Classification and Checklist of the Freshwater Oligochaetes
Occurring in the State of Florida, USA

Compiled by M.J. Wetzel, R.D. Kathman, S.V. Fend, and K.A. Coates.

Introduction. A current list of freshwater oligochaetes occurring in Florida, USA, is presented here; this is a subset of a website (established in November 2005, and updated periodically with this site) that presents a Classification and checklist of freshwater oligochaetes occurring in North America north of Mexico.

While this list is presently limited in scope to the freshwater oligochaetes occurring in Florida, we plan to expand this list (perhaps in a tabular format) to include the marine, estuarine, and brackish-water oligochaetes occurring in/adjacent to the State, then later add the terrestrial oligochaetes, leeches, branchobdellidans, and aeolosomatidans (now aligned with the Polychaeta). Because several megadrile oligochaetes are commonly found in aquatic and semi-aquatic habitats, those species will soon be included in this list. Recent publications by Gelder et al. (2002) and Reynolds and Wetzel (2004) provided lists of North American species of branchiobdellidans and terrestrial (megadrile) oligochaetes, respectively; both publications presented accurate classification and nomenclature for their respective groups. A 'second edition' of Reynolds and Wetzel (2008) included new records for species included in their 2004 paper, and expanded the scope to include published records of megadriles occurring in Mexico, Puerto Rico, Hawaii, and Bermuda, and distribution maps for all species.

Don Klemm, Bill Moser, and Mark Wetzel just established a website entitled Classification and Checklist of the Leeches (Phylum Annelida: Class Clitellata: Subclass Hirudinida) occurring in North America North of Mexico. This site includes updated information pertinent to Klemm's (1993) guide to leeches of Florida.

This list is made possible through the collective efforts of the authors of this website, and in acknowledgment of our years of collaboration with Ralph Brinkhurst, John W. Reynolds, Donald J. Klemm, Douglas R. Spencer, Christer Erséus, Tarmo Timm, Michael Milligan (now deceased), and numerous other oligochaetologists working in Florida, in North America, and elsewhere in the World. We hope to expand this list, including the leeches and branchiobdellidans occurring in Florida, with systematic and nomenclatural expertise for those groups provided by Donald J. Klemm and William Moser, and Stuart R. Gelder, respectively. Please note that a website listing the freshwater leeches occurring in North America was recently established by Klemm, Moser, and Wetzel, and is linked from the navigator bar at the bottom of this page.

Finally, our efforts to maintain an accurate and current checklist is dependent upon our continuing collaboration with aquatic biologists working in Florida, especially those associated with the Florida Department of Environmental Protection [FDEP], Southwest Florida Water Management District, Reedy Creek Improvement District, St. Johns River Water Management District, Environmental Protection Commission of Hillsborough County, Water and Air Research, Inc., Terra Environmental Services, Inc., Florida Association of Benthologists, and other public and private agencies and firms conducting benthological studies in the State.

As of 9 April 2008, 117 nominal species representing 46 genera and five families of freshwater (microdrile) oligochaetes are known to occur in Florida. We present here a general classification and checklist of these species based upon records published in the peer-reviewed literature, those published in Milligan (1997), and those specimens in the FDEP SBIO database that have been independently verified. While many additional records of aquatic oligochaetes exist--representing specimens identified to the genus and species levels during the completion of a variety of privately and publicly funded projects at the local, state, regional, federal, and provincial government levels--those records remain 'unavailable' to science in general because 1) preliminary identifications of specimens have not been verified by recognized taxonomists and systematists, and 2) these data have not been published in the peer-reviewed literature. The unfortunate reality here is that many of these identified yet unpublished records likely could represent new distributional information for taxa at the drainage, state, provincial, and possibly continental level. Furthermore, many of these records could represent a) new (undescribed) species, b) identified species that are considered rare throughout their known distributional range, c) species for which few mature specimens have been examined, and/or d) new ecological affiliations (geographical, physical, biological, water quality). Several species of megadrile oligochaetes (classified in traditional 'earthworm' families) also occur in Florida, and will be listed here soon.

Source of information for species included in this checklist. The freshwater oligochaete fauna of North America, particularly the species in the families Naididae and Tubificidae (see footnote 1, below), and Lumbriculidae, has been documented in publications such as Brinkhurst (1978, 1986), Brinkhurst and Jamieson (1971), Brinkhurst and Wetzel (1984), Hiltunen and Klemm (1980, 1985), Brinkhurst and Gelder (2001), Govedich et al. (2010), Kathman and Brinkhurst (1998), and Stimpson et al. (1982, 1985). State and regional keys, guides, and papers with species distributions include Howmiller and Loden (1976), Kathman and Wetzel (2003), Milligan (1997), Smith (1984), Spencer (1980), Spencer and Denton (2003), Spencer and Hudson (2003), Spencer and Wisseman (1993), Strayer (1990), Wetzel (1982, 1992), Wetzel and Taylor (1999), Wetzel et al. (2009), and Whitley (1982). Please consult the literature cited in each of the above publications for additional - and usually more specific - distributional information for taxa.

Historically, the family Lumbriculidae has received less attention than other families of microdrile oligochaetes. Fortunately, extensive work with this group by Steve Fend and others, particularly in the western and southeastern regions of the U.S., has resulted in a significant increase in our distributional knowledge of this group. In addition, their research and publications have provided us with descriptions of several new lumbriculid genera and species as well as redescriptions and new combinations of yet other previously described / problematical taxa (e.g., Fend and Brinkhurst 2000, 2010; Fend and Gustafson 2001; Fend and Rodriguez 2003; Fend 2005a, 2005b, 2006, 2009; and Fend and Lenat 2007). While a few of these papers are relatively specific to taxa occurring in northwestern North America, information presented therein provides insight to taxa known or thought likely to occur in Florida and elsewhere in the southeastern U.S.

