Introduction. 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 the last few 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, and G. Stinger Guala 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 entails correcting nomenclatural discrepancies, incorporating recent synonymies, and providing supportive references for all valid taxa. This project is nearly completed (January 2013).

The Taxonomic Certification Program [TCP] underwritten by the Society for Freshwater Science [SFS] (formerly, the North American Benthological Society [NABS], 1973-2011) 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; however, because I am now 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 the terrestrial oligochaetes, leeches, branchobdellidans, and the aeolosomatidans (a group now aligned with the Polychaeta).

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

Because several megadrile oligochaetes are commonly found in aquatic and semi-aquatic habitats, those species will soon be 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, deposition of specimens, identification), 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 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 (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.

Reynolds and Wetzel (2012) published a second update of known distributions for earthworm species occurring in North America, including Mexico, the Commonwealth of Puerto Rico, Hawaii, and Bermuda that had been published since Reynolds and Wetzel (2008), including distributional maps for all new species records.

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.

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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, and numerous other oligochaetologists working in North America and elsewhere in the World. As this list grows to include the leeches and branchiobdellidans occurring in North America, the systematic and nomenclatural expertise for those groups will be provided by Donald J. Klemm and William Moser, and Stuart R. Gelder, respectively.

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 have served on this Committee since 1989, and (in collaboration with many colleagues) have 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 have changed accordingly, the AFS CNAI Program is currently on hold.

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 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 taxonomy and systematics.

We especially thank our dear friend and colleague, Tarmo Timm (Estonian Insitute 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).

As of 01 August 2012, 220 nominal species of freshwater (microdrile) oligochaetes are known to occur in North America north of Mexico [several megadrile oligochaetes ('earthworms') also occur in aquatic and semi-aquatic habitats; these taxa will be added to this web-based list, soon]. 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 families Naididae and Tubificidae (see footnote 1, below), and Lumbriculidae, has been documented in publications such as Brinkhurst (1978, 1986), Brinkhurst and Jamieson (1971), Brinkhurst and Wetzel (1984), Hiltunen and Klemm (1980, 1985), Brinkhurst and Gelder (2001), Govedich et al. (2010), Kathman and Brinkhurst (1998), and Stimpson et al. (1982, 1985). State and regional keys, guides, and papers with species distributions include Howmiller and Loden (1976), Kathman and Wetzel (2003), Milligan (1997), Smith (1984), Spencer (1980), Spencer and Denton (2003), Spencer and Hudson (2003), Spencer and Wisseman (1993), Strayer (1990), Wetzel (1982, 1992), Wetzel and Taylor (1999), Wetzel et al. (2009), and Whitley (1982). Please consult the literature cited in each of the above publications for additional - and usually more specific - distributional information for taxa.

Historically, the family Lumbriculidae has received less attention than other families of microdrile oligochaetes. Fortunately, extensive work with this group by Steve Fend and others, particularly in the western and southeastern regions of the U.S., has resulted in a significant increase in our distributional knowledge of this group. In addition, their research and publications have provided us with descriptions of several new lumbriculid genera and species as well as redescriptions and new combinations of yet other previously described / problematical taxa (e.g., Fend and Brinkhurst 2000, 2010; Fend and Gustafson 2001; Fend and Rodriguez 2003; Fend 2005a, 2005b, 2006, 2009; and Fend and Lenat 2007, 2010, 2012). While a few of these papers are relatively specific to taxa occurring in northwestern North America, information presented therein provides insight to taxa known or thought likely to occur in Florida and elsewhere in the southeastern U.S.

Nomenclature. Nomenclatural changes and synonymies for North American species published since Kathman and Brinkhurst (1998) have been included herein; to the best of our knowledge, the nomenclature for species presented here is current as of 1 June 2007. Nomenclatura Oligochaetologica [N.O.] (Reynolds and Cook 1976) and its three supplements -- Nomenclatura Oligochaetologica Supplementum Primum [N.O.S.P.] (Reynolds and Cook 1981), Nomenclatura Oligochaetologica Supplementum Secundum [N.O.S.S.] (Reynolds and Cook 1989), and Nomenclatura Oligochaetologica Supplementum Tertium [N.O.S.T.] (Reynolds and Cook 1993) -- present a catalogue of names, descriptions, and type specimens of the Oligochaeta.

