrilus (Moore, 1895)
      - Pterodrilus distichus (Moore, 1895)
    - Genus Sathodrilus (Holt, 1968)
      - Sathodrilus elevatus (Goodnight, 1940)
  - Family Xironodrilidae (Holt, 1986)
    - Genus Xironodrilus (Ellis, 1919) (Note: This genus was listed under the family Branchiobdellidae in Wetzel (1992); it subsequently has been placed in its own family (refer to Gelder [1996]).
      - Xironodrilus formosus (Ellis, 1919)
Class Oligochaetous Clitellata
- Subclass Oligochaeta (Grube, 1850)
  - Order Lumbriculida (Brinkhurst & Jamieson, 1971)
    - Family Lumbriculidae (Claus, 1872)
      - Genus Eclipidrilus (Eisen, 1881)
        
        Eclipidrilus asymmetricus (Smith, 1896)
      - Genus Lumbriculus (Grube, 1844)
        
        Lumbriculus variegatus (Müller, 1774)
      - Genus Stylodrilus (Claparède, 1862)
        
        Stylodrilus heringianus (Claparède 1862) +
      - Genus Trichodrilus (Claparède, 1862)
        
        Trichodrilus allobrogum (Claparède, 1862)
  - Order Haplotaxida (Brinkhurst & Jamieson, 1971)
    - Family Haplotaxidae (Michaelsen, 1900)
      - Genus Haplotaxis (Hoffmeister, 1843)
        
        Haplotaxis gordioides (Hartmann, in Oken, 1819)
        
        (?= Haplotaxis emissarius [Forbes, 1890]; ?= Haplotaxis forbesi Smith, 1918)
  - Order Enchytraeida (Kasprzak, 1984)
    - Family Enchytraeidae (d’Udekem, 1855)
      - Genus Barbidrilus (Loden & Locy, 1980)
        
        Barbidrilus paucisetus (Loden & Locy, 1980)
      - Genus Lumbricillus (Ørsted, 1844)
        
        Lumbricillus rivalis (Levinsen, 1883; augmented Ditlevsen, 1904)
      - Genus Marionina (Pfeffer, 1890)
        
        Marionina forbesae (Smith & Welch, 1913 ?incertae sedis)
      - Genus Henlea (Michaelsen, 1899)
        
        Henlea urbanensis (Welch, 1914)
  - Order Tubificida (Jamieson, 1978)
    - Family Naididae
      - Subfamily Naidinae (Ehrenberg, 1828)
        
        Genus Allonais (Sperber, 1948)
        
        Allonais inaequalis (Stephenson, 1911) (initially misidentified as A. paraguayensis; see Kathman and Wetzel (2003)) **
        
        Allonais pectinata (Stephenson, 1910)
        
        Genus Amphichaeta (Tauber, 1879)
        
        Amphichaeta leydigi (Tauber, 1879) *
        
        Genus Bratislavia (Košel, 1976)
        
        Bratislavia unidentata (Harman, 1973)
        
        Genus Chaetogaster (von Baer, 1827)
        
        Chaetogaster diaphanus (Gruithuisen, 1828)
        
        Chaetogaster diastrophus (Gruithuisen, 1828)
        
        Chaetogaster limnaei (von Baer, 1827)
        
        Genus Dero (Oken, 1915)
        
        Dero (Aulophorus) furcata (Müller, 1773)
        
        Dero (Aulophorus) vaga (Leidy, 1880)
        
        Dero (Dero) digitata (Müller, 1773)
        
        Dero (Dero) lodeni (Brinkhurst, 1986) **
        
        (Note: [=Dero (Aulophorus) intermedius Loden and Harman, 1982, Freshwat. Invert. Biol. 1(1): 53; nom. nov pro: Aulophorus pectinatus Stephenson, 1931 nec Dero pectinatus Aiyer, 1929, vide Reynolds and Cook, 1976 – p. 153; et nec Dero intermedia Cragin, 1887, vide Reynolds and Cook, 1976 – p. 118; thus an unavailable name according to Harman in litt. 23 Sep. 1983]; the nomenclatural status of this species, as Dero lodeni, was resolved by Brinkhurst (1986, p. 253).)
        
        Dero (Dero) nivea (Aiyer, 1930)
        
        Dero (Dero) obtusa (d’Udekem, 1835)
        
        Dero (Dero) pectinata (Aiyer, 1930) *
        
        Dero (Dero) trifida (Loden, 1979) *
        
        Genus Haemonais (Bretscher, 1900)
        
        Haemonais waldvogeli (Bretscher, 1900) *
        
        Genus Nais (Müller, 1773)
        
        Nais barbata (Müller, 1773)
        
        Nais behningi (Michaelsen, 1923)
        
        Nais communis (Piguet, 1906)
        
        Nais elinguis (Müller, 1773)
        
        Nais pardalis (Piguet, 1906)
        
        Nais pseudobtusa (Piguet, 1906)
        
        Nais simplex (Piguet, 1906)
        
        Nais variabilis (Piguet, 1906)
        
        Genus Ophidonais (Gervais, 1838)
        
        Ophidonais serpentina (Müller, 1773)
        