Nomenclature. Nomenclatural changes and synonymies for North American species published since Kathman and Brinkhurst (1998) have been included herein; to the best of our knowledge, the nomenclature for species discussed herein is current as of 6 January 2014.

On Monday 6 January 2014, Mark J. Wetzel (INHS) and John Reynolds (Oligochaetology Lab) launched a new website presenting the second edition of Nomenclatura Oligochaetologica, as Nomenclatura Oligochaetologica Editio Secunda – a catalogue of names, descriptions, and type specimens of the Oligochaeta. This web-based Second Edition of Nomenclatura Oligochaetologica [N.O.2]: integrates the accounts included in the first volume (Reynolds and Cook, 1976 [N.O.]) with those presented in the three supplements (Reynolds and Cook, 1981 [N.O.S.P.], 1989 [N.O.S.S.], and 1993 [N.O.S.T.]) – together comprising the original N.O. series; updates and corrects accounts for the generic, subgeneric, specific, and infra-specific names of oligochaetes (Annelida, oligochaetous Clitellata) as presented in the original series; adds accounts for all oligochaete taxa described as new to science since 1993 – including barcode, GenBank, tissue repository, and other pertinent DNA sequencing information; expands the Index Auctorum, Index Auctoritatum, Index Museorum, Prolegomenon, Glossarium, and References sections of the original series; includes translations (in 11 languages) of the Prolegomena and Glossaria of the original series and this second edition; presents and expands Dedicatio sections in the original series with biographies and memoria for our historical and contemporary colleagues focused on oligochaetology, including bibliographies of their published scientific contributions; offers a forum for Current Perspectives in oligochaete phylogeny, taxonomy, systematics, and nomenclature; provides a Using This Nomenclator section with account examples and instructions for using and navigating this web-based catalogue; includes an annotated list of links to web-based annelid resources.

The advantage of a web based edition is that it allows for corrections and additions to occur immediately after the information reaches us. We encourage assistance from our colleagues around the World to correct errors, update the information for each taxon account as presented, and to provide complete account information for newly described taxa as well as those that we have overlooked.

Note that the N.O. series does not include synonymies of species; you are directed to the synonymies for aquatic oligochaetes presented in Brinkhurst and Jamieson (1971), and more recent publications in the primary literature specific to new descriptions and revisions at the family, generic, and species levels.

Classification. In recent years several proposed classifications for annelids have been presented in the literature; one such classification divided the phylum Annelida into two subphyla: the Aclitellata (with the two classes Aphanoneura and Polychaeta) and the Clitellata (including the Acanthobdellida, Branchiobdellida, 'Oligochaeta', and Hirudinea). Yet, because the name 'Aclitellata' is a general term -- established only as an equivalent ranking to the Clitellata rather than being based on phylogenetic studies -- we think it is better to refer to the two groupings as clitellates and non-clitellates. Regardless, this checklist is not the place to discuss or resolve the most appropriate or accurate classification for the 'Oligochaeta', let alone for the entire Phylum Annelida.

Indeed, the nominal taxon 'Oligochaeta' is paraphyletic, and thus - in a strict cladistic approach - invalid if using in exclusion of leeches. The corresponding monophylon, Clitellata, includes the leeches and leech-like successors of the 'Oligochaeta'. Thus, Oligochaeta (with a new meaning) and Clitellata are synonyms; however, because the Code (ICZN 1999) does not use priority rules for taxa above the family-group level, either name is acceptable for use. Clitellata may be preferred as it retains its original meaning. Based upon the extensive discussions presented by Martin et al. (2000), Martin (2001), Timm (2005), Erséus (2005), and the recent phylogenetic work of Erséus et al. (2000), Erséus and Gustavsson (2002), Erséus et al. (2002), Erséus and Källersjö (2004), and Envall et al. (2006), the term 'oligochaetous Clitellata' (oligochaetoid Clitellata) also could be used for the oligochaetes instead of the name 'Oligochaeta' (albeit these two terms seem awkward) -- at least until this issue is more formally addressed. Timm (2005, p. 57) does temper us with this observation: "However, any good idea can be discredited when developed into [the] absurd, and so does the principle of monophyly in systematics." If you are intrigued by this quandary, you are directed to read the above papers (as well as those cited therein, including Ashlock, 1971, and Valentine, 2004) [citations provided below]). We thus use the term "oligochaetous Clitellata" in the checklist, below.

Introduction to the checklist. The primary purpose of this checklist is to present a current list of freshwater oligochaetes occurring in Florida, using the current accepted nomenclature for each species as published in the primary literature. The authority(ies) and date of original description in the literature (as cataloged in the Nomenclatura Oligochaetologica series) are also provided for each species.

Please read the information associated with footnotes 1 and 2, below, regarding the recent ruling by the International Commission on Zoological Nomenclature (ICZN), and review the paper by Erséus, Wetzel, and Gustavsson (2008 - Zootaxa 1744: 66-68; published 9 April) and how it affects our use of the familial and subfamilial classifications of Naididae and Tubificidae.

Recent synonymies and other invalid names will follow the correct name for taxa - vis., [= Genus-name species-name authority(ies), date]; other notations after species entries will be explained at the bottom of the list.

-)(- = See section on Classification (in text, above).

* = species not yet reported in the literature as occurring in Florida, but thought likely to occur in the state because of their documented occurrence in other states in the southeastern U.S.

i = species considered to have been introduced to North America. [Note that several other species listed here could be cryptogenic (of unknown origin, possibly representing historical introductions)].

? = taxon whose status as a valid species is considered questionable.

?? = occurrence in North America considered questionable /or/ based upon erroneous record [misidentification or lost specimen(s)] (see footnote).