The fourth supplement, Nomenclatura Oligochaetologica Supplementum Quartum [N.O.S.Q.] (Reynolds and Wetzel in prep.) is still being compiled; as of 30 March, 2010, accounts for two new families, 83 genera, 27 subgenera, and 1,350+ species (all described since publication of N.O.S.T.) have been included. More information on this nomenclator series is available via the Nomenclatura Oligochaetologica website.

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

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

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

Introduction to the checklist. The primary purpose of this website 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 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, 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 (2008).

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

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

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

- 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).

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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).

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Genus Altmanella Fend, 2009

Altmanella freidris (Cook, 1966) [transferred from Kincaidiana by Fend (2009)]

Altmanella idahoensis Fend, 2009

Altmanella lenati Fend, 2009

Genus Eclipidrilus Eisen, 1881

Eclipidrilus asymmetricus (F. Smith, 1896)

Eclipidrilus breviatriatus Fend & Lenat, 2012

Eclipidrilus (Premnodrilus) daneus Cook, 1966

Eclipidrilus (Leptodrilus) fontanus Wassel, 1984

Eclipidrilus (Eclipidrilus) frigidus Eisen, 1881

Eclipidrilus (Leptodrilus) ithys Brinkhurst, 1998

Eclipidrilus (Leptodrilus) lacustris (Verrill, 1871)

Eclipidrilus macphersonae Fend & Lenat, 2012

Eclipidrilus microthecus Fend & Lenat, 2012

Eclipidrilus pacificus Fend, 2005

Eclipidrilus (Premnodrilus) palustris (F. Smith, 1900)

Genus Eremidrilus Fend & Rodriguez, 2003

Eremidrilus allegheniensis (Cook, 1971)

Eremidrilus coyote Fend & Rodriguez, 2003

Eremidrilus elegans Fend & Rodriguez, 2003

Eremidrilus felini Fend & Rodriguez, 2003

Eremidrilus ritocsi Fend & Rodriguez, 2003

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]

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 (Rhynchelmoides) alaskana Holmquist, 1976

Rhynchelmis aleutensis Fend, 2005

Rhynchelmis brooksi Holmquist, 1976

Rhynchelmis (Rhynchelmoides) elrodi Smith & Dickey, 1918

Rhynchelmis gilensis Fend & Brinkhurst, 2000

Rhynchelmis (Rhynchelmoides) glandula Altman, 1936

Rhynchelmis gustafsoni Fend & Brinkhurst, 2000

Rhynchelmis monsserratus Fend & Brinkhurst, 2000

Rhynchelmis (Rhynchelmoides) rostrata Eisen, 1888

Rhynchelmis (Rhynchelmoides) saxosa Fend & Brinkhurst, 2000

Rhynchelmis utahensis Fend & Brinkhurst, 2000

Rhynchelmis yakimorum Fend & Brinkhurst, 2000

Rhynchelmis (Rhynchelmoides) bolinensis Fend & Lenat, 2010 [ distr.: U.S. - NC]

Rhynchelmis (Rhynchelmoides) croatanensis 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 heringianus Claparède, 1862 - i -

Stylodrilus sovaliki Holmquist, 1976

Stylodrilus wahkeenensis Rodriguez & Coates, 1996

Genus Styloscolex Michaelsen, 1901

Styloscolex opisthothecus Sokolskaya, 1969

Genus Tenagodrilus Eckroth & Brinkhurst, 1996

Tenagodrilus musculus Eckroth & Brinkhurst, 1996

Genus Trichodrilus Claparède, 1862

Trichodrilus allobrogum Claparède, 1862 - [this species has not been seen since 1917]

Trichodrilus culveri Cook, 1975

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Subfamily Naidinae Ehrenberg, 1828 ^1,2

Genus Chaetogaster von Baer, 1827

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)