        Genus Paranais (Czerniavsky, 1880)
        
        Paranais frici (Hrabě, 1941)
        
        Genus Piguetiella (Sperber, 1939)
        
        Piguetiella michiganensis (Hiltunen, 1967) ⁺
        
        Genus Slavina (Vejdovský, 1883)
        
        Slavina appendiculata (d’Udekem, 1855)
        
        Genus Specaria (Sperber, 1939)
        
        Specaria josinae (Vejdovský, 1883)
        
        Genus Stephensoniana (Černosvitov, 1938)
        
        Stephensoniana trivandrana (Aiyer, 1926)
        
        Genus Stylaria (Lamarck, 1816)
        
        Stylaria lacustris (Linnaeus, 1767)
        
        Stylaria fossularis (Leidy, 1852)
        
        Genus Uncinais (Levinson, 1884)
        
        Uncinais uncinata (Ørsted, 1842) ⁺
        
        Genus Vejdovskyella (Michaelsen, 1903)
        
        Vejdovskyella intermedia (Bretscher, 1896)
      - Subfamily Pristininae (Lastočkin, 1921) (formerly, Family Pristinidae (Lastočkin, 1921)) (included in Naididae sensu lato in phylogenetical classification; see Erséus et al. 2008; Erséus et al. 2010; Erséus et al. 2017 (citations below))
        
        Genus Pristina (Ehrenberg, 1828)
        Please Note: The genus Pristinella, established by Brinkhurst (1985), was suppressed by Collado and Schmelz (2000) based upon their description of two new species in the genus Pristina, and their reassessment of the morphology of Pristina and Pristinella species.
        
        Pristina acuminata (Liang, 1958)
        
        Pristina aequiseta (Bourne, 1891) (= Pristina evelinae Marcus, 1943, = Pristina foreli Piguet, 1906) *
        
        Pristina breviseta (Bourne, 1891)
        
        Pristina jenkinae (Stephenson, 1931) (= Pristina idrensis Sperber, 1948) *
        
        Pristina leidyi (Smith, 1896)
        
        Pristina longidentata (Harman, 1965) **
        
        Pristina osborni (Walton, 1906)
        
        Pristina plumaseta (Turner, 1935)
        
        Pristina sima (Marcus, 1944) *
        
        Pristina synclites (Stephenson, 1925)
    - [Family Tubificidae] (suppressed; see Erséus, Wetzel & Gustavsson, 2008 (in Literature Cited section, below))
      - Subfamily Tubificinae (Vejdovský, 1876)
        
        Genus Aulodrilus (Bretscher, 1899)
        
        Aulodrilus americanus (Brinkhurst & Cook, 1966)
        
        Aulodrilus limnobius (Bretscher, 1899)
        
        Aulodrilus pigueti (Kowalewski, 1914)
        
        Aulodrilus pluriseta (Piguet, 1906)
        
        Genus Ilyodrilus (Eisen, 1879)
        
        Ilyodrilus templetoni (Southern, 1909)
        
        Genus Isochaetides (Hrabě, 1966; emmend. Brinkhurst, 1981)
        
        Isochaetides curvisetosus (Brinkhurst & Cook, 1966) ⁺
        
        Isochaetides freyi (Brinkhurst, 1965) ⁺
        
        Genus Limnodrilus (Claparède, 1862)
        
        Limnodrilus cervix (Brinkhurst, 1963)
        
        Limnodrilus claparedianus (Ratzel, 1868)
        
        Limnodrilus hoffmeisteri (Claparède, 1862)
        
        Limnodrilus maumeensis (Brinkhurst & Cook, 1966)
        
        Limnodrilus profundicola (Verrill, 1871)
        
        Limnodrilus rubripenis (Loden, 1977)
        
        Limnodrilus tortilipenis (Wetzel, 1987)
        
        Limnodrilus udekemianus (Claparède, 1862)
        
        Genus Potamothrix (Vejdovský & Mrázek, 1902)
        
        Potamothrix bavaricus (Oschmann, 1913)
        
        Potamothrix bedoti (Piguet, 1913) ⁺
        
        Potamothrix moldaviensis (Vejdovský & Mrázek, 1902)
        
        Potamothrix vejdovskyi (Hrabě, 1941)
        
        Genus Psammoryctides (Hrabě, 1964)
        
        Psammoryctides californianus (Brinkhurst, 1965)
        
        Genus Quistadrilus (Brinkhurst, 1981)
        
        Quistadrilus multisetosus (Smith, 1900)
        
        Genus Rhizodrilus (Smith, 1900)
        
        Rhizodrilus lacteus (Smith, 1900)
        
        Genus Spirosperma (Eisen, 1879)
        
        Spirosperma ferox (Eisen, 1879)
        
        Spirosperma nikolskyi (Lastočkin & Sokolskaya, 1935) (= Spirosperma variegatus Leidy, 1851 sensu Brinkhurst, 1962) ⁺
        
        Genus Tasserkidrilus (Holmquist, 1985)
        
        Tasserkidrilus kessleri (Hrabě, 1962)
        
        Tasserkidrilus superiorensis (Brinkhurst & Cook, 1966) – considered to be restricted to the Great Lakes, but collected [by MJW, 16 April 2008] from a small, polluted stream in Marion Co., Illinois.
        