$? = possible/probable new record for state of Florida. [Specimens of these two species, and of three others (all in the Family Naididae, subfamilies Naidinae and Pristininae) are considered new to Florida, with at least two of these considered to be new records for North America. This list will be updated to reflect these new records after taxonomic review has been completed (Wetzel et al., mss in prep.).

Classification and Checklist of the Freshwater Oligochaetes
occurring in Florida, USA


Phylum Annelida
Class Clitellata
Michaelsen, 1919
Oligochaetous Clitellata [= 'Oligochaeta'] -)(-
Superorder Microdrili

- - - - - Order Haplotaxida - - - - -

Family Haplotaxidae Michaelsen, 1900

    Genus Haplotaxis Hoffmeister, 1843
    Haplotaxis cf. gordioides (Hartmann, 1821)
    [ ? Haplotaxis ichthyophagus Gates, 1971 - incertae sedis]

    Note: the genera Metataxis and Tiguassu were removed
    from this family by Brinkhurst (1988).

- - - - - Order Lumbriculida - - - - -

Family Lumbriculidae

    Genus Eclipidrilus Eisen, 1881
    * Eclipidrilus (Premnodrilus) daneus Cook, 1966
    * Eclipidrilus (Premnodrilus) lacustris (Verrill, 1871)
    Eclipidrilus (Premnodrilus) palustris (F. Smith, 1900)

    Genus Eremidrilus Fend & Rodriguez, 2003
    * Eremidrilus allegheniensis (Cook, 1971)

    Genus Lumbriculus Grube, 1844
    * Lumbriculus variegatus variegatus (Müller, 1774)

    Genus Stylodrilus Claparède, 1862
    * Stylodrilus heringianus Claparède, 1862 - i -
    * Stylodrilus wahkeenensis Rodriguez & Coates, 1996
- - - - - Order Tubificida - - - - -

Family Naididae (=Naidina) Ehrenberg, 1828 1,2

    Subfamily Naidinae 1,2

    Genus Amphichaeta Tauber, 1879
    Amphichaeta americana Chen, 1944
    * Amphichaeta leydigi Tauber, 1879

    Genus Chaetogaster von Baer, 1827
    Chaetogaster diaphanus (Gruithuisen, 1828) 9
    Chaetogaster diastrophus (Gruithuisen, 1828)
    Chaetogaster limnaei von Baer, 1827

    Genus Paranais Czerniavsky, 1880
    Paranais frici Hrabe, 1941
    * Paranais grandis Harman, 1977
    Paranais litoralis (Müller, 1784)

    Genus Allonais Sperber, 1948 3
    Allonais inaequalis (Stephenson, 1911) 3
    ?? [Allonais paraguayensis (Michaelsen, 1905)] 3
    ?? Allonais pectinata (Stephenson, 1910) 3

    Genus Arcteonais Piguet, 1928
    Arcteonais lomondi (Martin, 1907)

    Genus Bratislavia Košel, 1976
    Bratislavia bilongata (Chen, 1944)
    Bratislavia dadayi (Michaelsen, 1905) [= Bratislavia unidentata (Harman, 1973) -- See Righi and Hamoui (2002)]

    Genus Dero Oken, 1815
    Dero (Allodero) hylae Goodchild, 1951
    Dero (Allodero) floridana Harman, 1971
    Dero (Dero) abranchiata Harman, 1977
    Dero (Dero) botrytis Marcus, 1943
    Dero (Dero) digitata (Müller, 1773)
    Dero (Dero) nivea Aiyer, 1930
    Dero (Dero) obtusa d'Udekem, 1855
    Dero (Dero) pectinata Aiyer, 1930
    Dero (Dero) trifida Loden, 1979
    Dero (Aulophorus) borellii Michaelsen, 1900
    Dero (Aulophorus) flabelliger (Stephenson, 1931)
    Dero (Aulophorus) furcata (Müller, 1773)
    Dero (Aulophorus) lodeni Brinkhurst, 1986
    Dero (Aulophorus) superterrenus Michaelsen, 1912 12
    Dero (Aulophorus) vaga (Leidy, 1880)

    Genus Haemonais Bretscher, 1900
    Haemonais waldvogeli Bretscher, 1900

    Genus Homochaeta Bretscher, 1896
    {{ ? Homochaeta naidina Bretscher, 1896}} 4

    Genus Nais Müller, 1773
    Nais barbata Müller, 1773
    Nais behningi Michaelsen, 1923
    Nais bretscheri Michaelsen, 1899
    Nais communis Piguet, 1906
    Nais elinguis Müller, 1773
    Nais pardalis Piguet, 1906
    Nais pseudobtusa Piguet, 1906
    Nais simplex Piguet, 1906
    Nais variabilis Piguet, 1906 10

    Genus Ophidonais Gervais, 1838
    Ophidonais serpentina (Müller, 1773)

    Genus Ripistes Dujardin, 1842
    $? Ripistes parasita (Schmidt, 1874) - i -

    Genus Slavina Vejdovský, 1883
    Slavina appendiculata (d'Udekem, 1855)

    Genus Specaria Sperber, 1939
    Specaria josinae (Vejdovský, 1883)

    Genus Stephensoniana Cernosvitov, 1938
    Stephensoniana tandyi Harman, 1975
    Stephensoniana trivandrana (Aiyer, 1926)

    Genus Stylaria Lamark, 1816
    Stylaria lacustris (Linnaeus, 1767)
    [= Stylaria fossularis Leidy, 1852]

    Genus Uncinais Levinsen, 1884
    * Uncinais uncinata (Ørsted, 1842)

    Genus Vejdovskyella Michaelsen, 1903
    Vejdovskyella comata (Vejdovský, 1883)

    Genus Stephensoniana Cernosvitov, 1938
    Stephensoniana tandyi Harman, 1975
    Stephensoniana trivandrana (Aiyer, 1926)

