Classification and Checklist of Freshwater Oligochaetes (Annelida: Clitellata) in North America North of Mexico

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

[ Copyright © 2005–2024; all rights reserved]
[page update: 20Nov2022; 01,21Jan; 20,22Feb; 04,09Mar;16,21Apr; 15Aug; 25oct; 23nov2023; 29jan, 2Feb, 17Mar 2024; mjw]

This current list of freshwater oligochaetes occurring in North America north of Mexico is being presented here as an interim step towards a major revision and update of information currently available for this group via the Integrated Taxonomic Information System [ITIS] website. For several years, I [MJW] have been serving as the steward for the freshwater oligochaete information being presented by ITIS. In 2011, I began working more closely with ITIS personnel Jenny K. Archibald, Craig C. Freeman, G. Stinger Guala, and David Nicolson (ITIS Data Development Coordinator) to assist them in updating their databases and web interface with a current list of freshwater oligochaetes occurring in North America north of Mexico. Specifically, this project for ITIS focused on correcting nomenclatural discrepancies, incorporating recent synonymies, and providing supportive references for all valid taxa. An extensive dataset was submitted to ITIS in 2014, with periodic updates and corrections integrated during 2015, 2016, and 2017.

The Taxonomic Certification Program [TCP] underwritten by the Society for Freshwater Science [SFS] (formerly, the North American Benthological Society [NABS], 1975-2011) will be using the ITIS database as its official source of current nomenclature for most groups of freshwater macroinvertebrate taxa associated with their taxonomic certification tests.  As noted above, I [MJW] have been working with ITIS personnel to correct and update their list of freshwater oligochaetes.  The list presented on this website, below, shall serve as the source of accepted names for species included on upcoming TCP tests. Please visit the SFS Taxonomic Certification Program website for additional information.

While the classification and checklist presented below is limited in scope to the freshwater oligochaetes occurring in North America north of Mexico, we plan to expand this list (perhaps in a tabular format) to include the marine, estuarine, and brackish-water oligochaetes occurring in all of continental North America, then later add aeolosomatidans (a group now aligned with the Polychaeta).  In 2008, Don Klemm, Bill Moser, and Mark Wetzel established a website entitled Classification and Checklist of the Leeches (Phylum Annelida: Class Clitellata: Subclass Hirudinida) occurring in North America North of Mexico. In 2018, Fred Govedich agreed to assist with this website.

Gelder et al. (2002) presented a listing of North American species of branchiobdellidans, an update on classification and nomenclature for this group.

Because several megadrile oligochaetes are commonly found in aquatic, semi-aquatic, and limicolous habitats, those species have been included in the listing, below.

Reynolds and Wetzel (2004) produced the first comprehensive publication on earthworm distribution in North America since Gates (1942). Although Gates did include the distributional records for earthworms occurring in Central America and the Caribbean, his checklist only reported their occurrence by country, without further geographic breakdown. The checklist in Reynolds and Wetzel (2004) included the province, state and territorial distributions for earthworms based upon published records of species occurring in Canada and the continental United States. Additionally, many topics related to earthworms were covered, viz.: global biogeography, native and introduced species, unique distributions and habitats, barriers to migration, habitat requirements, field and laboratory methodologies used in the study of megadriles (e.g., collecting, preservation, identification, and deposition of specimens), and nomenclature.

In their update four years later, Reynolds and Wetzel (2008) presented known distributions for earthworm species occurring in continental North America based upon records published since Reynolds and Wetzel (2004) and in addition, broadened their coverage by integrating distributional records for species occurring in Mexico, the Commonwealth of Puerto Rico, Hawaii, and Bermuda that had been published to date. That paper also included distributional maps for all species, and an index for all species, genera, and families discussed therein.

Reynolds and Wetzel (2012) compiled a second update of known distributions for earthworm species occurring in North America, including Mexico, the Commonwealth of Puerto Rico, Hawaii, and Bermuda.  Specifically, this 2012 update highlighted distributional records for species that had been published since Reynolds and Wetzel (2008), and included distributional maps for all new species records.  A fourth update is being prepared for publication.

Reynolds and Wetzel (2011) published an historical perspective and the first update of earthworms in Illinois in over 50 years; three new records for the state were reported, increasing the number of species in the state to 38 (18 considered native and 20 considered to have been introduced). Several earthworms considered limicolous (mud-loving) and others commonly found in aquatic habitats were discussed.  A list of Illinois earthworms is presented here.  A second paper, to include numerous historical as well as recent records of earthworms occurring in Illinois, is now in preparation.

Reynolds and Wetzel (2014) published the first comprehensive paper on earthworms occurring in Michigan, USA, which includes a few species that are commonly collected from aquatic and semiaquatic habitats.

Will Reeves, John Reynolds, and MJW are now completing a paper on the limicolous family Sparganophilidae in North America, with publication scheduled for late 2023 or early 2024.

The classification and checklist presented below 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, Patrick Martin, and numerous other oligochaetologists working in North America and elsewhere in the World.

Much of the recent distributional information for oligochaetes in North America has been summarized during our affiliation with the American Fisheries Society AFS) Committee on Names of Aquatic Invertebrates (CNAI). The primary objective of the CNAI, established on 30 September 1981 by AFS president John J. Magnuson, was to achieve uniformity and confusion in the vernacular [and scientific] nomenclature of aquatic invertebrates. Under the direction of Dr. Donna Turgeon, a series of volumes in the AFS Special Publications series [SP] focusing on invertebrate groups (by phylum) was begun; to date, several volumes have been published: SP 16 (Mollusks – Turgeon et al. 1988), SP 17 (Decapod Crustaceans – Williams et al. 1989), SP 22 (Cnidaria and Ctenophora – Cairns et al. 1991), SP 26 (Mollusks, Second Edition – Turgeon et al. 1998), SP 28 (Cnidaria and Ctenophora, Second Edition – Cairns et al. 2002), and SP 31 (Crustaceans, Second Edition – McLaughlin et al. 2005). These are available for purchase via this AFS website. Coates and Wetzel  served on this Committee (1989–2006), and during that time (in collaboration with many colleagues) had been preparing the Annelida volume for this series. The retirement of Dr. Turgeon several years ago as well as the restructuring of priorities and active administrative oversight by AFS resulted in the ending of the AFS CNAI Program.

We thank our annelidically inclined colleagues both in North America and elsewhere in the world for their contributions to our science, and for providing us with reprints and other information summarizing their research, thus allowing us to remain relatively current with the taxonomy, systematics, and ecology of oligochaetes and other annelid groups. However, we also recognize that—due the very nature of the science of taxonomy and systematics—virtually the moment a work is published it begins the ‘drift towards obsolescence’, depending on how active the research is in a particular group. An uncanny case in point can be found by reading footnote ‘4’ at the bottom of the checklist presented here; in mid-November 2005, the day I (MJW) had finalized the first version of this html file for posting live on our server, a reprint from two of our long-time colleagues, Tarmo Timm and Reinmar Grimm, arrived in the mail. Their paper (Timm & Grimm 2005) suppressed (perhaps forever) the lingering name of a naidine (Homochaeta naidina) for which there are no actual specimens (types or otherwise)—just a lot of mis-identified worms deposited in a lot of different institutions and museums and reported in a lot of diverse publications—and representing (based upon exhaustive re-evaluation of these specimens by Timm and Grimm) taxa in several families. I encourage you to read the paper by Timm and Grimm, and all footnotes, below; each lends pragmatic reality to the world of oligochaete taxonomy and systematics.

In addition, we thank our dear friend and colleague, Tarmo Timm (Estonian Institute of Life Sciences, Centre for Limnology, Rannu, Tartumaa, Estonia), for his assistance with all aspects of this list; Tarmo is a tireless oligochaetologist, always providing editorial insight, oversight, invaluable critique and advice, and philosophical guidance for our research and our presentation of information.

Finally, our efforts to maintain ‘currency’ in this checklist is dependent upon collaboration between and corroboration among our fellow oligochaetologists —in Florida and elsewhwere; the information compiled and presented below would not have been possible without their extensive and unselfish efforts in the global partnership dedicated to the enhancement of expertise in taxonomy, systematics, phylogeny, and nomenclature (emphasizing here the objectives and accomplishments of the NSF PEET Program) [a recent visit (April 2023) to the PEET website discovered a notice at the top of its home page….that the program has been archived]

As of August 2023, ~270 nominal species of freshwater (microdrile) oligochaetes representing 84 genera in six families are known to occur in North America – inclusive of Canada, continental United States and Mexico, and Hawai’i, Bermuda, and Puerto Rico. Several megadrile oligochaetes (‘earthworms’) also occur in aquatic and semi-aquatic habitats, soon to be added to this web-based list.  We present here a general classification and checklist of these species based upon records published in the peer-reviewed literature. 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 oligochaetologists, and / or  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).

Pragmatic aphorism: ”Too many worms…..too little time.”

Source of information for species included in this checklist

The freshwater oligochaete fauna of North America, particularly the species in the family Naididae and former family 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, 1989), 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.  Additionally, locality information for recently described taxa in the list, below, can be accessed via the citations in the Literature Cited section of this website that are associated with the authority(ies) of those new 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, 2009a, 2009b; Fend and Carter 2014; Fend and Lenat 2007, 2010, 2012; Fend et al. 2015; Fend et al. 2016; and Rodriguez et al. 2014). While many of these papers are specific to newly described taxa with unique and usually limited distributions, information presented therein provides insight to taxa known or thought likely to occur elsewhere in North America. 


Nomenclatural changes and synonymies for North American species published since Kathman and Brinkhurst (1998, 1999) have been included herein; to the best of our knowledge, the nomenclature for species discussed herein is current as of 23 November 2023.

On Monday 6 January 2014, Mark J. Wetzel (INHS) and John W. Reynolds (Oligochaetology Lab) launched a new website presenting the second edition of Nomenclatura Oligochaetologica, as Nomenclatura Oligochaetologica – a catalogue of names, descriptions, and type specimens of the Oligochaeta, Editio Secunda. 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.