        Genus Tubifex (Lamarck, 1816)
        
        Tubifex ignotus (Štolc, 1886) ⁺
        
        Tubifex tubifex (Müller, 1774)
        
        Genus Varichaetadrilus (Brinkhurst & Kathman, 1983)
        
        Varichaetadrilus angustipenis (Brinkhurst & Cook, 1966) ¹
      - Subfamily Rhyacodrilinae (Hrabě, 1963)
        
        Genus Bothrioneurum (Štolc, 1888)
        
        Bothrioneurum vejdovskyanum (Štolc, 1888)
        
        Genus Branchiura (Beddard, 1892)
        
        Branchiura sowerbyi (Beddard, 1892)
        
        Genus Rhyacodrilus (Bretscher, 1901)
        
        Rhyacodrilus coccineus (Vejdovský, 1875)
        
        Rhyacodrilus falciformis (Bretscher, 1901) *
        
        Rhyacodrilus montanus (Brinkhurst, 1965)
        
        Rhyacodrilus cf. sodalis (Eisen, 1879) **
        
        Rhyacodrilus subterraneus (Hrabě, 1963) **
- Order Crassiclitellata (Jamieson, 1988)
  - Superfamily Lumbricoidea
    - Family Lumbricidae
      - Genus Eisenia (Malm, 1877)
        
        Eisenia foetida (Savigny, 1826)
      - Genus Eiseniella (Michaelsen, 1900)
        
        Eiseniella tetraedra (Savigny, 1826)
  - Superfamily Megascolecoidea
    - Family Sparganophilidae
      - Genus Sparganophilus (Benham, 1892)
        
        Sparganophilus meansi (Reynolds, 1980)
        
        Sparganophilus tamesis (Behnam, 1892 (= Sp. eiseni Smith, 1895)) (see Rota et al. 2016, Reeves et al. (2024), and Reynolds (2025) – citations in Literature Cited section, below)
        
        Note: Please visit this webpage highlighting the earthworms present in Illinois.
- Subclass Hirudinea (Savigny, in Lamarck, 1818)
  - Order Rhynchobdellida
    Note: several nomenclatural changes to taxa listed below have occurred; corrections will soon be made to the list, below. A current list of valid species and nomenclature is provided here: Classification and checklist of the leeches occurring in North America north of Mexico.
    - Family Glossiphoniidae
      - Genus Actinobdella (Moore, 1901)
        
        Actinobdella inequiannulata (Moore, 1901)
      - Genus Alboglossiphonia (Luken, 1976)
        
        Alboglossiphonia heteroclita (Linnaeus, 1761)
      - Genus Desserobdella (Barta & Sawyer, 1990)
        
        Desserobdella phalera (Graf, 1899)
        
        Desserobdella picta (Verrill, 1872)
      - Genus Gloiobdella (Ringuelet, 1978)
        
        Gloiobdella elongata (Castle, 1900)
      - Genus Glossiphonia (Johnson, 1816)
        
        Glossiphonia complanata (Linnaeus, 1758)
      - Genus Helobdella (R. Blanchard, 1896)
        
        Helobdella fusca (Castle, 1900)
        
        Helobdella papillata (Moore, 1906)
        
        Helobdella stagnalis (Linnaeus, 1758)
        
        Helobdella triserialis (E. Blanchard, 1849)
      - Genus Placobdella (R. Blanchard, 1893)
        
        Placobdella montifera (Moore, 1906)
        
        Placobdella multilineata (Moore, 1953)
        
        Placobdella ornata (Verrill, 1872)
        
        Placobdella papillifera (Verrill, 1872)
        
        Placobdella parasitica (Say, 1824)
        
        Placobdella pediculata (Hemingway, 1908)
      - Genus Theromyzon (Phillippi, 1867)
        
        Theromyzon biannulatum (Klemm, 1977)
    - Family Piscicolidae
      - Genus Cystobranchus (Diesing, 1859)
        
        Cystobranchus verrilli (Meyer, 1940)
      - Genus Myzobdella (Leidy, 1851)
        
        Myzobdella lugubris (Leidy, 1851)
      - Genus Piscicola (de Blainville, 1818)
        
        Piscicola milneri (Verrill, 1874)
        
        Piscicola punctata (Verrill, 1871)
      - Genus Piscicolaria (Whitman, 1889)
        
        Piscicolaria reducta (Meyer, 1940)
  - Order Arhynchobdellida
    - Family Haemopidae
      - Genus Haemopis (Savigny, 1822)
        
        Haemopis marmorata (Say, 1824)
        
        Haemopis terrestris (Forbes, 1890)
    - Family Hirudinidae
      - Genus Macrobdella (Verrill, 1872)
        
        Macrobdella decora (Say, 1824)
      - Genus Philobdella (Verrill, 1874)
        
        Philobdella gracilis (Moore, 1901)
    - Family Erpobdellidae
      - Genus Dina (R. Blanchard, 1892)
        
        Dina dubia (Moore & Meyer, 1951)
        