    Subfamily Pristininae 1

    Genus Pristina Ehrenberg, 1828
    Pristina aequiseta Bourne, 1891
    Pristina breviseta Bourne, 1891
    Pristina jenkinae (Stephenson, 1931) 5
    Pristina leidyi F. Smith, 1896
    * Pristina longidentata Harman, 1965 5
    Pristina longiseta Ehrenberg, 1828
    Pristina longisoma Harman, 1977 5
    Pristina osborni (Walton, 1906) 5
    * Pristina plumaseta Turner, 1935
    $? Pristina proboscidea Beddard, 1896 14
    Pristina sima (Marcus, 1944) 5
    Pristina synclites Stephenson, 1925

    Genus Pristinella Brinkhurst, 1985 5
    this genus was suppressed by Collado and Schmelz (2000) 5

    Subfamily Tubificinae 1

    Genus Aulodrilus Bretscher, 1899
    Aulodrilus americanus Brinkhurst & Cook, 1966
    Aulodrilus limnobius Bretscher, 1899
    Aulodrilus pigueti Kowalewski, 1914
    Aulodrilus pluriseta (Piguet, 1906) /-/ Aulodrilus japonicus Yamaguchi, 1953 8

    Genus Haber Holmquist, 1978
    Haber speciosus (Hrabe, 1931)

    Genus Ilyodrilus Eisen, 1879
    Ilyodrilus templetoni (Southern, 1909)

    Genus Isochaetides Hrabe, 1966
    * Isochaetides curvisetosus (Brinkhurst & Cook, 1966)
    * Isochaetides freyi (Brinkhurst, 1965)

    Genus Limnodrilus Claparède, 1862
    * Limnodrilus cervix Brinkhurst, 1963
    * Limnodrilus claparedeianus Ratzel, 1868
    Limnodrilus hoffmeisteri Claparède, 1862
    * Limnodrilus profundicola (Verrill, 1871)
    * Limnodrilus rubripenis Loden, 1977
    * Limnodrilus udekemianus Claparède, 1862

    Genus Potamothrix Vejdovský & Mrázek, 1902
    ?? Potamothrix hammoniensis (Michaelsen, 1901) - i - 7
    Potamothrix vejdovskyi (Hrabe, 1941) - i -

    Genus Psammoryctides Hrabe, 1964
    Psammoryctides californianus Brinkhurst, 1965
    Psammoryctides convolutus Loden, 1978

    Genus Quistadrilus Brinkhurst, 1981
    Quistadrilus multisetosus (F. Smith, 1900)
    Genus Spirosperma Eisen, 1879
    * Spirosperma carolinensis Brinkhurst, 1965
    Spirosperma ferox Eisen, 1879 - i -
    Spirosperma nikolskyi (Lastockin & Sokolskaya, 1935)

    Genus Tasserkidrilus Holmquist, 1985
    {Tasserkidrilus harmani (Loden, 1979)} 11

    Genus Tubifex Lamark, 1816
    Tubifex ignotus (Stolc, 1886)
    Tubifex nerthus Michaelsen, 1908
    Tubifex tubifex (Müller, 1774)

    Genus Varichaetadrilus Brinkhurst & Kathman, 1983
    * Varichaetadrilus angustipenis (Brinkhurst & Cook, 1966)
    * Varichaetadrilus fulleri Brinkhurst & Kathman, 1983
    Varichaetadrilus harmani (Loden, 1979) 11
    Varichaetadrilus psammophilus (Loden, 1977)15

    Subfamily Telmatodrilinae

      None yet reported from Florida

    Subfamily Rhyacodrilinae

    Genus Bothrioneurum Michaelsen, 1900
    * Bothrioneurum vejdovskyanum Stolc, 1888

    Genus Branchiura Beddard, 1892
    * Branchiura sowerbyi Beddard, 1892 - i -

    Genus Rhizodrilus F. Smith, 1900
    * Rhizodrilus lacteus F. Smith, 1900

    Genus Rhyacodrilus Bretscher, 1901
    Rhyacodrilus coccineus (Vejdovský, 1875)
    Rhyacodrilus sodalis (Eisen, 1879)
    * Rhyacodrilus subterraneus Hrabe, 1963

    Subfamily Phallodrilinae

      No freshwater species in Florida

    Family Opistocystidae Cernosvitov, 1936

      Genus Crustipellis Harman & Loden, 1978
      Crustipellis tribranchiata (Harman, 1970) 6 (a)

      Genus Trieminentia Harman & Loden, 1978
      Trieminentia corderoi (Harman, 1970) 6 (b)

      Genus Opistocysta Cernosvitov, 1936
      Opistocysta flagellum (Leidy, 1880) - species inquirenda 6 (c)

    Family Parvidrilidae Erséus, 1999
      None yet reported from Florida

- - - - - Order Enchytraeida - - - - -

Family Enchytraeidae

    Genus Barbidrilus Loden & Locy, 1980
    Barbidrilus paucisetus Loden & Locy, 1980

    Genus Cernosvitoviella Nielsen & Christensen, 1959
    Cernosvitoviella christenseni Dash, 1970
    Cernosvitoviella pusilla Nurminen, 1973

    Genus Cognettia Nielsen & Christensen, 1959
    Cognettia spp. -- in banks of rivers and in lakes, also upper intertidal or tidal areas

    Genus Fridericia Michaelsen, 1889
    Fridericia canadensis Dash, 1972

    Genus Guaranidrilus Cernosvitov, 1937
    Guaranidrilus oregonensis Coates & Diaz, 1988

    Genus Hemienchytraeus Cernosvitov, 1934
    Hemienchytraeus spp. -- possible in wet soils, in southern regions