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 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]). 

Introduction to the checklist

The primary purpose of this webpage is to present an accurate list of freshwater oligochaetes occurring in North America (north of Mexico), 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.

Distribution records (state, province) for many species described since 1 January 2007 are included at the end of their listing.

The megadrile (‘earthworm’) family Sparganophilidae is represented in North America by one genus, 14 species, and two subspecies. Sparganophilus eiseni Smith, 1895—now considered a junior synonym of Sparganophilus tamesis Benham, 1892 [see Rota et al. 2016] 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; four occur only in Florida. All species in the genus Sparganophilus are limicolous, or mud-loving — most 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 (2004, 2008, 2012). As noted above, Will Reeves, John Reynolds, and MJW are now completing a paper on the limicolous family Sparganophilidae in North America.

Please read the information associated with footnotes 1 and 2, below, regarding the 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).

– 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 / based upon erroneous record [misidentification or lost specimen(s)] (see footnote).

Classification and checklist of the freshwater oligochaetes
(Annelida: Clitellata) occurring in North America North of Mexico

Phylum Annelida
Class Clitellata
 Michaelsen, 1919
Subclass Oligochaeta Grube, 1850

– – – – – 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 Vejdovský, 1884
Genus Altmanella Fend, 2009
Altmanella freidris (Cook, 1966) [ transferred from Kincaidiana by Fend (2009)]
Altmanella idahoensis Fend, 2009
Altmanella lenati Fend, 2009
Genus Cookidrilus Rodriguez & Giani, 1987
Cookidrilus pocosinus Rodriguez, Fend & Lenat, 2014
Genus Eclipidrilus Eisen, 1881
Eclipidrilus asymmetricus (F. Smith, 1896)
Eclipidrilus breviatriatus Fend & Lenat, 2012
Eclipidrilus (Premnodrilusdaneus Cook, 1966
Eclipidrilus (Leptodrilusfontanus Wassel, 1984
Eclipidrilus (Eclipidrilusfrigidus Eisen, 1881
Eclipidrilus (Leptodrilusithys Brinkhurst, 1998
Eclipidrilus (Leptodriluslacustris (Verrill, 1871)
Eclipidrilus macphersonae Fend & Lenat, 2012
Eclipidrilus microthecus Fend & Lenat, 2012
Eclipidrilus pacificus Fend, 2005
Eclipidrilus (Premnodriluspalustris (F. Smith, 1900)
Genus Eremidrilus Fend & Rodriguez, 2003
Eremidrilus allegheniensis (Cook, 1971)
Eremidrilus chalonensis Rodriquez & Fend, 2022

Eremidrilus coyote Fend & Rodriguez, 2003
Eremidrilus elegans Fend & Rodriguez, 2003
Eremidrilus felini Fend & Rodriguez, 2003
Eremidrilus owyhee Rodriquez & Fend, 2022
Eremidrilus parvitheca [typical specimens] Fend 2023; [distr.: U.S. – TX: Hays Co., Comal Co.]
Eremidrilus parvitheca {variant specimens} Fend 2023 [distr.: U.S. – TX: Comal Co., Bell Co.]
Eremidrilus pinedai Rodriquez & Fend, 2022
Eremidrilus ritocsi Fend & Rodriguez, 2003
Eremidrilus samacos Fend 2023 [distr.: U.S. – TX: Hays Co., Comal Co.]

Genus Kincaidiana Altman, 1936 [ diagnosis of this genus emended by Fend (2009) ]
     [ Kincaidiana freidris Cook, 1966] [ transferred to genus Altmanella by Fend (2009) ]
Kincaidiana hexatheca Altman, 1936  [ diagnosis emended by Fend (2009) ]
Genus Lumbriculus Grube, 1844
Lumbriculus ambiguus Holmquist, 1976
Lumbriculus genitosetosus Holmquist, 1976
Lumbriculus inconstans (F. Smith, 1895)
Lumbriculus variegatus variegatus (Müller, 1774)
Genus Martinidrilus Fend & Lenat, 2007
Martinidrilus arenosus Fend & Lenat, 2007 [ distr.: U.S. – NC ]
Martinidrilus carolinensis Fend & Lenat, 2007 [ distr.: U.S. – NC,VA ]
Genus Pararhynchelmis Fend & Lenat, 2010
Pararhynchelmis murdocki Fend & Lenat, 2010 [ distr.: U.S. – VA ]

Pararhynchelmis texana Fend 2023 [distr.: U.S. – TX: Hays Co.]

Genus Pilaridrilus Fend & Lenat, 2007
Pilaridrilus uliginosus Fend & Lenat, 2007 [ distr.: U.S. – NC ]
Genus Phagodrilus McKey-Fender, 1988
Phagodrilus phoebe McKey-Fender, 2001
Phagodrilus balchi McKey-Fender, 2001
Phagodrilus baueri McKey-Fender, 2001
Phagodrilus chetcoensis McKey-Fender, 2001
Phagodrilus hauserensis McKey-Fender, 2001
Phagodrilus johnsoni McKey-Fender, 2001
Phagodrilus klamathensis McKey-Fender, 2001
Phagodrilus laqueus McKey-Fender, 2001
Phagodrilus macnabi McKey-Fender, 1988
Phagodrilus minimus McKey-Fender, 2001
Phagodrilus oregonensis McKey-Fender, 2001
Phagodrilus parvus McKey-Fender, 2001
Phagodrilus perditus McKey-Fender, 2001
Phagodrilus secundus McKey-Fender, 2001
Phagodrilus stellatus McKey-Fender, 2001
Genus Rhynchelmis Hoffmeister, 1843
Rhynchelmis (Rhynchelmoidesalaskana Holmquist, 1976
Rhynchelmis (Sutroa) aleutensis Fend, 2005
Rhynchelmis brooksi Holmquist, 1976
Rhynchelmis (Sutroadiespluviae Fend, 2014
Rhynchelmis (Rhynchelmoides) elrodi Smith & Dickey, 1918
Rhynchelmis (Sutroa) gilensis Fend & Brinkhurst, 2000
Rhynchelmis (Rhynchelmoidesglandula Altman, 1936
Rhynchelmis (Sutroa) gustafsoni Fend & Brinkhurst, 2000
Rhynchelmis (Sutroaklamathensis Fend, 2014
Rhynchelmis (Sutroa) monsserratus Fend & Brinkhurst, 2000
Rhynchelmis rostrata (Eisen, 1888)
Rhynchelmis (Rhynchelmoidessaxosa Fend & Brinkhurst, 2000
Rhynchelmis (Sutroa) utahensis Fend & Brinkhurst, 2000
Rhynchelmis (Sutroa) yakimorum Fend & Brinkhurst, 2000
Rhynchelmis (Rhynchelmoidesbolinensis Fend & Lenat, 2010 [ distr.: U.S. – NC]
Rhynchelmis (Rhynchelmoidescroatanensis Fend & Lenat, 2010 [ distr.: U.S. – NC ]
Genus Secubelmis Fend & Gustafson, 2001
Secubelmis limpida Fend & Gustafson, 2001
Genus Spelaedrilus Cook, 1975
Spelaedrilus multiporus Cook, 1975
Genus Stylodrilus Claparède, 1862
Stylodrilus beattiei Cook, 1975
Stylodrilus californianus Rodriguez, 1996
Stylodrilus coreyi Rodriguez, Fend & Lenat, 2014
Stylodrilus heringianus Claparède, 1862 – i –
Stylodrilus sovaliki Holmquist, 1976
Stylodrilus wahkeenensis Rodriguez & Coates, 1996
Genus Styloscolex Michaelsen, 1901
Styloscolex opisthothecus Sokolskaya, 1969
Genus Sylphella Rodriguez, Fend & Lenat, 2014
Sylphella puccoon Rodriguez, Fend & Lenat, 2014
Genus Tenagodrilus Eckroth & Brinkhurst, 1996
Tenagodrilus musculus Eckroth & Brinkhurst, 1996
Genus Trichodrilus Claparède, 1862
Trichodrilus allobrogum Claparède, 1862

Trichodrilus comalensis Fend, 2023 [distr.: U.S. – TX: Comal Co., Hays Co.]
Trichodrilus culveri Cook, 1975

Genus Utkena Fend, Rodriguez & Lenat, 2015
Utkena riparia Fend, Rodriguez & Lenat, 2015 [ distr.: U.S. – NC]