        Dina parva (Moore, 1912)
      - Genus Erpobdella (de Blainville, 1818)
        
        Erpobdella punctata punctata (Leidy, 1870)
      - Genus Mooreobdella (Pawlowski, 1955)
        
        Mooreobdella fervida (Verrill, 1871)
        
        Mooreobdella microstoma (Moore, 1901)
      - Genus Nephelopsis (Verrill, 1872)
        
        Nephelopsis obscura (Verrill, 1872)

¹ Notes on V. angustipenis: A Nearctic species, V. angustipenis originally was described as Limnodrilus angustipenis from material collected by J. K. Hiltunen from Saginaw Bay, Lake Huron; it was assigned to the genus Varichaetadrilus by R. O. Brinkhurst in 1989. This rarely collected species presently is known to occur in Alabama, Illinois, Louisiana, Manitoba, Michigan, North Carolina, New Brunswick, New Jersey, Ohio, Ontario, and Tennessee (Coates and Wetzel, manuscript in prep.); many of these distributional records, however, are from one or two collections, primarily oligotrophic lakes. During a continuing faunal survey of springs in southern Illinois (1990-), I have collected V. angustipenis from nine of 17 springs and their associated springruns (Webb et al. 1995; Webb et al. 1996, 1998b). Recently, I identified specimens from five springs in Texas [manuscript in prep.], from additional springs in western Illinois (Webb et al. 1998a), from caves in Illinois and Missouri [INHS Annelida Collection records; Wetzel and Taylor (2001)], and from several other streams in Illinois (many with groundwater influence). With the exception of seven stream sites in southern Illinois (Wetzel 1982a), extensive collecting and identification of aquatic oligochaetes throughout North America have revealed few specimens of this taxon.

Literature Cited and Selected References

Barta J.R., and R.T. Sawyer. 1990. Definition of a new genus of glossiphoniid leech and a redescription of the type species, Clepsine picta Verrill, 1872. Can. J. Zool. 68(9): 1942-1950.

Bartolomaeus, T. 1997. Chaetogenesis in polychaetous Annelida – significance for annelid systematics and the position of the Pogonophora. Zoology – Analysis of Complex Systems 100(4): 348-364.

Brinkhurst, R O. 1982. Evolution in the Annelida. Can. J. Zool. 60(5): 1043-1059.

Brinkhurst, R O. 1985. The generic and subfamilial classification of Naididae (Annelida: Oligochaeta). Proceedings of the Biological Society of Washington 98(2): 470-475.

Brinkhurst, R.O. 1986. Guide to the freshwater aquatic microdrile oligochaetes of North America. Can. Spec. Publ. Fish. Aquat. Sci. 84. vi + 259 pp.

Brinkhurst, R.O. 1988. A taxonomic analysis of the Haplotaxidae. Can. J. Zool. 66(10): 2243-2252.

Brinkhurst, R.O. 1989. A phylogenetic analysis of the Lumbriculidae (Annelida, Oligochaeta). Can. J. Zool. 67(11): 2731-2739.

Brinkhurst, R. O. 1991a. Ancestors. Mitt. Hamburg Mus. Inst. 88(1): 97-110.

Brinkhurst, R.O. 1991b. A phylogenetic analysis of the Tubificinae (Annelida, Oligochaeta). Can. J. Zool. 69(2): 392-397.

Brinkhurst, R.O. 1991c. A phylogenetic analysis of the Phreodrilidae (Annelida, Oligochaeta). Can. J. Zool. 69(8): 2031-2040.

Brinkhurst, R.O. 1992. Evolutionary relationships within the Clitellata. Soil. Biol. Biochem. 24(12): 1202-1205.

Brinkhurst, R.O. 1994. Evolutionary relationships within the Clitellata: an update. Megadrilogica 5(10: 109-112.

Brinkhurst, R.O. 1999. Lumbriculids, branchiobdellidans and leeches: an overview of recent progress in phylogenetic research on clitellates. Hydrobiologia 406: 281-290.

Brinkhurst, R.O., and S.R. Gelder. 1989. Did the lumbriculids provide the ancestors of the branchiobdellidans, acanthobdellidans and leeches? Hydrobiologia 180: 7-15.

Brinkhurst, R.O., and B.G.M. Jamieson. 1971. Aquatic Oligochaeta of the world. University of Toronto Press, Toronto, ON, and Buffalo, NY. xi + 860pp. [out of print].

Brinkhurst, R.O., and A.F.L. Nemec. 1987. A comparison of phenetic and phylogenetic methods applied to the systematics of Oligochaeta. Hydrobiologia 155: 65-74.

Brinkhurst, R.O., and M.J. Wetzel. 1984. Aquatic Oligochaeta of the world: Supplement. A catalogue of new freshwater species, descriptions, and revisions. Canadian Tech. Rpt. Hydrography Ocean Sci. No. 44. v + 101 pp.