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 -

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

Dero (Allodero) hylae Goodchild, 1951

Dero (Allodero) floridana Harman, 1971

Dero (Dero) abranchiata Harman, 1977

Dero (Dero) botrytis Marcus, 1943

Dero (Dero) digitata (Müller, 1773)

Dero (Dero) nivea Aiyer, 1930

Dero (Dero) obtusa d'Udekem, 1855

Dero (Dero) pectinata Aiyer, 1930

Dero (Dero) trifida Loden, 1979

Dero (Aulophorus) borellii Michaelsen, 1900

Dero (Aulophorus) flabelliger (Stephenson, 1931)

Dero (Aulophorus) furcata (Müller, 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)

Genus Vejdovskyella Michaelsen, 1903

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

this genus 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 (Hrabe, 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 Limnodrilus Claparède, 1862

Limnodrilus cervix Brinkhurst, 1963

Limnodrilus claparedeianus Ratzel, 1868

Limnodrilus hoffmeisteri Claparède, 1862

Limnodrilus maumeensis Brinkhurst & Cook, 1966

Limnodrilus profundicola (Verrill, 1871)

Limnodrilus rubripenis Loden, 1977

Limnodrilus silvani Eisen, 1879

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)} ¹¹

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)

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 brevidentatus Brinkhurst, 1965

Rhyacodrilus coccineus (Vejdovský, 1875)

Rhyacodrilus falciformis Bretscher, 1901

Rhyacodrilus montana (Brinkhurst, 1965) ['montanus' is incorrect]

Rhyacodrilus sodalis (Eisen, 1879)

Rhyacodrilus subterraneus Hrabě, 1963

Subfamily Phallodrilinae Brinkhurst, 1971

Genus Gianius Erséus, 1992

Gianius aquaedulcis (Hrabě, 1960)

Genus Thalassodrilus Brinkhurst, 1963

Thalassodrilus hallae (Cook & Hiltunen, 1975)

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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)}

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Genus Parvidrilus Erséus, 1999

Parvidrilus strayeri Erséus, 1999

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Genus Achaeta Vejdovský, 1878

Achaeta spp.

Genus Barbidrilus Loden & Locy, 1980

Barbidrilus paucisetus Loden & Locy, 1980

Genus Bryodrilus Ude, 1892

Bryodrilus spp.

Genus Buchholzia Michaelsen, 1886

Buchholzia spp.

Genus Cernosvitoviella Nielsen & Christensen, 1959

Cernosvitoviella christenseni Dash, 1970

Cernosvitoviella pusilla Nurminen, 1973

Genus Cognettia Nielsen & Christensen, 1959

Cognettia spp. -- in banks of rivers and in lakes, also upper intertidal or tidal areas

Genus Fridericia Michaelsen, 1889

Fridericia canadensis Dash, 1972

Genus Guaranidrilus Černosvitov, 1937

Guaranidrilus oregonensis Coates & Diaz, 1988

Genus Hemienchytraeus Černosvitov, 1934

Hemienchytraeus spp. -- possible in wet soils, in southern regions

Genus Henlea Michaelsen, 1889

Henlea urbanensis Welch, 1914

Genus Lumbricillus Ørsted, 1844

Lumbricillus arenarius (Michaelsen, 1889) sensu Knöllner, 1935

Lumbricillus charae (Tynen, 1970)

Lumbricillus franciscanus Eisen, 1904 -- incertae sedis

Lumbricillus kalatdlitus Nurminen, 1970

Lumbricillus rivalis (Levinsen, 1883) augm. Ditlevsen, 1904

[= L. rutilus Welch, 1914]

Lumbricillus santaeclarae Eisen, 1904

Genus Marionina Michaelsen 1890, in Pfeffer, 1890

Marionina forbesae F. Smith & Welch, 1913 -- incertae sedis

Genus Mesenchytraeus Eisen, 1878

Mesenchytraeus armatus (Levinsen, 1884)

Mesenchytraeus franciscanus Eisen, 1904

Mesenchytraeus fuscus Eisen, 1904

Mesenchytraeus fuscus inermis Eisen, 1904

Mesenchytraeus gelidus Welch, 1916

Mesenchytraeus kincaidi Eisen, 1904

Mesenchytraeus pedatus Eisen, 1904

Mesenchytraeus rhithralis Healy & Fend, 2002

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Genus Propappus Michaelsen, 1905

Propappus volki Michaelsen, 1922 - i -

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Footnotes

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 Lastockin's (1921) paper has erroneously been recorded as '1924' in numerous publications over the years].