– – – – – Order Tubificida – – – – –

Family Naididae (=Naidina) Ehrenberg, 1828 1,2,14
Subfamily Naidinae Ehrenberg, 1828 1,2
Genus Branchiodrilus Michaelsen, 1900
          Branchiodrilus cf. hortensis (Stephenson, 1910) 21
Genus Chaetogaster von Baer, 1827 25 
Chaetogaster cristallinus Vejdovský, 1883 9
Chaetogaster diaphanus (Gruithuisen, 1828) 9
Chaetogaster diastrophus (Gruithuisen, 1828)
Chaetogaster limnaei von Baer, 1827
Chaetogaster setosus Svetlov, 1925
Genus Amphichaeta Tauber, 1879
Amphichaeta americana Chen, 1944
Amphichaeta leydigi Tauber, 1879
Amphichaeta raptisae (Chapman, 1981)
Genus Homochaeta Bretscher, 1896
{{ ? Homochaeta naidina Bretscher, 1896}} 4
Genus Ophidonais Gervais, 1838
Ophidonais serpentina (Müller, 1773) 22 
Genus Uncinais Levinsen, 1884
Uncinais uncinata (Ørsted, 1842)
Genus Piguetiella Sperber, 1939
Piguetiella blanci (Piguet, 1906)
Piguetiella michiganensis Hiltunen, 1967
Genus Specaria Sperber, 1939
Specaria fraseri Brinkhurst, 1978
Specaria hellei (Brinkhurst, 1971)
Specaria josinae (Vejdovský, 1883)
Genus Ripistes Dujardin, 1842
Ripistes parasita (Schmidt, 1874)  i – 11
Genus Stylaria Lamark, 1816
Stylaria lacustris (Linnaeus, 1767)
Stylaria fossularis Leidy, 1852
Genus Arcteonais Piguet, 1928
Arcteonais lomondi (Martin, 1907)
Genus Haemonais Bretscher, 1900
Haemonais waldvogeli Bretscher, 1900
Genus Dero Oken, 1815 24 
Dero (Allodero) hylae Goodchild, 1951 23
Dero (Allodero) floridana Harman, 1971 23
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, 1774)
Dero (Aulophorus) lodeni Brinkhurst, 1986
Dero (Aulophorus) superterrenus (Michaelsen, 1912) 12
Dero (Aulophorus) vaga (Leidy, 1880)
Genus Allonais Sperber, 1948 3
Allonais inaequalis (Stephenson, 1911) 3
   ?? [Allonais paraguayensis (Michaelsen, 1905)] 3
   ?? Allonais pectinata (Stephenson, 1910) 3
Genus Slavina Vejdovský, 1883
Slavina appendiculata (d’Udekem, 1855)
     note that another species in this genus, Slavina evelinae (previously known from
      Central and South America, has been identified by MJW based on specimens
      (collected in Florida) that were received from Dr. J.H. Epler; a paper discussing
      the global distribution of known Slavina species is in prep. 
Genus Vejdovskyella Michaelsen, 1903 26
Vejdovskyella comata (Vejdovský, 1883)
Vejdovskyella intermedia (Bretscher, 1896)
Genus Nais Müller, 1773
Nais alpina Sperber, 1948
Nais barbata Müller, 1773
Nais behningi Michaelsen, 1923
Nais bicuspidalis Grimm & Fend, 1997
Nais bretscheri Michaelsen, 1899
Nais communis Piguet, 1906
Nais christinae Kasprzak, 1973 10
Nais elinguis Müller, 1773
Nais magnaseta Harman, 1973
Nais pardalis Piguet, 1906
Nais pseudobtusa Piguet, 1906
Nais simplex Piguet, 1906
Nais variabilis Piguet, 1906 10
Genus Stephensoniana Černosvitov, 1938
Stephensoniana tandyi Harman, 1975
Stephensoniana trivandrana (Aiyer, 1926)
Genus Bratislavia Košel, 1976
Bratislavia bilongata (Chen, 1944)
Bratislavia dadayi (Michaelsen, 1905)
[= Bratislavia unidentata (Harman, 1973) – See Righi and Hamoui (2002)]
Genus Paranais Czerniavsky, 1880
Paranais birsteini Sokolskaya, 1971
Paranais frici Hrabě, 1941
Paranais grandis Harman, 1977
Paranais litoralis (Müller, 1784)
Subfamily Pristininae Lastočkin, 1924 1
Genus Pristina Ehrenberg, 1828
Pristina acuminata Liang, 1958 5
Pristina aequiseta Bourne, 1891
Pristina americana Černosvitov, 1937
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 minuta (Stephenson, 1914) 5
Pristina notopora Černosvitov, 1937 5
Pristina osborni (Walton, 1906) 5
Pristina plumaseta Turner, 1935
? Pristina proboscidea Beddard, 1896
Pristina sima (Marcus, 1944) 5
Pristina synclites Stephenson, 1925
Genus Pristinella Brinkhurst, 1985 5
Note:  the genus Pristinella was suppressed by Collado and Schmelz (2000) 5

 Subfamily Tubificinae 1 Vejdovský, 1876
Genus Arctodrilus Brinkhurst & Kathman, 1983
Arctodrilus wulikensis Brinkhurst & Kathman, 1983
Genus Aulodrilus Bretscher, 1899
Aulodrilus americanus Brinkhurst & Cook, 1966
Aulodrilus limnobius Bretscher, 1899
Aulodrilus paucichaeta Brinkhurst & Barbour, 1985
Aulodrilus pigueti Kowalewski, 1914
Aulodrilus pluriseta (Piguet, 1906) /-/ Aulodrilus japonicus Yamaguchi, 1953 8
Genus Haber Holmquist, 1978
     Haber speciosus (Hrabě, 1931)
Genus Ilyodrilus Eisen, 1879
     Ilyodrilus frantzi Brinkhurst, 1965
     Ilyodrilus perrieri Eisen, 1879
     Ilyodrilus templetoni (Southern, 1909)
Genus Isochaetides Hrabě, 1966
     Isochaetides curvisetosus (Brinkhurst & Cook, 1966)
     Isochaetides freyi (Brinkhurst, 1965)
Genus Kopilrok Holmquist, 1985
     Kopilrok flexipenis Holmquist, 1985 – AK
     Kopilrok sagavanirktoki Holmquist, 1985 – AK
Genus Limnodrilus Claparède, 1862
Limnodrilus cervix Brinkhurst, 1963
Limnodrilus claparedeianus Ratzel, 1868
Limnodrilus hoffmeisteri Claparède, 1862 17
Limnodrilus maumeensis Brinkhurst & Cook, 1966
Limnodrilus profundicola (Verrill, 1871)
Limnodrilus rubripenis Loden, 1977
Limnodrilus silvani Eisen, 1879
Limnodrilus sulphurensis Fend, Liu & Erséus, 2016 [ distr.: U.S. – CO ]
Limnodrilus tortilipenis Wetzel, 1987
Limnodrilus udekemianus Claparède, 1862
Genus Potamothrix Vejdovský & Mrázek, 1902
Potamothrix bavaricus (Oschmann, 1913) 13 – i –
Potamothrix bedoti (Piguet, 1913) – i (?) –
   ?? Potamothrix hammoniensis (Michaelsen, 1901) 7– i –
Potamothrix moldaviensis Vejdovský & Mrázek, 1902 – i –
Potamothrix vejdovskyi (Hrabě, 1941) – i –
Genus Psammoryctides Hrabě, 1964
Psammoryctides barbatus (Grube, 1891)
Psammoryctides californianus Brinkhurst, 1965
Psammoryctides convolutus Loden, 1978
Genus Quistadrilus Brinkhurst, 1981
Quistadrilus multisetosus (F. Smith, 1900)
Genus Spirosperma Eisen, 1879
Spirosperma beetoni Brinkhurst, 1965
Spirosperma carolinensis Brinkhurst, 1965
Spirosperma ferox Eisen, 1879 – i –
Spirosperma nikolskyi (Lastočkin & Sokolskaya, 1935)
Genus Tasserkidrilus Holmquist, 1985
Tasserkidrilus americanus (Brinkhurst & Cook, 1966)
{Tasserkidrilus harmani (Loden, 1979)} 16
Tasserkidrilus superiorensis (Brinkhurst & Cook, 1966)
Genus Teneridrilus Holmquist, 1985
Teneridrilus calvus Erséus & Brinkhurst, 1990
Teneridrilus columbiensis (Brinkhurst & Diaz, 1985)
Teneridrilus flexus Erséus & Hiltunen, 1990
Teneridrilus mastix (Brinkhurst, 1978)
Genus Tubifex Lamarck, 1816
Tubifex fissidens Holmquist, 1983
Tubifex ignotus (Štolc, 1886)
Tubifex nerthus Michaelsen, 1908
?? Tubifex newaensis (Michaelsen, 1903) 7
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 minutus (Brinkhurst, 1965)
Varichaetadrilus pacificus (Brinkhurst, 1981)
Varichaetadrilus psammophilus (Loden, 1977) 15

Subfamily Telmatodrilinae Eisen, 1879

Genus Telmatodrilus Eisen, 1879
     Telmatodrilus vejdovskyi Eisen, 1879
Genus Alexandrovia Hrabě, 1962
     Alexandrovia onegensis Hrabě, 1962

Subfamily Rhyacodrilinae Hrabě, 1963
Genus Bothrioneurum Michaelsen, 1900
     Bothrioneurum vejdovskyanum Štolc, 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 alcyoneus Rodriguez & Fend, 2013 [ distr.: U.S. – CA ]
Rhyacodrilus brevidentatus Brinkhurst, 1965
Rhyacodrilus clio Rodriguez & Fend, 2013 [ distr.: U.S. – CA ]
Rhyacodrilus coccineus (Vejdovský, 1875)
Rhyacodrilus falciformis Bretscher, 1901
Rhyacodrilus longichaeta Rodriguez & Fend, 2013 [ distr.: U.S. – CA ]
Rhyacodrilus montana (Brinkhurst, 1965) [‘montanus’ is incorrect]
Rhyacodrilus propiporus Rodriguez & Fend, 2013 [ distr.: U.S. – NC ]
Rhyacodrilus saelonae Rodriguez & Fend, 2013 [ distr.: U.S. – CA ]
Rhyacodrilus sodalis (Eisen, 1879)
Rhyacodrilus subterraneus Hrabě, 1963
Rhyacodrilus quileuticus Rodriguez & Fend, 2013 [ distr.: U.S. – WA ]
Subfamily Phallodrilinae Brinkhurst, 1971
Genus Gianius Erséus, 1992
Gianius aquaedulcis (Hrabě, 1960)
Genus Thalassodrilus Brinkhurst, 1963
Thalassodrilus hallae (Cook & Hiltunen, 1975)
Subfamily Opistocystinae Černosvitov, 1936
     [formerly, Family Opistocystidae Černosvitov, 1936] 
            see Erséus, C., I. Envall, M. Marchese, and L.M. Gustavsson (2010), 
              and Erséus, C., I. Envall, P. De Wit, L.M. Gustavsson. 2017 [citations below]
Genus Crustipellis Harman & Loden, 1978
     Crustipellis tribranchiata (Harman, 1970) 6 (a)
Genus Trieminentia Harman & Loden, 1978
     Trieminentia corderoi (Harman, 1970) 6 (b)
Genus Opistocysta Černosvitov, 1936
     Opistocysta flagellum (Leidy, 1880) – species inquirenda 6 (c)

Family Parvidrilidae Erséus, 1999
Genus Parvidrilus Erséus, 1999
     Parvidrilus strayeri Erséus, 1999
For additional information on other species in this genus (those occurring outside of North America), please see: Martínez-Ansemil et al. (2002) and  Martínez-Ansemil et al. (2012).