Claus, C.F.W. 1872. Grundzüge der Zoologie, zum Gebrauche an Universitäten und höheren Lehranstalten. Leitfaden zur Einführung in das wissenschaftliche Studium der Zoologie. 2nd Edition. N. G. Elwert’sche Universitäts-Buchhandlung, Marburg and Leipzig, 1170 pp. https://doi.org/10.5962/bhl.title.3783

Coates, K.A. 1987. Phylogenetics of some Enchytraeidae (Annelida: Oligochaeta): A preliminary investigation of relationships to the Haplotaxidae. Hydrobiologia 155: 91-106.

Coates, K.A. 1989. Phylogeny and origins of Enchytraeidae. Hydrobiologia 180: 17-33.

Coates, K.A. and M.J. Wetzel. 1998. Manuscript in prep. Common and scientific names of aquatic invertebrates from the United States and Canada: Clitellate annelids, including Aphanoneura, Branchiobdellae, Hirudinea, and Oligochaeta.

Collado, R., and R.M. Schmelz. 2000. Pristina silvicola and Pristina terrena spp. nov., two new soil-dwelling species of Naididae (Oligochaeta, Annelida) from the tropical rain forest near Manaus, Brazil, with comments on the genus Pristinella. Journal of Zoology, London 252: 509-516.

Ehrenberg [C.G.] 1831. Symbolae physicae, animalia evertebrata, exclusis insectis. Series prima cum tabularum decade prima. Phytozoa. Ex Officina Academica, venditur a Mittlero, Berlin, 70 pp.

Erséus, C. 1987. Phylogenetic analysis of the aquatic Oligochaeta under the principle of parsimony. Hydrobiologia 155: 75-89.

Erséus, C. 1990. Cladistic analysis of the subfamilies within the Tubificidae (Oligochaeta). Zool. Scr. 19(1): 57-63.

Erséus, C., I. Envall, P. De Wit, L.M. Gustavsson. 2017. Molecular data reveal a tropical freshwater origin of Naidinae (Annelida, Clitellata, Naididae). Molecular Phylogenetics and Evolution 115: 115–127.

Erséus, C., I. Envall, M. Marchese, L.M. Gustavsson. 2010. The systematic position of Opistocystidae (Annelida, Clitellata) revealed by DNA data. Molecular Phylogenetics and Evolution 54(1): 309–313.

Erséus, C., M.J. Wetzel, and L. Gustavsson. 2008. ICZN rules – a farewell to Tubificidae (Annelida, Clitellata). Zootaxa 1744: 66-68 [9 April].

Evans, C.R. 1939. The Branchiobdellidae (Annelida) on crayfishes of Champaign County, Illinois. Journal of Parasitology, 25: 448.

Gelder, S.R. 1996. A review of the taxonomic nomenclature and a checklist of the species of the Branchiobdellae (Annelida: Clitellata). Proc. Biol. Soc. Washington 109(4): 653-663.

Gelder, S.R. 2020. An ongoing saga: Endemic branchiobdellidans (Annelida: Clitellata) on translocated commercial North American crayfish. Zoosymposia 17(1): 141-158.

Gelder, S.R., and R.O. Brinkhurst. 1990. An assessment of the phylogeny of the Branchiobdellida (Annelida: Clitellata), using PAUP. Can. J. Zool. 68(6): 1318-1326.

Gelder, S.R., N.L. Gagnon, and K. Nelson. 2002. Taxonomic considerations and distribution of the Branchiobdellida (Annelida: Clitellata) of the North American continent. Northeastern Naturalist 9(4): 451-468.

Goodnight, C.J. 1940. The Branchiobdellidae (Oligochaeta) of North American crayfish. Illinois Biological Monographs 17(3): 1–75.

Govedich, F.R., B.A. Bain, W.E. Moser, S.R. Gelder, R.W. Davies, and R.O. Brinkhurst. 2010. Annelida (Clitellata) Oligochaeta, Branchiobdellida, Hirudinida, and Acanthobdellida. Pages 385-436, In: J.H. Thorp and A.P. Covich (eds). Ecology and classification of North American freshwater invertebrates. Third Edition xiv + 1,021 pp. Academic Press / Elsevier, San Diego, CA. ISBN 978-0-12-374855-3. [elsevier.com] USD$ 139.95 / 93.95 Euro.

Grube, A.E. 1850. Die Familien der Anneliden. Archiv für Naturgeschichte, 16, 249–364.

Holt, P.C. 1986. Newly established families of the order Branchiobdellida (Annelida: Clitellata) with a synopsis of the genera. Proc. Biol. Soc. Washington 99(4): 676-702.

Holt, P.C. 1989. Comments on the classification of the Clitellata. Hydrobiologia 180: 1-5.

Holt, P.C. 1982. A new species of the genus Cambarincola (Clitellata: Branchiobdellida) from Illinois with remarks on the bursa of Cambarincola vitreus Ellis, 1919, and the status of Sathodrilus Holt, 1968. Proceedings of the Biological Society of Washington 95( ): 251–255.

Holt, P.C., and B.D. Opell. 1993. A checklist of and illustrated key to the genera and species of the Central and North American Cambarincolidae (Clitellata: Branchiobdellida). Proc. Biol. Soc. Washington 106(2): 251-295.

Jamieson, B.G.M. 1978. Phylogenetic and phenetic systematics of the opisthoporous Oligochaeta (Annelida: Clitellata). Evol. Theor. 3(4): 195-233.