    Genus Henlea Michaelsen, 1889
    Henlea urbanensis Welch, 1914

    Genus Lumbricillus Ørsted, 1844
    Lumbricillus arenarius (Michaelsen, 1889) sensu Knöllner, 1935
    Lumbricillus charae (Tynen, 1970)
    Lumbricillus franciscanus Eisen, 1904 -- incertae sedis
    Lumbricillus kalatdlitus Nurminen, 1970
    Lumbricillus rivalis (Levinsen, 1883) augm. Ditlevsen, 1904
    [= L. rutilus Welch, 1914]
    Lumbricillus santaeclarae Eisen, 1904

    Genus Marionina Michaelsen, 1890, in Pfeffer, 1890
    Marionina forbesae F. Smith & Welch, 1913 -- incertae sedis

    Genus Mesenchytraeus Eisen, 1878
    Mesenchytraeus armatus (Levinsen, 1884)
    Mesenchytraeus franciscanus Eisen, 1904
    Mesenchytraeus fuscus Eisen, 1904
    Mesenchytraeus fuscus inermis Eisen, 1904
    Mesenchytraeus gelidus Welch, 1916
    Mesenchytraeus kincaidi Eisen, 1904
    Mesenchytraeus pedatus Eisen, 1904
    Mesenchytraeus rithralis Healy & Fend, 2002

Family Propappidae

    Genus Propappus Michaelsen, 1905
    None yet reported from Florida


Subclass Metagynophora

- - - - Order Opisthopora - - - -
Suborder Lumbricina
Superfamily Sparganophiloidea

Family Sparganophilidae13

    Genus Sparganophilus Benham, 1892
    Sparganophilus eiseni Smith, 1895
    Sparganophilus gatesi Reynolds, 1980 - endemic to FL
    Sparganophilus meansi Reynolds, 1980
    Sparganophilus pearsei Reynolds, 1975
    Sparganophilus pearsei libertiensis Reynolds, 1980 - endemic to FL
    Sparganophilus pearsei sarasotae Reynolds, 1980 - endemic to FL
    Sparganophilus tennesseensis Reynolds, 1977
    Sparganophilus wilmae Reynolds, 1980 - endemic to FL


1 Current Status of the families Naididae and Tubificidae (Annelida, Clitellata, 'Oligochaeta')

Based on sequences of 18S rDNA and other molecular and morphological data, Erséus et al. (2002) concluded that the family Naididae is more correctly placed within a subfamily [Naidinae] of the Tubificidae. As the name Naididae (=Naidina Ehrenberg, 1828) is older than Tubificidae Vejdovský 1876, Drs. C. Erséus, L. Gustavsson, and R.O. Brinkhurst submitted an application to the International Commission on Zoological Nomenclature (ICZN), requesting the conservation of the family-group names Tubificidae and Naididae, and that Tubificidae take precedence because of the large number of species (~800) in the family compared to those within the Naididae (~175) (C. Erséus, pers. comm.; see also Erséus and Gustavsson 2002). This application (Erséus et al. 2005) was published in December 2005 in the 'Abstracts of Cases' section of the Bulletin of Zoological Nomenclature [Vol. 62, part 4]. One paper in support of this application (Timm 2006a) was published in March 2006 in the 'Comments' section of the Bulletin of Zoological Nomenclature [Vol. 63, part 1]. No other papers, either in support of or in opposition to the petition, were published by the ICZN.

On 1 September 2006, the Commission was invited to vote on this proposal; at the close of the voting period (1 December 2006), 10 affirmative and 18 negative votes had been recorded. Thus, the ICZN has voted against the proposal to give precedence to Tubificidae over Naididae, maintaining precedence for Naididae over Tubificidae; their decision (ICZN 2007) was published in March 2007 in the 'Opinions' section of the Bulletin of Zoological Nomenclature [Vol. 64, Part 1: Opinion # 2167]. In this same article, the Commission included the original references for the names placed on Official Lists by the above ruling.

The ICZN stated that priority rule must be followed in this case (per the Code: ICZN 1999 - Article 23). Strictly speaking (Erséus, pers. comm. to MJW, October 2007), the ICZN did rule in favor of the reverse of their (Erséus et al. 2005) petition. However, the Commission took no position on the 18S rDNA data, nor on the idea that the Naididae represent a derived branch within the Tubificidae.

In November 2007, Erséus, Wetzel, and Gustavsson submitted a paper for publication that summarized the above information (history of this issue, DNA evidence, supportive publications, ICZN petition, ICZN ruling, consequences, and recommendations). This paper was published (9 April 2008) by Erséus, Wetzel, and Gustavsson, in Zootaxa 1744: 66-68; a complete citation for this paper is provided in the literature cited section of this page, below. A downloadable PDF of this paper is available free (through open access) by clicking on the journal's name, immediately above in this paragraph.

In late November 2007, Erséus, Wetzel, and Gustavsson submitted a paper for publication in which we summarize the above information (history of this issue, DNA evidence, supportive publications, ICZN petition, ICZN ruling, consequences, and recommendations). This paper was subsequently published on 9 April 2008 in Zootaxa 1744: 66-68; a complete citation for this paper is provided in the literature cited section of this page, below. A downloadable PDF of this paper is available free (through open access) by clicking on the journal's name, immediately above in this paragraph.

The familial rank of Naididae is maintained, with the subfamilies of the Tubificidae now included within the Naididae.

We encourage oligochaetologists to accept this new hirearchy and classification (as presented in the list, above).


Erséus et al. (2002) also presented parsimonious trees (resulting from cladistic analyses) showing the genus Pristina to be completely separate from the rest of the Naididae, and thus suggesting polyphyly of the family. However, because no particular support (e.g., jackknife analysis) for polyphyly was presented in that paper, Erséus (pers. comm., 2003) suggested [at that time] it may be premature to conclude that Naididae was polyphyletic.