² On earlier versions of this website (< April 2007), and also presented in a workshop workbook (Wetzel et al. 2006), the above list of North American freshwater microdriles included an interim subfamilial ranking, with [at that time of posting on the website] all 'former' Naididae placed in sub-familial groupings. This interim ranking was suggested by Dr. Gustavsson (pers. comm. to Wetzel, August 2005), pending a published opinion by the ICZN on the application by Erséus et al. (2005), and/or until such time that the monophyly of these groups could be tested definitively. Interim groupings (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 recently published (Timm and Grimm 2005) an exhaustive review of the taxon Homochaeta naidina Bretscher, 1896; I paraphrase here the abstract from that paper: Homochaeta naidina has never been redescribed. No type material exists for this taxon, and all subsequent material identified as this taxon, when available for review [by Timm and Grimm], had been misidentified. The original description by Bretscher likely was based upon different, immature specimens in the families Naididae and Tubificidae [probably Uncinais uncinata (Ørsted, 1842) and Bothrioneurum vejdovskyanum 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; to designate a neotype from [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.

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

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

⁹ Chaetogaster cristallinus could easily be confused with Chaetogaster diaphanus. Ch. cristallinus has a median prostomial incision (easily overlooked / unobservable due to orientation of mounted specimens), and {perhaps?} smaller / shorter chaetae than Ch. diaphanus; see comments in Kathman and Brinkhurst (1998: p. 42). This incision occurs also in typical Ch. diaphanus; see Poddubnaya (1966). 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). Tarmo stated 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, Tarmo 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 Tasserkidrilus harmani was transferred to Varichaetadrilus by Timm (2006b; citation provided, below).

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

¹³ The 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 his name was without the umlaut in the original description for P. bavaricus (Zoologischer Anzieger 42: 559. [1913]), but had been accidentally 'created' later by somebody, and 'loyally' copied by subsequent writers [including MJW]. This will be corrected in Supplementum Quartum of Nomenclatura Oligochaetologica (still in prep.).

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

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., 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., 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]. A downloadable PDF of this paper is available free (through open access) HERE.

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

Fend, S.V. 2009. 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., 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 D.L. Gustafson. 2001. A new lumbriculid genus and species from North America (Clitellata, Lumbriculidae). Hydrobiologia 463: 13-22.

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., 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.

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. [www.elsevier.com] 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. [www.elsevier.com] USD$ 139.95 / 93.95 Euro.

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. ]

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 Thiere, Jena, 163: 377-431.

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

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.

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].

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. 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. As of 01 March 2013, all inquiries regarding this guide should be addressed to: Dr. R.D. Kathman, 5125 Rock Bridge Lane, Thompsons Station, Tennessee 37179 USA, or via E-mail [ ** R.Deedee.Kathman{AT}tn.gov ]; cost: USD$55.00 (inclusive of postage to addresses in North America; payment via personal / business check or money order only; no purchase orders or charge card sales).

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.

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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).

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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.

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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 2008 (citation below)].

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.

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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. in prep. Nomenclatura oligochaetologica. A catalogue of names, descriptions and type specimens of the Oligochaeta. Second Edition.

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

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Spencer, D.R. 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.

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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. 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].

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.

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

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

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

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

Wetzel, M.J., S.V. Fend, K.A. Coates, R.D. Kathman, and S.R. Gelder. 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.}

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. [to order, contact Aleta Holt via telephone, at (217) 897-1274; cost: USD$43.00 (postage included; discount on multiple copy purchases)].

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Suggested citation for this electronic web page: Wetzel, M.J., R.D. Kathman, S.V. Fend, and K.A. Coates. 2013. Classification and checklist of the freshwater oligochaetes occurring in North America north of Mexico. 04 March.
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