Jamieson, B.G.M. 1980. Preliminary discussion of an Hennigian analysis of the phylogeny and systematics of opisthoporous oligochaetes. Rev. Ecol. Biol. Sol 17(2): 261-275.

Jamieson, B.G.M. 1988. On the phylogeny and higher classification of the Oligochaeta. Cladistics 4: 367-410.

Jones, S.R.M., and P.T.K. Woo. 1990. Redescription of the leech Desserobdella phalera (Graf, 1899) n.comb. (Rhynchobdellida: Glossiphoniidae), with notes on its biology and occurrence on fishes. Can. J. Zool. 68(9): 1951-1955.

Kasprzak, K. 1982. Problems of the origin of oligochaetes (Annelida: Oligochaeta). Prezgl. Zool. 26: 145-160. (English translation, Can. Transl. Fish. Aquat. Sci. No. 4996).

Kasprzak, K. 1984. The previous and contemporary conceptions on phylogeny and systematic classifications of Oligochaeta (Annelida). Annales Zoologici, Warszawa [Ann. Zool. Polska Akad. Nauk Inst. Zool.] 38: 205–223.

Kathman, R.D., and R.O. Brinkhurst. 1998, 1999. Guide to the freshwater oligochaetes of North America. Aquatic Resources Center, College Grove, TN 37046 USA. iv + 264 pp. To ordering this guide, contact Kathman, Ph.D., [r.deedee.kathman'[AT]’gmail.com] ***

Kathman, R.D., and M.J. Wetzel. 2003. Allonais inaequalis (Annelida: Oligochaeta: Tubificidae) in North America. Proceedings of the Biological Society of Washington 116(3): 548-556.

Klemm, D.J., ed. 1985. A guide to the freshwater Annelida (Polychaeta, naidid and tubificid Oligochaeta, and Hirudinea) of North America. Kendall/Hunt Publishing Company, Dubuque, IA. xiii + 198 pp.

Lamarck, J.B.P.A. 1818. Histoire naturelle des animaux sans vertèbres, présentant les caractères généraux et particuliers de ces animaux, leur distribution, leurs classes, leurs familles, leurs genres, et la citation des principales espèces qui s’y rapportent; précédée d’une introduction offrant la détermination des caractères essentiels de l’animal, sa distinction du végétal et des autres corps naturels; enfin, l’exposition des principes généraux de zoologie. Tome cinquième. Arachnides, Crustacés, Annélides, Cirrhipèdes. J. B. Baillière, Paris, 612 pp.

Lastočkin, D.A. 1921. Issledovanija po faune Ivanovo-Voznesenskoj gubernii. 3. Fauna Oligochaeta limicola rajona issledovanija. Izvestija Ivanovo-Voznesenskogo Politehničeskogo Instituta, 4, 70–77.

Madill, J., K.A. Coates, M.J. Wetzel, S.R. Gelder. 1992. Common and scientific names of aphanoneuran and clitellate annelids of the United States and Canada. Soil Biol. Biochem. 24(12): 1259-1262.

Martin, P., I. Kaygorodova, D.Yu. Sherbakov and E. Verheyen. 2000. Rapidly evolving lineages impede the resolution of phylogenetic relationships among Clitellata (Annelida). Molecular Phylogenetics and Evolution 15(3): 355-368.

Martin, P., E. Martinez-Ansemil, A. Pinder, T. Timm, and M.J. Wetzel. 2008. Global diversity of oligochaetous clitellates (“Oligochaeta”; Clitellata) in freshwater. Pp. 117-127, In E. V. Balian, C. Léveque, H. Segers & K. Martens, eds. Freshwater Animal Diversity Assessment. Hydrobiologia 595(1).

McHugh, D. 1999. Phylogeny of the Annelida: Sidall et al. (1998) rebutted. Cladistics 15: 85-89.

McHugh, D. 2000. Molecular phylogeny of the Annelida. Canadian Journal of Zoology 78(11): 1873-1884.

Nemec, A.F.L., and R.O. Brinkhurst. 1987. A comparison of methodological approaches to the subfamilial classification of the Naididae (Oligochaeta). Can. J. Zool. 65(3): 691–707.

Oken, L. 1819. Versammlung der Schweizer Naturforscher zu St. Gallen, vom 26, 27. und 28. Julius 1819. In: Oken, L. (Ed.), Isis oder Encyclopädische Zeitung von Oken. Litterarischer Anzeiger, Jena, Nr. XLV, Erster Band, Heft I–VI, pp. 178–179.

Omodeo, P. 1998. History of Clitellata. Italian Journal of Zoology 65: 51-73.

Omodeo, P. 2000. Evolution and biogeography of megadriles (Annelida, Clitellata). Italian Journal of Zoology 67(2): 179-201.

Parker, S.P. 1982. Synopsis and classification of living organisms. McGraw Hill, New York. 1,232 pp.

Purschke, G. 1999. Terrestrial polychaetes — models for the evolution of the Clitellata (Annelida)? Hydrobiologia 406: 87-99.