However, in recently published research by Envall et al. (2006) -- which included additional taxa and genes in their analyses -- substantive molecular support for polyphyly was established; they are now rather sure that Pristina represents a separate branch of the Tubificidae - one that has acquired asexual reproduction independently from the other naidids. Furthermore, Envall et al. concluded that most of the old naidids indeed make ONE good group (with the exception of Pristina); their tree corresponds most closely to Lastockin's (1921) division of Naididae into Naidinae and Pristininae (only two subfamilies), with no real support for other subfamilial classification (exclusive of the now-included 'Tubificidae'). Note: the year of publication for Lastockin's (1921) paper has erroneously been recorded as '1924' in numerous publications over the years].

2 On earlier versions of this website (< April 2007), and also presented in a workshop workbook (Wetzel et al. 2006), the above list of North American freshwater microdriles included an interim subfamilial ranking, with [at that time of posting on the website] all 'former' Naididae placed in sub-familial groupings. This interim ranking was suggested by Dr. Gustavsson (pers. comm. to Wetzel, August 2005), pending a published opinion by the ICZN on the application by Erséus et al. (2005), and/or until such time that the monophyly of these groups could be tested definitively. Interim groupings (< April 2007) were as follows: Chaetogastrinae --> Chaetogastrinae Group, Paranaidinae --> Paranaidinae Group, Pristininae --> Pristininae Group, and with those taxa in the pre-existing subfamily Naidinae --> Naidinae Group -- all four of these groups were ranked within the one subfamily Naidinae.

However, in response to and in acceptance of the ICZN ruling (ICZN 2007) on that petition, and in consideration of the results of Envall et al. (2006) [see footnote 1, above] these interim groupings have now been removed from this classification, and the subfamilies Chaetogastrinae and Paranaidinae have been suppressed.

3 Kathman and Wetzel (2003) published a reassessment of Allonais species previously reported in North America, and included a re-description of A. inaequalis. Based upon their examinations of newly collected material, type specimens deposited in the USNM-Smithsonian Institution, and additional material held in other collections, they determined that all previous identifications of Allonais paraguayensis specimens collected in North America were incorrect. We therefore recommend the re-examination of all specimens that have been identified as Allonais species.

4 Oligochaete 'detectives' Tarmo Timm and Rheinar Grimm published (Timm and Grimm 2005) an exhaustive review of the taxon Homochaeta naidina Bretscher, 1896; I paraphrase here the abstract from that paper: Homochaeta naidina has never been redescribed. No type material exists for this taxon, and all subsequent material identified as this taxon, when available for review [by Timm and Grimm], had been misidentified. The original description by Bretscher likely was based upon different, immature specimens in the families Naididae and Tubificidae [probably Uncinais uncinata (Ørsted, 1842) and Bothrioneurum vejdovskyanum Stolc, 1886, respectively]. Timm and Grimm suggested that H. naidina, although formerly considered a valid species, may not in fact exist in nature; furthermore, the other species in the genus Homochaeta, based on the type species Homochaeta naidina, are either synonyms of Aulodrilus limnobius, or synonyms of other valid naidids that, themselves, have descriptions based only on immature specimens -- and thus whose generic positions remain uncertain until they have been thoroughly redescribed. Therefore, Timm and Grimm have concluded that the genus Homochaeta -- and in particular the taxon H. naidina -- should not be included in the routine identification keys for oligochaetes. We thus have double-bracketed {{ }} this taxon in the list, above, so that users of the list can be enlightened to the recent review by Timm and Grimm (2005) [citation provided below]. A rather humorous observation / summation of this taxon was also presented by Timm and Grimm in their paper: "....the unambitious name of Homochaeta naidina, included in all identification keys, has become some sort of a dustbin for small, immature oligochaete specimens with trivial bifid chaetae in all segments beginning in [segment] II." Timm and Grimm concluded with the suggestion that Homochaeta naidina (=Paranais naidina) can be regarded as a ghost name circulating from one hydrobiological paper to another. In a subsequent paper highlighting some commonly missused names of aquatic oligochaetes, Timm (2005) stated that it was unfortunate that the nominal species Homochaeta naidina could not be invalidated on formal reasons, since no type specimens are available to support a proper invalidation; to designate a neotype from [material previously misidentified as H. naidina] would be obsurd.

5 These Pristina species, all without a proboscis, were placed in the genus Pristinella by Brinkhurst (1985a) to separate them from other Pristina species with a proboscis. This separation was later suppressed by Collado and Schmelz (2000) based upon their research.

6 (a) The correct year of publication for W.J. Harman paper in which these descriptions were first presented [and thus, the date to be associated with these descriptions] is 1970, not '1969' (which was a misdating on all papers in the fourth issue of that volume). Actual publication of this paper - and all other papers in that fourth issue - occurred on 28 January 1970; the editor of the journal, J.O. Corliss, clarified this situation in the subsequent volume (see Transactions of the American Microscopical Society, vol. 89, p. 347), stating the 1970 date was correct and should be used.

6 (b) In his guide to freshwater oligochaetes of Florida, Michael Milligan (1997: 19-21) included Trieminentia corderoi based upon observations made on immature specimens only; however, no specific information [e.g., collecting location(s), specimen repository], was included with the account for this species in his guide. Unfortunately, Mike passed away in November 2005, so it is assumed that the specimen(s) he observed are forever lost. To date, the inclusion of T. corderoi report in Milligan's guide remains the only one for this species in North America (Kathman and Brinkhurst 1998, Wetzel et al. 2009). This species has been reported from Uruguay and Costa Rica (Harman and Loden 1978), and from Argentina (Christoffersen, 2008).