– – – – – Order Enchytraeida – – – – –

This listing includes genera and species collected from terrestrial, limicolous, semi-aquatic, and aquatic habitats.

Family Enchytraeidae Vejdovský, 1879

 Genus Achaeta Vejdovský, 1878
     Achaeta spp.
Genus Barbidrilus Loden & Locy, 1980
     Barbidrilus paucisetus Loden & Locy, 1980
Genus Bryodrilus Ude, 1892
     Bryodrilus librus (Nielsen & Christensen, 1959) 20 WI
     Bryodrilus cf. diverticulatus Černosvitov, 1929 20 MN, WI
     Bryodrilus cf. ehlersi Ude, 1892 20 WI
     Bryodrilus spp. 20 MN
Genus Buchholzia Michaelsen, 1886
Buchholzia appendiculata (Buchholz, 1862) 20 MN
Buchholzia spp. 20 MN
Genus Cernosvitoviella Nielsen & Christensen, 1959
Cernosvitoviella christenseni Dash, 1970
Cernosvitoviella pusilla Nurminen, 1973
Genus Cognettia Nielsen & Christensen, 1959 18
Cognettia glandulosa (Michaelsen, 1888) 18  MN, WI
Cognettia sphagnetorum
(Vejdovský, 1878) 19  MI, FL? WA

Genus Enchytraeus Henle, 1837
Enchytraeus buchholzi sensu lato Vejdovský, 1879 20 MN, WI
Enchytraeus dichaetus Schmelz & Collado, 2010 20 MN
Enchytraeus spp. 20 MN


Genus Enchytronia Nielsen & Christensen, 1959
Enchytronia parva Nielsen & Christensen, 1959 20 WI

Genus Fridericia Michaelsen, 1889
Fridericia bulboides Nielsen & Christensen, 1959 20 MN, WI
Fridericia canadensis Dash, 1972
Fridericia cylindrica Springett, 1971 20 WI
Fridericia galba (Hoffmeister, 1843) 20 MN
Fridericia paroniana Ussel, 1904 20 MN
Fridericia ratzeli (Eisen, 1872) 20 MN
Fridericia sp. 1 (cf. agilis/agricola20 MN
Fridericia spp. 20 MN, WI
Genus Guaranidrilus Černosvitov, 1937
Guaranidrilus oregonensis Coates & Diaz, 1988
Genus Hemienchytraeus Černosvitov, 1934
Hemienchytraeus spp. — possible in wet soils, in southern regions
Genus Hemifridericia Nielsen & Christensen, 1959
Hemifridericia bivesiculata Christensen & Dózsa-Farkas, 1999 20 MN, WI
Hemifridericia parva Nielsen & Christensen, 1959 20 MN, WI
Hemifridericia spp.

Genus Henlea Michaelsen, 1889

Henlea conchifera Christensen & Dózsa-Farkas, 1999 20 MN
Henlea nasuta (Eisen, 1878) 20 MN
Henlea perpusilla Friend, 1911 20 MN, WI
Henlea ventriculosa (Udekem, 1854) 20 MN, WI
Henlea urbanensis Welch, 1914
Henlea spp. 20 MN, WI
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
Marionina sp. 20 MN, WI
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 rhithralis Healy & Fend, 2002
Genus Oconnorella Rota, 1995
Oconnorella spp. 20 MN, WI
Genus Stercutus Michaelsen, 1888
Stercutus niveus Michaelsen, 1888 20 MN, WI

Family Propappidae Coates, 1986
Genus Propappus Michaelsen, 1905
Propappus volki Michaelsen, 1922 – i –

Although the vast majority of oligochaetes in this order are terrestrial, several families include taxa (listed below) that are often collected from aquatic, semiaquatic, and/or limicolous (muddy) habitats. 
Family Lumbricidae
Genus Allolobophora Eisen, 1873
     Allolobophora chlorotica (Savigny, 1826)
Genus Eiseniella Michaelsen, 1900
     Eiseniella tetraedra (Savigny, 1826)
Family Sparganophilidae

Genus Sparganophilus Benham, 1892

Sparganophilus gatesi Reynolds, 1980
Sparganophilus helenae Reynolds, 1980
Sparganophilus komareki Reynolds, 1980
Sparganophilus kristinae Reynolds, 1980
Sparganophilus meansi Reynolds, 1980
Sparganophilus pearsei Reynolds, 1975
Sparganophilus pearsei libertiensis Reynolds, 1980
Sparganophilus pearsei sarasotae Reynolds, 1980
Sparganophilus smithi Eisen, 1896
Sparganophilus sonomae Eisen, 1896
Sparganophilus tamesis Benham, 1892 [= Sparganophilus eiseni Smith, 1895;
            see Rota et al. 2016]
Sparganophilus tennesseensis Reynolds, 1977b
Sparganophilus wilmae Reynolds, 1980

– – – – – Class Polychaeta – – – – –
Subclass Sedenteria
Order Sabellida
Family Fabricidae
Genus Manayunkia Leidy, 1859
Manayunkia speciosa (Leidy, 1859)
Subclass Polychaeta incertae sedis
Family Parergordrilidae
Genus Parergodrilus Reisinger, 1925

Parergodrilus heideri Reisinger, 1925 20 MN, WI


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.

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 hierarchy 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 recent research published 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 Lastočkin’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 Lastočkin’s (1921) paper has erroneously been recorded as ‘1924’ in numerous publications over the years; citations for both of these publications by Lastočkin  are included in the Literature Cited and References section].

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. L. Gustavsson (Swedish Museum of Natural History, Stockholm, 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 (prior to 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 — with all four of these groups 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 Reinmar Grimm (Timm and Grimm 2005) published 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 Štolc, 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 misused 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 – that to designate a neotype [based only on material previously misidentified as H. naidina] would be absurd.

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.” We have included this sp. inq. here as reference for those who may have seen this name in old publications or project reports. The World List of Marine Species (WoRMS Editorial Board, 2023) database record for O. flagellum provides additional information.

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). You are encouraged to review a recently published taxonomical study of the genus Aulodrilus by Ohtaka (2021).

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, 1999: p. 42). This incision occurs also in typical Ch. diaphanus; see Poddubnaya (1966).  Dr. Tarmo Timm (pers. comm. to MJW, 2 February 2012) translated some of Poddubnaya’s observations and some additional insight (based upon his own extensive studies) for this taxon:

In her well-illustrated paper, Poddubnaya demonstrated different versions of the prostomial incision in specimens of the otherwise typical, large Ch. diaphanus. Declaring that all records of ‘Ch. crystallinus’ [a false spelling of cristallinus by her] in SSSR (after Sperber and Cekanovskaja) were usually based on this incision, they may in fact belong to Ch. diaphanus. Poddubnaya hesitated in declaring that Ch. ‘usually’ crystallinus [sic] is a synonym of Ch. diaphanus, since these species must have also other differences, including the reproductive organs (following Sperber’s [1948] discussions).

Dr. Timm continued, stating that he can confirm Poddubnaya’s observation — that the incision is common in Ch. diaphanus. Sperber wrote that Ch. cristallinus is also considerably smaller. However, Dr. Timm also stated that he has seen smaller individuals of Ch. cristallinus — that could just as easily be young Ch. diaphanus.
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 Ripistes parasita, an introduction to North America, has been reported from two Canadian provinces, most of the Great Lakes, and several Midwestern, northeastern, southern, and southeastern states in the U.S. Globally, this species occurs in several European and Asian countries, Lake Baikal, Central and South America, Africa, and Australia. Wetzel currently has a manuscript in prep discussing global and North American distributional records for R. parasita, including a diagnosis, morphological, physiological, and behavioral observations, and habitat associations noted by those who have sent specimens to MJWetzel for verification. If you have collected and / or identified specimens of R. parasita, please contact Mark via the email address at the bottom of this webpage. Thanks!

12 Dero superterrenus (Michaelsen, 1912)— a Neotropical species originally described by Michaelsen in 1912 as Aulophorus superterrenus —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, in his PhD dissertation). 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 [however, neither Milligan nor Wetzel were aware of the paper by Frank and Lounibos at the time Milligan’s 1997 guide was sent to press]. Later, Frank and Fish (2008) published a paper in which they discussed 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 Dero superterrenus occurring in state were shared with MJW by 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 had collected specimens from a bucket used as a cistern to capture / retain rainwater runoff from a roof in Melrose, Alachua Co., FL). Several specimens sent to MJW by Doug Strom were  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 (Malaysia). [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]. Larry Hribar and MJW recently published a paper discussing the collection of aquatic oligochaetes and other invertebrates from a variety of natural phytotelmata and anthropogenic water bodies in Monroe County, Florida [historically monikered ‘The Conch Republic’ by those indigenous to the region]; most of the specimens that have been sorted from these collections and subsequently mounted on slides have been identified as D. superterrenus. This ms was published in the Florida Scientist [Vol. 86(4): 501–507] in December 2023 (citation below). 

13 The last name of Albertus Oschmann, the authority for Potamothrix bavaricus, is correctly spelled without an umlaut over the ‘O’. Tarmo Timm (pers. comm. to MJW) clarified — that the umlaut was absent from his last name in the original description for P. bavaricus (Zoologischer Anzieger 42: 559 [1913]) but had been accidentally ‘created’ later by one or more person(s), then ‘loyally’ copied and thus perpetuated by subsequent writers [including MJW]. This has been corrected in Nomenclatura Oligochaetologica – a catalogue of names, descriptions, and type specimens of the Oligochaeta. Editio Secunda.