Purschke, G., W. Westheide, D. Rohde, and R.O. Brinkhurst. 1993. Morphological reinvestigation and phylogenetic relationship of Acanthobdella peledina (Annelida, Clitellata). Zoomorphology 113(2): 91-101.

Readel, A.M., C.A. Phillips, and M.J. Wetzel. 2008. Leech parasitism in a turtle assemblage: effects of host and environmental characteristics. Copeia 2008(1): 225-231.

Reeves, W.K., J.W. Reynolds and M.J. Wetzel. 2024. Sparganophilus (Annelida, Oligochaeta, Sparganophilidae) in North America. Megadrilogica 28(5): 53–80. April. [paper in English; abstracts in English, French, Spanish, German; with 18 colour plates].

Reynolds, J.W. 2025. The Genus Sparganophilus is Exploding in North America! Global Academic and Scientific Journal of Multidisciplinary Studies (GASJMS) 3(1): 22–27.

Reynolds, J.W., and D.G. Cook. 1976. Nomenclatura oligochaetologica. A catalogue of names, descriptions and type specimens of the Oligochaeta. Univ. New Brunswick, Fredericton, New Brunswick. x + 217 pp. [This catalog, and the first supplement, cited below, are available from University of New Brunswick Bookstore, P. O. Box 4400, Fredericton, NB, Canada E3B 5A3, for CAN$20.00 (which includes GST {Canada’s goods and services tax}; the second and third supplements, also cited below, are available, at no charge, from the New Brunswick Museum, 277 Douglas Avenue, Saint John, NB, Canada E2K 1E5.].

Reynolds, J.W., and D.G. Cook. 1981. Nomenclatura oligochaetologica. Supplementum primum. A catalogue of names, descriptions and type specimens of the Oligochaeta. [published by the University of New Brunswick] Fredericton, New Brunswick. v + 39 pp. [to order, see note above].

Reynolds, J.W., and D.G. Cook. 1989. Nomenclatura oligochaetologica. Supplementum secundum. A catalogue of names, descriptions and type specimens of the Oligochaeta. New Brunswick Mus. Monogr. Ser. (Nat. Sci.) No. 8. v + 37 pp. [this second supplement is available, at no charge, from the New Brunswick Museum, 277 Douglas Avenue, Saint John, NB, Canada E2K 1E5.].

Reynolds, J.W., and D.G. Cook. 1993. Nomenclatura oligochaetologica. Supplementum tertium. A catalogue of names, descriptions and type specimens of the Oligochaeta. New Brunswick Mus. Monogr. Ser. (Nat. Sci.) No. 9. vi + 33 pp. [this third supplement is available, at no charge, from the New Brunswick Museum, 277 Douglas Avenue, Saint John, NB, Canada E2K 1E5.].

Reynolds, J.W. and M.J. Wetzel. 2004. Terrestrial Oligochaeta (Annelida: Clitellata) in North America north of Mexico. Megadrilogica 9(11): 71–98.

Reynolds, J.W., and M.J. Wetzel. 2008. Terrestrial Oligochaeta (Annelida: Clitellata) in North America, including Mexico, Puerto Rico, Hawaii, and Bermuda. Megadrilogica 12 (12): 157–208.

Reynolds, J.W., and M.J. Wetzel. 2011. The earthworms (Oligochaeta: Acanthodrilidae, Glossoscolecidae, Komarekionidae, Lumbridicae, Megascolecidae, and Sparganophilidae) of Illinois, USA. Megadrilogica 15(4): 35–67.

Reynolds, J.W., and M.J. Wetzel. 2012. Terrestrial Oligochaeta (Annelida: Clitellata) in North America, including Mexico, Puerto Rico, Hawaii, and Bermuda. III. Megadrilogica 15(8): 191–209.

Reynolds, J.W. and M.J. Wetzel. 2025. Nomenclatura Oligochaetologica – A catalogue of names, descriptions and type specimens of the Oligochaeta. Editio Secunda.
URL: https://nomenclatura-oligochaetologica.inhs.illinois.edu (date accessed: xx-xx-2025).

Rota, E., S. Martinsson, M. Bartoli, A. Beylich, U. Graefe, A. Laini, M.J. Wetzel, and C. Erséus. 2016. Mitochondrial evidence supports a Nearctic origin for the spreading limicolous earthworm Sparganophilus tamesis Benham, 1892 (Clitellata, Sparganophilidae). Contributions to Zoology 85(1): 113-119.

Sawyer, R.T. 1986a. Leech biology and behaviour. Volume I. Anatomy, physiology, and behaviour. Clarendon Press, Oxford. xiv + pp. 1-417 + index.

Sawyer, R.T. 1986b. Leech biology and behaviour. Volume II. Feeding biology, ecology, and systematics. Clarendon Press, Oxford. xiv + pp. 419-793 + index.

Sawyer, R.T. 1986c. Leech biology and behaviour. Volume III. Bibliography. Clarendon Press, Oxford. x + pp. 799-1,065.

Schmelz, R.M., C. Erséus, P. Martin, T. van Haaren, and T. Timm. 2021. A proposed order-level classification in Oligochaeta (Annelida, Clitellata). Zootaxa 5040(4): 589–597. https://doi.org/10.11646/zootaxa.5040.4.9

Siddall, M.E. and E.M. Burreson. 1995. Phylogeny of the Euhirudinea: independent evolution of blood feeding by leeches? Canadian Journal of Zoology 73: 1048-1064.