6 (c) The taxon Opistocysta flagellum (Leidy, 1880) has long been considered a species inquirenda. As stated in Brinkhurst and Jamieson (1971: 642-643), "Both W.J. Harman and the author [R.O. Brinkhurst] have sought the type specimens in vain, and a search of the type locality has failed to yield fresh material. In view of the diversity of characters displayed by S. American material and material from the southern states of the U.S.A., it is no longer possible to accept this as a valid species. I [Brinkhurst] agree with Harman (1970) that the best course of action is to remove this contentious name from consideration in the current nomenclature of the family." As best we can determine, there have never been any published reports of this taxon in Florida; we have included this sp. inq. here as reference for those who may have seen this name in old publications or project reports.

7 Records of Potamothrix hammoniensis and Tubifex newaensis in North America are thought erroneous, but have been perpetuated in the literature (pers. comm., R.O. Brinkhurst to MJW, Feb. 2006); careful review of all specimens identified as P. hammoniensis and T. newaensis is thus warranted prior to considering them to be extant species in North America.

8 Aulodrilus pluriseta could easily be confused with Aulodrilus japonicus (A. Ohtaka, pers. comm. to S.V. Fend and M.J. Wetzel). This opinion based upon Ohtaka's observations of several Nearctic specimens from the western U.S., and specimens of these taxa from Europe and Japan; see comments in Kathman and Brinkhurst (1998, p. 184). Dr. Ohtaka is currently preparing a paper with redescriptions of both species (pers. comm., May 2007).

9 Chaetogaster cristallinus could easily be confused with Chaetogaster diaphanus. Ch. cristallinus has a median prostomial incision (easily overlooked / unobservable due to orientation of mounted specimens), and {perhaps?} smaller / shorter chaetae than Ch. diaphanus; see comments in Kathman and Brinkhurst (1998: p. 42).
Hiltunen and Klemm (1980: pp. 7, 11, 33) included Chaetogaster cristallinus in their guide to North American Naididae. In the 'Annotations' section of Hiltunen and Klemm (1985: p. 39), the authors noted that - although Kasprzak (1972) synonymized Ch. cristallinus with Ch. diaphanus - further investigation is needed to substantiate this synonymy.
Thus, pending the publication of information to the contrary, we maintain the validity of Ch. cristallinus.

10 Several specimens initially identified as Nais variabilis (collected from the western U.S.) were later identified as Nais christinae (C. Erséus, pers. comm. to S.V. Fend) - a species not previously reported from North America. Certainly, additional morphological studies, in conjunction with gene sequencing, will be necessary to resolve the 'plasticity' and intergrades of many aquatic oligochaetes, including these (and other) Nais taxa.

11 Tasserkidrilus harmani was transferred to Varichaetadrilus by Timm (2006b; citation provided, below).

12 Dero (Aulophorus) superterrenus Michaelsen, 1912 -- a Neotropical species -- was newly added to this list in April 2008; however, it has long been known to occur in Florida, having been reported at least as far back as 1976 (as unidentified) by Fish (1976). Fish later sent some of his specimens to Jarl K. Hiltunen for identification, and those records were then noted in Frank and Lounibos (1987). Milligan (1997, p. 80) noted an unconfirmed record (as reported in Hiltunen and Klemm, 1985, p. 40) of D. superterrenus in North America. Milligan also mentioned the possibility of its occurrence in Florida, given that the state shares a rich naidid community with tropical South America [neither Milligan nor Wetzel were aware of the paper by Frank and Lounibos]. Just recently, Frank and Fish (2008) published a paper in which they discuss the potential loss of phylotelmata in Florida bromeliads due to destruction of host plants by an invasive dryophthorid beetle; in their paper, they note the observations of Lopez et al. (2005) of Dero superterrenus, which crawls onto the skin of frogs and thus using them as a mode of dispersal. Other unpublished records and specimens of D. superterrenus occurring in state were shared with me by my floridian colleagues Dr. John Epler (in 1999; specimens collected from phytotelmata of terrestrial bromeliads [cf. Billbergia] at the Riverwoods Field Laboratory, near Lorida, on the Kissimmee River), and Doug Strom [Water & Air Research, Inc.] (in March 2008; specimens were brought to his attention by a coworker, Efrain Tavarez, who obtained specimens from a bucket used as a cistern to collect / retain rainwater runoff from a roof in Melrose, Alachua Co., FL). Several specimens recently sent to me by Doug Strom are being mounted on slides, while other specimens were given to Dr. Christer Erséus, Göteborg University, Sweden, for gene sequencing. Dero superterrenus have been collected from bromeliads, tree holes, water basins, cisterns, and other epiphytes in Costa Rica, South America (Argentina and Brazil), from other areas in the Dutch West Indies, and from Malacca (Maylasia). [Previous exclusion of Dero superterrenus from this list, and from the species list on the North American oligochaete website, was an inadvertent oversight by MJW]. Wetzel is now working with Larry Hribar to identify oligochaetes collected from a variety of natural and anthropogenic phytotelmata in Monroe County, Florida ['Conch Republic']; we anticipate several specimens in these collections to be identified as D. superterrenus.

13 The megadrile ('earthworm') family Sparganophilidae is represented in North America by one genus, 14 species, and two subspecies. Sparganophilus eiseni Smith, 1895, is the only species in this family that is widespread in distribution (most states east of the Mississippi River, as well as Arkansas, Iowa, Missouri, Oklahoma, and Texas, and the eastern Canadian provinces of New Brunswick, Ontario, and Quebec). The rest of the taxa in this genus are restricted to five or fewer states; of the eight Sparganophilus species known to occur in Florida, four are (to date) endemic to the state. All species in this genus are limicolous, or mud-loving -- commonly found in wet soils, mud, muck and other moist substrates adjacent to streams, ponds, and lakes, and substrates at the bottom of these environs, up to a meter in water depth (Reynolds 2008). Additional information on this family can be found in Reynolds (2001, 2008) and Reynolds and Wetzel (2008).