14 In following Article 36, Principle of Coordination (ICZN 1999: p. 45), the authority and date for the subfamily name, Naidinae….is Ehrenberg, 1828not Lastočkin, 1921 [‘Lastockin, 1824’ [sic, 1924] on previous versions of this webpage]. {Pers. comm., Tarmo Timm to MJW}.

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 / justification for this new combination. Similar comment on this unsupported new combination was noted in Timm (2006b: 29–30).

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

17 Liu et al. (2017) summarized the extensive cryptic diversity of Limnodrilus hoffmeisteri.  Their study confirmed that the common taxon L. hoffmeisteri, as previously viewed, is not a single cosmopolitan species but rather a group of at least ten more or less cryptic species.  A neotype for L. hoffmeisteri sensu stricto was established which will provide a baseline for further revisions of the L. hoffmeisteri complex.

18 Molecular studies published by Martinsson and Erséus (2014) concluded that Cognettia sphagnetorum (Vejdovský, 1878), the morphotaxon by Nielsen and Christensen (1959), is a complex of several species that do not form a monophyletic group.  Martinsson, Rota and Erséus (2015), in their revision of the genus Cognettia Nielsen & Christensen, 1959, transferred the majority of species in this genus [including the type species, C. sphagnetorum (Vejdovský, 1878)] to the genus Chamaedrilus Friend, 1913; a few other former Cognettia species were transferred to the genus Euenchytraeus Bretscher, 1906.  In addition, the authors re-described several species, including the types for Chamaedrilus and Cognettia, and described two species new to science.  The authors also emphasized the importance of using molecular methods (e.g., DNA barcoding) to attain reliable identification of these enchytraeid taxa.  Schmelz and Collado (2015) affirmed Cognettia as a junior synonym of Chamaedrilus.  The following year, Martinsson et al. (2015a) revised the genus Cognettia, recognizing again the genus Chamaedrilus and describing the cryptic species in the sphagnetorum complex. In this same paper, Martinsson and Erséus established a neotype for Cognettia sphagnetorum because Vejdovský’s original description of Pachydrilus sphagnetorum was brief, and no type material is known.  Soon after this revision, an application to conserve the generic name Cognettia Nielsen & Christensen, 1959 was received (and assigned Case No. 3689) by the International Commission on Zoological Nomenclature (ICZN – the Commission). After additional correspondence pertinent to this application had been received, this Case was published by Schmelz et al. (2015). Three supportive comments and one adverse comment were then received by the Commission, followed by a response by Schmelz and Römbke (2016). The ruling to conserve the generic name Cognettia Nielsen & Christensen, 1959 by giving it precedence over two senior subjective synonyms, Euenchytraeus Bretscher, 1906 and Chamaedrilus Friend, 1913 was published by the Commission (ICZN 2018).  Martinsson et al. (2017) showed that Cognettia, Euenchytraeus, and a third enchytraeid genus, Stercutus Michaelsen, 1888, form a monophyletic group, but that the relationships between these three genera remain unresolved.  Martinsson (2019) recently published a morphological key to the Cognettia species of the world. In this key, he emphasized that Euenchytraeus is still considered to be a valid genus separate from Cognettia.

19 Our enchytraeid colleague, Jiří Schlaghamerský (2013a) documented what is believed to be the first record of Cognettia sphagnetorum in North America, based upon his faunistic and ecological studies of enchytraeid assemblages in old growth forests in the upper peninsula of the state of Michigan. Previously, our dear colleague, Brenda Healy (1989, 1996) tentatively reported Cognettia sphagnetorum from western Florida—based on just a few juvenile specimens.  However, C. sphagnetorum specimens were absent from Schlaghamerský’s other surveys in Nebraska (Schlaghamerský 2013b), in northern Wisconsin and Minnesota (Schlaghamerský’ et al. 2014), and then summarized in Schlaghamerský (2014).

20 Records of enchytraeids reported by Jiří Schlaghamerský et al. (2014) from Minnesota and/or Wisconsin.

21 A note about Br. hortensis will be posted here soon.  See van Haaren et al. (2005), Martin et al. (2018), and the World Register of Marine Species (WoRMS Editorial Board (2024) for additional information.

22 Ophidonais serpentina (Müller, 1773), a naidid common in European countries, North America, Africa, Japan, Iran, Korea, and Siberia, was recently redescribed based on specimens from the Xinkai River in Zhejiang Province, China by Yu et al. (2023) – the first report of this taxon from China. Previous morphological descriptions related to O. serpentina in the world were integrated, suggesting three morphological groupings:  a) specimens with dorsal chaetae starting from VI, b) specimens without dorsal chaetae, and c) specimens with an unstable starting position of the dorsal chaetae. After comparing the mitochondrial DNA (16S rDNA, COI), nuclear DNA (ITS2), and histones (H3) from the three groups, Bayesian inference and maximum likelihood phylogenetic analyses were performed based on the combined data set. Different analyses gave almost consistent phylogenetic trees. All of the genetic distances between the three groups were 0.00%. No genetic variation was detected between the specimens – regardless of the presence and starting position of dorsal chaetae. This result suggests that a single lineage of O. serpentina is widespread worldwide. You are encouraged to read this paper by Yu et al. (2023), and the literature cited therein. 

23 D. (Allodero) hylae Goodchild, 1951 and D. (Allodero) floridana Harman, 1971 are commensal on frogs and toads, living in the ureters and – in rare cases – in the cloaca of the host amphibian (Harman 1971, 1973).  Dero (Allodero) specimens have been collected in Florida, Louisiana, and Mississippi, as well as from Central and South American frogs in zoos, yet little is known about their life history.  Specimens of free-living Dero (Allodero) have been collected (R.O. Brinkhurst and J.H. Epler, pers. comm. to MJW) prompting the need for taxonomic and ecological re-assessment of these two species.

24 The extant species assigned to the genus Dero and its three subgenera – Allodero, Dero, and Aulophorus – present several a complicated nomenclatural, and systematic conundrums.  More to be added here…..

25 Historically, only three species of the genus Chaetogaster (C. diaphanus, C. limnaei, and C. diastrophus) have been commonly referenced in the literature, yet additional species have been described and prior molecular data suggests that there is cryptic diversity within named species. Recently, Mack et al. (2023) generated the first molecular phylogeny of the genus Chaetogaster based on mitochondrial and nuclear sequence data from 128 oligochaetes collected primarily from North American and European sites. Results of these phyolgenetic analyses suggest that the three commonly referenced species are a complex of 24 mostly cryptic species, with the dataset generated from the analyses suggesting that C. “diaphanus” is represented by two species, C. “limnaei” is represented by 3 species, and that C. “diastrophus” is represented by 19 species. You are encouraged to read this excellent paper (citation below) and consider the additional information and recommendations presented therein.

26 In their paper focusing on non-indigenous aquatic species, Grigorovich et al. (2003) reported the presence of both Vejdovskyella comata and V. intermedia in samples they collected from Lake Superior – each a Holarctic taxon considered native to the Laurentian Great Lakes.  In addition, one immature specimen of an anomalous, previously unreported form of Vejdovskyella – with morphological features of both V. intermedia and V. macrochaeta Lastoćkin, 1937. This specimen, co-occurring with V. intermedia, was collected in silty sand substrates from an off-shore site near Duluth, Minnesota.  Grigorovich et al. (2003) suggested that the morphological observations of this single specimen support Sergei Hrabe’s (1934) contention that V. intermedia and V. macrochaeta are one species.  However, given that only one immature specimen of this form was collected, and that there have been inconsistencies in morphological observations of V. comata by other North American oligochaetologists (e.g., Brinkhurst and Kathman, 1983, Kathman and Brinkhurst, 1998) who had lumped it with V. macrochaeta – the conventional wisdom is that additional, and hopefully mature specimens (those with penial chaetae present),considered critical to resolve this issue, need to be collected, properly preserved, and prepared for both morphological observations and genetic sequencing to resolve these taxonomic questions.

Literature Cited and Selected References

Ashlock, P.D. 1971. Monophyly and associated terms. Systematic Zoology 20: 63-69.

Brinkhurst, R.O. 1978. Freshwater Oligochaeta in Canada. Canadian Journal of Zoology 56(10): 2166-2175.

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. Canadian Special Publication of Fisheries and Aquatic Sciences 84. vi + 259 pp. [Note: This guide out of print; now superseded by Kathman and Brinkhurst (1998) (citation provided below)]

Brinkhurst, R.O. 1988. A taxonomic analysis of the Haplotaxidae. Canadian Journal of Zoology 66(10): 2243-2252.

Brinkhurst, R.O., and S.R. Gelder. 2001. Annelida: Oligochaeta, including Branchiobdellidae. Pages 431-463, In: J.H. Thorp and A.P. Covich (eds). Ecology and classification of North American freshwater invertebrates. Second Edition. Academic Press, San Diego, CA [superseded by Govedich et al. 2010, citation below].

Brinkhurst, R.O., and B.G.M. Jamieson. 1971. Aquatic Oligochaeta of the world. Univ. Toronto Press, Buffalo, New York. 860 pp. [Out of print; a classic reference if you can find a used copy].

Brinkhurst, R. O. & R. D. Kathman, 1983. A contribution to the taxonomy of the Naididae (Oligochaeta) of North America. Canadian Journal of Zoology 61: 2307–2312.

Brinkhurst, R.O., and M.J. Wetzel. 1984. Aquatic Oligochaeta of the world: Supplement. Canadian Technical Report of Hydrography and Ocean Sciences No. 44. v + 101 pp.

des Châtelliers, M.C., J. Juget, M. Lafont, and P. Martin. 2009. Subterranean aquatic Oligochaeta. Freshwater Biology 54: 678-690.

Cook, D.G. 1975. Cave dwelling aquatic Oligochaeta (Annelida) from the eastern United States. Transactions of the American Microscopical Society 94(1): 24-37.