Siddall, M.E. and E.M. Burreson. 1996. Leeches (Oligochaeta?: Euhirudinea), their phylogeny and the evolution of life-history strategies. Hydrobiologia 334: 277-285.

Siddall, M. E. and E. M. Burreson. 1998. Phylogeny of leeches (Hirudinea) based on mitochondrial cytochrome c oxidase subunit I. Molecular Phylogenetics and Evolution 9(1): 156-162.

Siddall, M.E., K. Fitzhugh and K.A. Coates. 1998. Problems determining the phylogenetic position of echiurans and pogonophorans with limited data. Cladistics 14: 401-410.

Sima, P. 1993. Phylogeny and classification of annelids. Pages 13-25, In: V. Vetvicka, P. Sima, E. L. Cooper, M. Bilej and P. Roch. 1993. Immunology of annelids. CRC Press, Boca Raton, FL (USA), Ann Arbor, MI (USA), London, Tokyo. 299 pp.

Timm, T. 1981. On the origin and evolution of aquatic Oligochaeta. Eesti NSV Tead. Akad. Toim. Biologia 30(3): 174-181.

Trapido-Rosenthal, H.G., I.B. Kuffner, L. McDowell and K.A. Coates. 1997. Paraphyly of the annelids: all annelids are not segmented worms. Bulletin of Marine Science 60(2): 624.

Udekem, J.d‘ 1855. Nouvelle classification des Annélides sétigères abranches. Bulletins de l’Académie Royale Belgique, classe des sciences, 22, 533–555.

Webb, D.W., M.J. Wetzel, P.C. Reed, L.R. Phillippe, and M.A. Harris. 1995. Aquatic biodiversity in Illinois springs. Journal of the Kansas Entomological Society 68(2)suppl.: 93-107.

Webb, D. W., M.J. Wetzel, P.C. Reed, L.R. Phillippe, and T.C. Young. 1996. Biodiversity, hydrogeology, and water quality of 10 karst springs in the Salem Plateau Section of Illinois. Pp. 146-185, In: Davis, M., Editor. Research on agricultural chemicals in Illinois groundwater: Status and future directions VI. Illinois Groundwater Consortium, Office of Research Development and Administration, Southern Illinois University, Carbondale.

Webb, D.W., M.J. Wetzel, and L.R. Phillippe. 1998. [Webb et al. 1998a]. The aquatic biota and groundwater quality of Illinois springs in the Lincoln Hills, Wisconsin Driftless, and Northern Till Plains Sections of Illinois. Illinois Natural History Survey Center for Biodiversity Technical Report 1998 (6). 164 pp. Prepared for the Environmental Protection Trust Fund Commission and the Illinois Department of Natural Resources, Division of Energy and Environmental Assessment, Office of Realty and Environmental Planning, Springfield.

Webb, D.W., M.J. Wetzel, P.C. Reed, L.R. Phillippe, and T.C. Young. 1998. [Webb et al. 1998b]. The macroinvertebrate biodiversity, water quality, and hydrogeology of ten karst springs in the Salem Plateau Section of Illinois, USA. Pp. 39-48, In: L. Botosaneanu (ed.). Studies in Crenobiology – The biology of springs and springbrooks. Backhuys Publishers, Leiden, The Netherlands. 261 pp. ISBN 90-73348-04-8.

Westheide, W., D. McHugh, G. Purschke and G. Rouse. 1999. Systematization of the Annelida: different approaches. Hydrobiologia 402: 291-307.

Wetzel, M.J. 1982a. The distribution and relative abundance of aquatic Oligochaeta in the Cache River system, southern Illinois, in relation to water quality. Unpublished MSc. thesis, Eastern Illinois University, Charleston. x + 182 pp.

Wetzel, M.J. 1982b. Aquatic Oligochaeta (Annelida: Clitellata) in Kansas, with notes on their distribution and ecology. Technical Publication of the State Biological Survey of Kansas 12:112-130.

Wetzel, M.J. 1992. Aquatic Annelida of Illinois: Introduction and checklist of species. Trans. Illinois St. Acad. Sci. 85(1 and 2): 87-101.

Wetzel, M.J., S.V. Fend, K.A. Coates, R.D. Kathman, and S.R. Gelder. 2009. Taxonomy, systematics, and ecology of the freshwater oligochaetes and branchiobdellidans (Annelida, Clitellata) of North America. A workbook. 1 November 2009. vi + 280 pp. + color plates.

Wetzel, M.J., and S.J. Taylor. 2001. First records of freshwater oligochaetes (Annelida, Clitellata) from caves in Illinois and Missouri, USA. Journal of Cave and Karst Studies 63(3): 99-104.

Wetzel, M.J. and J.W. Reynolds. 2021. A preliminary inventory of earthworms (Annelida, Clitellata) of the Nachusa Grasslands Area, Lee and Ogle Counties, Illinois, USA. Megadrilogica 26(8): 91–125.