14 As stated in Kathman & Brinkhurst (1998, p. 58), Pristina proboscidea Beddard, 1896, was included in the regional key for aquatic oligochaetes occurring in the Carolinas (Whitley, 1982), and later from Florida (Milligan, 1997); unfortunately, no specimens are available (lost, discarded) to corroborate these reports, and to date, no other North American records of this species have been published. Recently, several specimens in the genus Pristina that fit the published description of P. proboscidea were collected from two locations in Florida and loaned to M.J. Wetzel for examination. This information will be published in 2011.

15 The taxon Limnodrilus psammophilus Loden, 1977 was presented under this original name in Stimpson et al. (1985). Although Brinkhurst (1986), Kathman and Brinkhurst (1998), and Brinkhurst (1989) subsequently presented this taxon as Varichaetadrilus psammophilus (Loden, 1977) -- none of these three publications included reference to, discussion of, or formal explanation or justification for this new combination. Similar comment on this unsupported new combination was noted in Timm (2006b, pp.29-30).

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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. 2012. Terrestrial Oligochaeta (Annelida: Clitellata) in North America, including Mexico, Puerto Rico, Hawaii, and Bermuda. III. Megadrilogica 15(8): 191-211.

Reynolds, J.W., and M.J. Wetzel. in prep. Nomenclatura oligochaetologica. A catalogue of names, descriptions and type specimens of the Oligochaeta. Second Edition.

Righi, G., and V. Hamoui. 2002. Oligochaeta, Naididae of the West Indies and adjacent regions. Papéis Avulsos de Zoologia, S. Paulo 42(6): 119-167.

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Spencer, D.R., and R.L. Denton. 2003. Aquatic Oligochaeta (Annelida: Lumbriculidae, Haplotaxidae, Naididae, Tubificidae) of Utah. Western North American Naturalist 63(3):343-352.

Spencer, D.R., and P.L. Hudson. 2003. The Oligochaeta (Annelida, Clitellata) of the St. Lawrence Great Lakes region: an update. Journal of Great Lakes Research 29(1):89-104.

Spencer, D.R., and R.E. Wisseman. 1993. Some new records of Naididae and Tubificidae (Annelida: Oligochaeta) from Washington. Great Basin Naturalist 53(4):395-401.

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Stimpson, K.S., D.J. Klemm, and J.K. Hiltunen. 1982. A guide to the freshwater Tubificidae (Annelida: Clitellata: Oligochaeta) of North America. EPA-600/3-82-033 U.S. Environmental Protection Agency, Environmental Monitoring & Support Laboratory, Cincinnati, OH. x + 61 pp.

Stimpson, K.S., D.J. Klemm, and J K. Hiltunen. 1985. Freshwater Tubificidae (Annelida: Oligochaeta). Pages 44-69, In: D.J. Klemm (ed). A guide to the freshwater Annelida (Polychaeta, naidid and tubificid Oligochaeta, and Hirudinea) of North America. Kendall/Hunt Publishing Company, Dubuque, IA.

Strayer, D. 1990. Aquatic Oligochaeta. Pages 373-397, In: B.L. Peckarsky, P.R. Fraissinet, M.A. Penton, and D.J. Conklin, Jr. (eds). Freshwater macroinvertebrates of northeastern North America. Comstock Publishing Associates, Ithaca, NY. . xi + 442 pp. Paper softcover. ISBN 0-8014-9688-8. [available from Cornell University Press; To order, tel. 607.277-2211. USD$36.95]

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Wetzel, M.J. 1982. 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.

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Wetzel, M.J. 2008. In memoriam: Charlie D. Drewes (1946-2005). Pp. 10-17, In: Wang , H.-Z. et al. (eds.). Aquatic Oligochaete Biology X. Proceedings of the 10th International Symposium on Aquatic Oligochaeta. Acta Hydrobiologica Sinica 31 (suppl.).

Wetzel, M.J. 2008. In memoriam: Michael Robert Milligan (1951-2005). Pp. 18-20, In: Wang , H.-Z. et al. (eds.). Aquatic Oligochaete Biology X. Proceedings of the 10th International Symposium on Aquatic Oligochaeta. Acta Hydrobiologica Sinica 31 (suppl.).

Wetzel, M.J., S.V. Fend, K.A. Coates, R.D. Kathman, and S.R. Gelder. 2006. Taxonomy, systematics, and ecology of the aquatic Oligochaeta and Branchiobdellidae (Annelida, Clitellata) of North America, with emphasis on the fauna occurring in Florida. A workbook. 1 August. vi + 269 pp. + 4 color plates. [this workbook was prepared for / used during two workshops presented by the authors in 2006: 1) underwritten by / covened at the Florida Department of Environmental Protection, Tallahassee, 21-23 February, and 2) underwritten by the Florida Association of Benthologists, convened at the University of Florida Department of Entomology and Nematology, 9-11 May. Note: the 2006 versions of this workbook are now out of print; it has been superseded by the workbook cited below.

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. vi + 280 pp. + 4 color plates. Copyright (c) 2009. {The overall scope of this workbook is the North American continent; this serves as a compliment to the most recent guide published by R.D. Kathman and R. Brinkhurst in 1998.}

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. Available in PDF, at: HERE.

Whitley, L.S. 1982. Aquatic Oligochaeta. Pages 2.1-2.29, In: A. R. Brigham, W. U. Brigham and A. Gnilka (eds). Aquatic insects and oligochaetes of North and South Carolina. Midwest Aquatic Enterprises, Mahomet, Illinois. [chapters independently numbered]. ISBN 0-936416-00-9. [to order, contact Aleta Holt via telephone, at (217) 897-1274; cost: USD$43.00 (postage included; discount on multiple copy purchases)].

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Suggested citation for this electronic web page: Wetzel, M.J., R.D. Kathman, S.V. Fend, and K.A. Coates. 2015. Classification and Checklist of the Freshwater Oligochaetes Occurring in the State of Florida, USA. 3 February. World Wide Web URL: < >.

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