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.

Corliss, J.O. 1970. Date of publication of “October 1969” Transactions. Transactions of the American Microscopical Society 89: 347.

Eckroth, M.C., and R.O. Brinkhurst. 1996. Tenagodrilus musculus, a new genus and species of Lumbriculidae (Clitellata) from a temporary pond in Alabama, USA. Hydrobiologia 334: 1-9.

Envall, I., M. Källersjö, and C. Erséus. 2006. Molecular evidence for the non-monophyletic status of Naidinae (Annelida, Clitellata, Tubificidae). Molecular Phylogenetics and Evolution 40: 570-584.

Erséus, C. 1999. Parvidrilus strayeri, a new genus and species, an enigmatic interstitial clitellate from underground waters in Alabama. Proceedings of the Biological Society of Washington 112(2): 327-337.

Erséus, C. 2005. Phylogeny of oligochaetous Clitellata. Hydrobiologia 535: 357-372.

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., and L. Gustavsson. 2002. A proposal to regard the former family Naididae as a subfamily within Tubificidae (Annelida, Clitellata). Hydrobiologia 485(1-3): 253-256.

Erséus, C., L. Gustavsson, and R.O. Brinkhurst. 2005. Case 3305. Tubificidae Vejdovský, 1876 (Annelida, Clitellata): proposed precedence over Naididae Ehrenberg, 1828. Bulletin of Zoological Nomenclature 62(4): 226-231.

Erséus, C., and M. Källersjö. 2004. 18s rDNA phylogeny of Clitellata (Annelida). Zoologica Scripta 33(2): 187-196.

Erséus, C., M. Källersjö, M. Ekman, and R. Hovmöller. 2002. 18S rDNA phylogeny of the Tubificidae (Clitellata) and its constituent taxa: dismissal of the Naididae. Molecular Phylogenetics and Evolution 22: 414-422.

Erséus, C., T. Prestegaard, and M. Källersjö. 2000. Phylogenetic analysis of Tubificidae (Annelida, Clitellata) based on 18S rDNA sequences. Molecular Phylogenetics and Evolution 15(3): 381-389.

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

Fend, S.V. 2005a. A review of the genus Eclipidrilus (Annelida: Clitellata: Lumbriculidae), with description of a new species from western North America. Zootaxa 969: 1-42.

Fend, S.V. 2005b. Rhynchelmis aleutensis n. sp. (Clitellata: Lumbriculidae) from Adak Island, Alaska. Zootaxa, 1093: 45-53.

Fend, S.V. 2009a. An evaluation of the genus Kincaidiana Altman, 1936, with the designation of Altmanella n. gen. (Annelida, Clitellata, Lumbriculidae). Zootaxa 2077: 1-30.

Fend, S.V. 2009b. Chapter 9. Lumbriculidae. Pp. 119-147, In: Wetzel, Fend, Coates, Kathman, and Gelder (complete citation below in this literature section).

Fend, S.V., and R.O. Brinkhurst. 2000. New species of Rhynchelmis (Clitellata, Lumbriculidae), with observations on the Nearctic species. Hydrobiologia 428: 1-59.

Fend, S.V., and R.O. Brinkhurst. 2010. Contributions towards a review of the genus Rhynchelmis Hoffmeister (Clitellata: Lumbriculidae). Zootaxa 2407: 1-27.

Fend, S.V., and J.J. Carter. 2014. Rhynchelmis subgenus Sutroa Eisen new rank, with two new species from western North America (Annelida, Clitellata, Lumbriculidae). Zootaxa 3760(2): 180-210.

Fend, S.V., and D.L. Gustafson. 2001. A new lumbriculid genus and species from North America (Clitellata, Lumbriculidae). Hydrobiologia 463: 13-22.

Fend, S., B.T. Hutchins, R. Gibson, B.F. Schwartz, and P.H. Diaz. 2023. New species of Lumbriculidae (Clitellata) from groundwater-dependent habitats in Texas. Zoosymposia 23: 52–77. ZooBank.^

Fend, S.V., and D.R. Lenat. 2007. Two genera of Lumbriculidae (Annelida, Clitellata) from North Carolina, USA. Zootaxa 1666: 1-22.

Fend, S.V., and D.R. Lenat. 2010. New southeastern Nearctic Rhynchelmis (Rhynchelmoides) species and the description of Pararhynchelmis n. gen. (Annelida: Clitellata: Lumbriculidae). Zootaxa 2554: 1-22.

Fend, S.V., and D.R. Lenat. 2012. New Eclipidrilus species (Annelida, Clitellata, Lumbriculidae) from southeastern North America. Zootaxa 3194: 51-67.

Fend, S.V., Y. Liu, D. Steinmann, O. Giere, H.A. Barton, F. Luiszer, and C. Erséus. 2016. Limnodrilus sulphurensis n. sp., from a sulfur cave in Colorado, USA, with notes on the morphologically similar L. profundicola (Clitellata, Naididae, Tubificinae). Zootaxa 4066(4): 451-468.

Fend, S.V., and P. Rodriguez. 2003. Eremidrilus n. gen. (Annelida, Clitellata, Lumbriculidae) and new species from California, U.S.A. Canadian Journal of Zoology 81(3): 515-542.

Fend, S.V., P. Rodriguez, and D.R. Lenat. 2015. Uktena riparia n. gen., n. sp. (Annelida, Clitellata, Lumbriculidae), a new spermatophore-producing oligochaete. Zootaxa 3994(3): 411-424.

Fish, D. 1976. Structure and composition of the aquatic invertebrate community inhabiting epiphytic bromeliads in south Florida and the discovery of an insectivorous bromeliad. Ph.D. Dissertation, University of Florida. [as referenced in Frank, J.H., and D. Fish, 2008; citation below].

Frank, J.H., and D. Fish. 2008. Potential biodiversity loss in Florida bromeliad phytotelmata due to Metamasius callizona (Coleoptera: Dryophthoridae), an invasive species. Florida Entomologist 91(1): 1-8.

Frank, J.H., and L.P. Lounibos. 1987. Phytotelmata: swamps or islands? Florida Entomologist 70: 14-20.

Gates, G.E. 1942. Check list and bibliography of North American earthworms. American Midl. Nat. 27(1): 86-108.

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.

Gelder, S.R. 2010. Branchiobdellida, section III (pp. 402-410), in Govedich, F.R., B.A. Bain, W.E. Moser, S.R. Gelder, R.W. Davies, and R.O. Brinkhurst. Chapter 12. 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. [] USD$ 139.95 / 93.95 Euro.

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. [] USD$ 139.95 / 93.95 Euro.

Grigorovich, I.G., A.V. Korniushin, D.K. Gray, I.C. Duggan, R.I. Colautti, and H.J. MacIsaac. 2003. Lake Superior: an invasion coldspot? Hydrobiologia 499: 191–210.

Harman, W.J. 1970. Revision of the family Opistocystidae (Oligochaeta). Transactions of the American Microscopical Society 88(4): 472-478. [ Note: ‘1970’ is the correct year of publication for this citation (not ‘1969’, a misdating on all papers in the fourth issue of this volume). Actual publication of this paper, and all other papers in this issue, occurred in January 1970; the editor of the journal clarified this situation in the subsequent volume (see Transactions of the American Microscopical Society, volume 89, page 347), stating that the 1970 date was correct and should be used. ]

Healy, B. 1989. Preliminary report on the Enchytraeidae (Oligochaeta) of West Florida. – Hydrobiologia 180: 47–56. ^

Healy, B. 1996. Records of Enchytraeidae (Annelida: Oligochaeta) from West Florida.1. Mesenchytraeus, Cognettia, Bryodrilus, Hemienchytraeus, Henlea and Buchholzia. – Proceedings of the Biological Society of Washington 109(1): 118–137. ^

Hiltunen, J.K., and D.J. Klemm. 1980. A guide to the Naididae (Annelida: Clitellata: Oligochaeta) of North America. EPA-600/4-80-031. Environmental Monitoring and Support Laboratory, Office of Research and Development, U.S. Environmental Protection Agency, Cincinnati, OH. 48 pp.

Hiltunen, J. K., and D. J. Klemm. 1985. Freshwater Naididae (Annelida: Oligochaeta). Pages 24-43, In: D.J. Klemm (ed). A guide to the freshwater Annelida (Polychaeta, naidid and tubificid Oligochaeta, and Hirudinea) of North America. Kendall/Hunt Publishing Co., Dubuque, IA.

Holmquist, C. 1976. Lumbriculids (Oligochaeta) of Northern Alaska and Northwestern Canada. Zoologische Jahrbücher–Abteilung für Systematik, Geographie und Biologie der Tiere, Jena, 163: 377-431.

Holmquist, C. 1983. What is Tubifex tubifex (O.F. Müller) (Oligochaeta, Tubificidae)? Zoologica Scripta 12(3): 187-201.

Holmquist, C. 1985.  A revision of the genera Tubifex Lamark, Ilyodrilus Eisen, and Potamothrix Vejdovský & Mrázek (Oligochaeta, Tubificidae), with extensions to some connected genera. Zoologische Jahrbücher–Abteilung für Systematik Ökologie und Geographie der Tiere, Jena, 112(3): 311-366.

Howmiller, R.P., and M.S. Loden. 1976. Identification of Wisconsin Tubificidae and Naididae. Transactions of the Wisconsin Academy of Sciences Arts and Letters 64: 185-197.

Hribar, L. J. and M. J. Wetzel. 2023. Dero superterrenus (Michaelsen, 1912) (Annelida, Clitellata, Naididae) associated with phytotelmata and artificial containers in Monroe County, Florida, USA.  Florida Scientist 86(4): 501–507.

International Commission on Zoological Nomenclature [ICZN]. 1999. International Code of Zoological Nomenclature, 4th ed. Published by the International Trust for Zoological Nomenclature, c/o The Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom. xxix + 306 pp. [provisions of this Code supersede those of previous editions with effect from 1 January 2000].  Note: Please visit the ICZN website to access the Code Online, and to see two important updates to the Code:  1)  incorporating Declaration 44, amendments of Article 74.7.3 – specific to declaration of lectotypes (with effect from 31 December 1999),  and  2)  the amendment on e-publication, amendments to Articles 8, 9, 10, 21 and 78 (with effect from 1 January 2012).

International Commission on Zoological Nomenclature [ICZN]. 2007. Opinion 2167 (Case 3305). Naididae Ehrenberg, 1828 (Annelida, Clitellata): precedence over Tubificidae Vejdovský, 1876 maintained. Bulletin of Zoological Nomenclature 64: 71-72.

Kasprzak, K. 1972. Variability of Chaetogaster diaphanus (Gruithuisen, 1828) (Oligochaeta, Naididae) in different environments. Zoologica Poloniae 22: 43-51.

Kathman, R.D. 1985. Synonymy of Pristinella jenkinae (Naididae: Oligochaeta). Proceedings of the Biological Society of Washington 98(4): 1022-1027.

Kathman, R.D., and R.O. Brinkhurst. 1998, 1999. Guide to the freshwater oligochaetes of North America. Aquatic Resources Center, College Grove, TN. iv + 264 pp. [Note: an updated version of this guide was printed in May 1999, with addenda noted in subsequent printings.  All inquiries regarding this guide should be addressed to: Dr. R.D. Kathman via E-mail [ ** r.deedee.kathman{AT} ].

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.

Kernall, M.J. 1914. An undescribed species of Tubifex from Illinois. (Unpublished Thesis). Graduate School, University of Illinois. 21 pp.

Lastočkin, D.A. 1921. Issledovanija po faune Ivanovo-Voznesenskoj gubernii, organizovannye sel’sko- hozjajstvennym fakul’tetom I.-V. P. I. letom 1920 goda. 2. Fauna Oligochaeta limicola rajona issledovanija. Izvestija Ivanovo-Voznesenskogo Politehniceskogo Instituta 4: 70-77.

Lastočkin, D.A. 1924. Novye i redkie Copepoda i Oligochaeta v faune Ivanovo-Voznesenskoi gubernii. [New and rare Copepoda and Oligochaeta from central Russia (Ivanovo-Voznesensk Government)]. Izvestiya Rossijskogo Gidrologiceskogo Instituta (Bulletin de l’Institut Hydrologique de Russie [Leningrad]) 9(1):1-22, figs. 1-7. [in Russian with English summary].

Lopez, L., B. Elizola, I. Deiss, and R. Rios. 2005. Phoretic behaviour of bromeliad annelids (Dero) and ostracods (Elpidium) using frogs and lizards as dispersal vectors. Hydrobiologia 549: 15-22.

Liu, Y., S.V. Fend, S. Martinsson and C. Erséus. 2017. Extensive cryptic diversity in the cosmopolitan sludge worm Limnodrilus hoffmeisteri (Clitellata, Naididae). Organisms Diversity & Evolution DOI 10.1007/s13127-016-0317-z.

Mack, J.M., M. Klinth, S. Martinsson, R. Lu, H. Stormer, P. Hanington, H.C. Proctor, C. Erséus, and A.E. Bely. 2023. Cryptic carnivores: Intercontinental sampling reveals extensive novel diversity in a genus of freshwater annelids. Molecular Phylogenetics and Evolution 182 (2023) 107748.

Magalhães, W.F., Hutchings, P., Oceguera-Figueroa, A., Martin, P., Schmelz, R.M., Wetzel, M.J., Wiklund, H., Maciolek, N.J., Kawauchi, G.Y. & Williams, J.D. 2021. Segmented worms (Phylum Annelida): a celebration of twenty years of progress through Zootaxa and call for action on the taxonomic work that remains. Zootaxa, 4979 (1), 190–211.

Martin, P. 2001. On the origin of the Hirudinea and the demise of the Oligochaeta. Proceedings of the Royal Society of London 268: 1089-1098.

Martin, P., I. Kaygorodova, D. Yu. Sherbakov, and E. Verheyen. 2000. Rapidly evolving lineages impede the resolution of phylogenetic relationships among Clitellata (Annelida). Mol. Phyl. Evol. 15: 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. Lévêque, H. Segers & K. Martens, eds. Freshwater Animal Diversity Assessment. Hydrobiologia 595(1).

     The above publication is superseded by:

Martin, P., Martinez-Ansemil E., Pinder, A., Timm T. & Wetzel, M.J., 2016. World checklist of freshwater Oligochaeta species. World Wide Web electronic publication. Available online at [date accessed] [this is an on-line update (first posted 15 December 2016) of our 2008 publication {Hydrobiologia, 595, 117-127; complete citation provided below}, with support from FADA, Freshwater Animal Diversity Assessment Initiative].

Martin, P., J. Wuillot, S. Martinsson, and C. Erséus. 2018. Integrative species delimitation and phylogeny of the branchiate worm Branchiodrilus (Clitellata, Naididae). Zoologica Scripta 47(6):727-742.

Martínez-Ansemil, E., B. Samburgar, and N. Giani.  2002. First record of Parvidrilidae (Annelida, Oligochaeta) in Europe with a description of a new species (Parvidrilus spelaeus sp. nov.) and comments on the family and its phyletic relationships. Journal of Zoology, London 256: 495-503.

Martínez-Ansemil, E., M. Creuze Des Châtelliers, P. Martin, and B. Sambugar. 2002. The Parvidrilidae – a diversified groundwater family: description of six new species from southern Europe, and clues for its phylogenetic position within Clitellata (Annelida). Zoological Journal of the Linnean Society 166: 530-558.

Martinsson, S., and C. Erséus. 2014. Cryptic diversity in the well-studied terrestrial worm Cognettia sphagnetorum (Clitellata: Enchytraeidae). Pedobiologia 57: 27–35.

Martinsson, S.; Rota, E.; Erséus, C. 2015. Revision of Cognettia (Clitellata, Enchytraeidae): re-establishment of Chamaedrilus and description of cryptic species in the sphagnetorum complex. Systematics and Biodiversity 13(3): 257-277.

McKey-Fender, D., and W.M. Fender. 1988. Phagodrilus gen.nov. (Lumbriculidae): systematics and biology of a predaceous oligochaete from western North America. Canadian Journal of Zoology 66(10): 2304-2311.

McKey-Fender, D., and W.M. Fender. 2001. Descriptions of new species of the predaceous lumbriculid oligochaete Phagodrilus from western North America. Megadrilogica 8(11): 57-84.

Michaelsen, W. 1900. Oligochaeta. Das Tierreich – Eine Zusammenstellung und Kennzeichnung der rezenten Tierformen. Vol. 10. [The Animal Kingdom. A compilation and identification of the extant animal forms]. xxix + 575 pp. Berlin.

Michaelsen, W. 1919. Über die Beziehungen der Hirudineen zu den Oligochaeten. Mitteilungen aus dem Naturhistorischen Museum, Hamburg [as Mitt. Naturh. Mus. Hamburg] 36: 131-153.

Milligan, M.R. 1997. Identification manual for the aquatic Oligochaeta of Florida, Volume I. Freshwater oligochaetes. 187 pp. Florida Department of Environmental Protection, Tallahassee. [This manual is available free, as a pdf document you can download from the Florida Department of Environmental Protection – Bureau of Laboratories List of Keys and Guides website; you must scroll down to the listing for this manual, then click on the entry ”oligofw.pdf” to begin the download; this file is 5.7 MB in size, so will be a very slow download if you are using a dial-up internet connection]. [A memoriam for Michael, who passed away in November 2005, is presented in Wetzel 2008b (citation below)].

Ohtaka, A. 2021. Taxonomical study of Japanese Aulodrilus Bretscher (Annelida, Clitellata, Tubificinae) with descriptions of two new species. Zootaxa 4952(1): 1–32.

Poddubnaja, T.L. 1966. K sistematike Chaetogaster diaphanus Gruith. (Oligochaeta, Naididae). In: Plankton i bentos vnutrennih vodoemov. Nauka, Moskva–Leningrad. Akademija Nauk SSSR, Institut biologii vnutrennih vod, Trudy 12(15): 120-124. [in Russian; well illustrated].

Reynolds, J.W. 2001. Sparganophilidae — are terrestrial oligochaetologists missing the habitat in North America? Megadrilogica 8(11): 82-84.

Reynolds, J.W. 2008. The earthworm family Sparganophilidae (Annelida: Oligochaeta) in North America, revisited. Megadrilogica 12(9): 125-143.  [Note: an update of this paper coauthored by W.K. Reeves, J.W. Reynolds, and M.J. Wetzel is in preparation, likely to be published in late 2023 or early 2024].

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 (below), are not currently available; existing copies have either been misplaced, or accidentally discarded. Dr. Reynolds is attempting to locate copies. MJW, October 2006]

Reynolds, J.W., and D.G. Cook. 1981. Nomenclatura oligochaetologica. Supplementum primum. A catalogue of names, descriptions and type specimens of the Oligochaeta. [Publ. by Univ. New Brunswick] Fredericton, New Brunswick. v + 39 pp. [not currently available; 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 Museum Monographic Series (Natural Science) No. 8. v + 37 pp. [this second supplement, and the third (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. 1993. Nomenclatura oligochaetologica. Supplementum tertium. A catalogue of names, descriptions and type specimens of the Oligochaeta. New Brunswick Museum Monographic Series (Natural Science) No. 9. vi + 33 pp. [To order, see note above.]

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, Lumbricidae, Megascolecidae, and Sparganophilidae) of Illinois, USA. Megadrilogica 15(4): 135-67.

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): 191-211.

Reynolds, J.W., and M.J. Wetzel. 2014.  A checklist by counties of earthworms (Oligochaeta: Acanthodrilidae, Lumbricidae, Megascolecidae, and Sparganophilidae) in Michigan, USA.  Megadrilogica 17(5): 51-72.

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

Rodriguez, P. 2006. Stylodrilus californianus n. sp., a new lumbriculid (Annelida: Oligochaeta) from North America. Hydrobiologia 333: 161-164.

Rodriguez, P., and K. A. Coates. 2006. A new American Stylodrilus species (Lumbriculidae, Oligochaeta). Canadian Journal of Zoology 74(1): 92-96.

Rodriguez, P., S.V. Fend, and D.R. Lenat. 2014. Sylphella puccoon gen. n., sp. n. and two additional new species of aquatic oligochaetes (Lumbriculidae, Clitellata) from poorly-known lotic habitats in North Carolina (USA). ZooKeys 451: 1-32. {Cookidrilus pocosinus sp.n., Stylodrilus coreyi sp.n.} doi: 10.3897/zookeys.451.7304.

Rodriguez, P., and N. Giani. 1987. Sur deux espèces de Lumbriculidae (Oligochaeta) cavernicoles: Cookidrilus speluncaeus n. gen., sp. n. et Trichodrilus diversisetosus Rodriguez, 1986. Bulletin de la Société d’Histoire Naturelle de Toulouse 123: 45–49.

Rousset, V., F. Pleijel, G.W. Rouse, C. Erséus, and M.E. Siddall. 2007. A molecular phylogeny of annelids. Cladistics 23: 41-63.

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.

Rota, E., Martinsson, S., Erséus, C. 2015. Comments on Cognettia Nielsen & Christensen, 1959 (Annelida, Oligochaeta, Enchytraeidae): proposed precedence over Euenchytraeus Bretscher, 1906 and Chamaedrilus Friend, 1913 (Case 3689). Bulletin of Zoological Nomenclature 72: 303–307.

Schlaghamerský, J. 2013a. Enchytraeid assemblages (Annelida: Clitellata: Enchytraeidae) of two old growth forests in the Porcupine Mountains (Michigan, USA). – Soil Organisms 85 (2): 85–96.

Schlaghamerský, J. 2013b. The enchytraeid assemblage (Annelida: Clitellata: Enchytraeidae) of a Sandhill prairie site in Nebraska, USA. – Soil Organisms 85 (2): 97–102.

Schlaghamerský, J. 2014. A brief history of research on potworms (Annelida: Clitellata: Enchytraeidae) of North America. Pp. 177–188, In: Pavlíček, T., P. Cardet, M.T. Almeida, C. Pascoal, and F. Cássio (Eds.). 2014.  Advances in Earthworm Taxonomy VI (Annelida: Oligochaeta). Proceedings of the 6th International Oligochaete Taxonomy Meeting (6th IOTM), Palmeira de Faro, Portugal, 22- 25 April, 2013. Publisher: Kasparek Verlag, Heidelberg. ISBN 9783-925064-69-2.  Please contact Dr. Schlaghamerský via e-mail { jiris'[AT]’ } if you would like a reprint of this paper. 

Schlaghamerský, J., Eisenhauer, N. & Frelich, L. E. 2014. Earthworm invasion alters enchytraeid community composition and individual biomass in Northern Hardwood Forests of North America. – Applied Soil Ecology 83: 159–169.

Schmelz, R.M.; Collado, R. 2015. Checklist of taxa of Enchytraeidae (Oligochaeta): an update. Soil Organisms 87(2): 149-152.

Schmelz, R.M., Collado, R., Roembke. J. 2015. Case 3689. Cognettia Nielsen & Christensen, 1959 (Annelida, Oligochaeta, ENCHYTRAEIDAE): proposed precedence over Euenchytraeus Bretscher, 1906 and Chamaedrilus Friend, 1913. Bulletin of Zoological Nomenclature 72: 186-192.

Schmelz, R.M., Roembke, J. 2016. Comment on Cognettia Nielsen & Christensen, 1959 (Annelida, Oligochaeta, Enchytraeidae): giving precedence to the name promotes stability. Bulletin of Zoological Nomenclature 73: 42-45.

Sjölin, E., C. Erséus, and M. Källersjö. 2005. Phylogeny of Tubificidae (Annelida, Clitellata) based on mitochondrial and nuclear sequence data. Molecular Phylogenetics and Evolution 35: 431-441.

Smith, M.E. 1984. Aquatic Oligochaeta (Naididae) of Washington. Northwest Science 58(3): 165-170.

Spencer, D.R. 1980. The aquatic Oligochaeta of the St. Lawrence Great Lakes region. Pages 115-164, In: Brinkhurst, R.O., and D.G. Cook (eds). 1980. Aquatic oligochaete biology. Plenum Press, New York and London.

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.

Sperber, C. 1948. A taxonomic guide to the Naididae. Zoologiska Bidrag från Uppsala 28: 1-296 + 21 plates.

Sperber, C. 1950. A guide for the determination of European Naididae. Zoologiska Bidrag från Uppsala 29: 44-78 + 3 plates.

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]

Timm, T. [2005] 2007. Some misused names in aquatic Oligochaeta. Pp. 53–60, In: V.V. Pop and A.A. Pop (eds). Advances in earthworm taxonomy II (Annelida: Oligochaeta). [Proceedings of the 2nd International Oligochaeta Taxonomy Meeting dedicated to Victor Pop, Cluj-Napoca, Romania, 4-8 September 2005]. Cluj University Press, Institute of Biological Research, 48 Republicii Street, Cluj-Napoca, Romania. ISBN 973-610-395-1. This volume is available for purchase [ order via E-mail, to: ]

Timm, T. 2006a. Comment on the proposed precedence of Tubificidae Vejdovský, 1876 over Naididae Ehrenberg, 1828 (Annelida, Clitellata). Bulletin of Zoological Nomenclature 63(1): 48 [March].

Timm, T. 2006b. A Nearctic tubificid Varichaetadrilus harmani (Loden) n. comb. in a Dutch wetland, with remarks on Tubifex tubifex (Müller) Annelida: Oligochaeta). Zootaxa 1281: 21-39.

Timm, T., and R. Grimm. 2005. What is Homochaeta naidina Bretscher, 1896 (Annelida, Oligochaeta, Naididae)? Zoosystema 27(3): 469-482.

Valentine, J.W. 2004. On the origin of phyla. University of Chicago Press, Chicago and London. 614 pp.

van Haaren, T., M. Soes, and R. Munts. 2005.  Branchiodrilus hortensis, een nieuwe exotische borstelworm in Nederland (Annelida: Oligochaeta). Nederlandse Faunistische Mededeling 22: 17-21.

Vejdovský, F. 1884. System und Morphologie der Oligochaeten. Franz Rivnac, Prague. 1-172.

Wassell, J.T. 1984. Revision of the lumbriculid oligochaete Eclipidrilus Eisen, 1881, with descriptions of three subgenera and Eclipidrilus (Leptodrilusfontanus n. subg., n. sp. from Pennsylvania. Proceedings of the Biological Society of Washington 97(1): 78–85.

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.

Wetzel, M.J. 1992. Aquatic Annelida of Illinois: Introduction and checklist of species. Transactions of the Illinois State Academy of Science 85(1 & 2): 87–101.

Wetzel, M.J. 2006. In Memoriam: Professor Walter James Harman, PhD (1928-2002). Hydrobiologia 564: 1–4.

Wetzel, M.J. 2008a. 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. 2008b. 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. 2009. Taxonomy, systematics, and ecology of the freshwater oligochaetes and branchiobdellidans (Annelida, Clitellata) of North America. A workbook. vi + 280 pp. + 4 color plates. 1 November. 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.}  Note: This 2009 guide now out of date / out of print; a new version is being considered.

Wetzel, M.J., and P. Morgan. 2002. Aquatic Oligochaeta (Annelida, Clitellata) of the Great Smoky Mountains National Park, North Carolina and Tennessee. Southeastern Biology 49(4): 389–392.

Wetzel, M.J., and M.A.P. Morgan. 2007. Aquatic microdrile oligochaetes (Annelida, Clitellata) of Great Smoky Mountains National Park, North Carolina and Tennessee. Pp. 153–158, In: P. Cox, (Ed.). The Great Smoky Mountains National Park All Taxa Biodiversity Inventory: A Search for Species in Our Own Backyard. Southeastern Naturalist 6 (Special Issue 1). 238 pp.

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.

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. [if interested in obtaining a copy, please contact me [MJW] via the email address, below; cost: USD$43.00 (postage included; discount on multiple copy purchases)].

WoRMS Editorial Board (2023). World Register of Marine Species. An authoritative classification and catalogue of marine names [now including accounts for freshwater and terrestrial oligochaetes, acanthobdellidans, and branchiobdellidans].

Yu, J., T. Zhou, H. Wang, and Y. Cui. 2022. First record of Ophidonais serpentina (Müller, 1773) (Oligochaeta: Naididae) in China: The occurrence or absence of needles are intraspecific differences. Diversity 2022, 14, 265.

Thank you for your interest! If you have questions, comments, suggestions, or ideas regarding this website, please send them to me via E-Mail: mjwetzel{AT}

** NOTE:
The ‘@’ symbol in the email addresses on this page have been replaced with ‘{AT}’ to deter the ‘harvesting, mining, or sweeping’ of active webpages by programs initiated by virtual telemarketers, spammers, hackers, skimmers, spoofers, bots, and similar ilk who/that perpetuate malware, adware, and/or otherwise create and flood the internet with programs to search the virtual world to auto-collect valid e-mail, social media handles, and other website addresses.  Thus, you must replace the ‘{AT}’ (in email addresses noted above on this page) with the ‘@’ symbol in order for your email message to be sent and received. 

Suggested citation for this electronic webpage: Wetzel, M.J., R.D. Kathman, S.V. Fend, and K.A. Coates. 2024. Classification and checklist of the freshwater oligochaetes occurring in North America north of Mexico. February.
World Wide Web URL: