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, 03Dec 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.
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, systematics, and nomenclature.
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, ~272 nominal species of freshwater (microdrile) oligochaetes representing 85 genera in seven 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.
Nomenclature
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.
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 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), Reynolds and Wetzel (2004, 2008, 2012), and Reeves, Reynolds and Wetzel (2024).
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
Class Clitellata Michaelsen, 1919
Subclass Oligochaeta Grube, 1850
– – – – – Order Haplotaxida – – – – –
Brinkhurst and Jamieson, 1971
Haplotaxis emissarius (Forbes, 1890)
Haplotaxis forbesi Smith, 1918
Notes:
> the genera Metataxis and Tiguassu were removed from this family by Brinkhurst (1988).
> see Martin, Fend, Martinsson, Klinth, Torii, and Erséus, 2024 regarding recognized species in the genus Haplotaxis, and an integrative revision of the family Haplotaxidae.
– – – – – Order Haplotaxoidida – – – – –
Martin, Fend, Martinsson, Klinth, Torii, and Erséus, 2024
Haplotaxoides tehama Fend, 2024 [distr: U.S. – CA: Tehama Co., Shasta Co. ]
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)
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.]
Lumbriculus genitosetosus Holmquist, 1976
Lumbriculus inconstans (F. Smith, 1895)
Lumbriculus variegatus variegatus (Müller, 1774)
Martinidrilus carolinensis Fend & Lenat, 2007 [ distr.: U.S. – NC, VA ]
Pararhynchelmis texana Fend 2023 [distr.: U.S. – TX: Hays Co. ]
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
Rhynchelmis (Sutroa) aleutensis Fend, 2005
Rhynchelmis brooksi Holmquist, 1976
Rhynchelmis (Sutroa) diespluviae Fend, 2014
Rhynchelmis (Rhynchelmoides) elrodi Smith & Dickey, 1918
Rhynchelmis (Sutroa) gilensis Fend & Brinkhurst, 2000
Rhynchelmis (Rhynchelmoides) glandula Altman, 1936
Rhynchelmis (Sutroa) gustafsoni Fend & Brinkhurst, 2000
Rhynchelmis (Sutroa) klamathensis Fend, 2014
Rhynchelmis (Sutroa) monsserratus Fend & Brinkhurst, 2000
Rhynchelmis rostrata (Eisen, 1888)
Rhynchelmis (Rhynchelmoides) saxosa Fend & Brinkhurst, 2000
Rhynchelmis (Sutroa) utahensis Fend & Brinkhurst, 2000
Rhynchelmis (Sutroa) yakimorum Fend & Brinkhurst, 2000
Rhynchelmis (Rhynchelmoides) bolinensis Fend & Lenat, 2010 [ distr.: U.S. – NC]
Rhynchelmis (Rhynchelmoides) croatanensis Fend & Lenat, 2010 [ distr.: U.S. – NC ]
Stylodrilus californianus Rodriguez, 1996
Stylodrilus coreyi Rodriguez, Fend & Lenat, 2014
Stylodrilus heringianus Claparède, 1862 – i –
Stylodrilus sovaliki Holmquist, 1976
Stylodrilus wahkeenensis Rodriguez & Coates, 1996
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 – – – – –
Branchiodrilus ? cf. hortensis (Stephenson, 1910) 21
Chaetogaster limnaei von Baer, 1827
Chaetogaster setosus Svetlov, 1925
Amphichaeta leydigi Tauber, 1879
Amphichaeta raptisae (Chapman, 1981)
Piguetiella michiganensis Hiltunen, 1967
Specaria hellei (Brinkhurst, 1971)
Specaria josinae (Vejdovský, 1883)
Stylaria fossularis Leidy, 1852
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)
?? [Allonais paraguayensis (Michaelsen, 1905)] 3
?? Allonais pectinata (Stephenson, 1910) 3
note that another species in this genus, Slavina evelinae (previously known from
Central and South America, has been identified by MJW based on specimens
the global distribution of known Slavina species is in prep.
Vejdovskyella intermedia (Bretscher, 1896)
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
Stephensoniana trivandrana (Aiyer, 1926)
Bratislavia dadayi (Michaelsen, 1905)
[= Bratislavia unidentata (Harman, 1973) – See Righi and Hamoui (2002)]
Paranais frici Hrabě, 1941
Paranais grandis Harman, 1977
Paranais litoralis (Müller, 1784)
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
Aulodrilus limnobius Bretscher, 1899
Aulodrilus paucichaeta Brinkhurst & Barbour, 1985
Aulodrilus pigueti Kowalewski, 1914
Aulodrilus pluriseta (Piguet, 1906) /-/ Aulodrilus japonicus Yamaguchi, 1953 8
Haber speciosus (Hrabě, 1931)
Ilyodrilus frantzi Brinkhurst, 1965
Ilyodrilus perrieri Eisen, 1879
Ilyodrilus templetoni (Southern, 1909)
Isochaetides curvisetosus (Brinkhurst & Cook, 1966)
Isochaetides freyi (Brinkhurst, 1965)
Kopilrok flexipenis Holmquist, 1985 – AK
Kopilrok sagavanirktoki Holmquist, 1985 – AK
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
Potamothrix bedoti (Piguet, 1913) – i (?) –
?? Potamothrix hammoniensis (Michaelsen, 1901) 7– i –
Potamothrix moldaviensis Vejdovský & Mrázek, 1902 – i –
Potamothrix vejdovskyi (Hrabě, 1941) – i –
Psammoryctides californianus Brinkhurst, 1965
Psammoryctides convolutus Loden, 1978
Spirosperma carolinensis Brinkhurst, 1965
Spirosperma ferox Eisen, 1879 – i –
Spirosperma nikolskyi (Lastočkin & Sokolskaya, 1935)
Teneridrilus columbiensis (Brinkhurst & Diaz, 1985)
Teneridrilus flexus Erséus & Hiltunen, 1990
Teneridrilus mastix (Brinkhurst, 1978)
Tubifex ignotus (Štolc, 1886)
Tubifex nerthus Michaelsen, 1908
?? Tubifex newaensis (Michaelsen, 1903) 7
Tubifex tubifex (Müller, 1774)
Varichaetadrilus fulleri Brinkhurst & Kathman, 1983
Varichaetadrilus harmani (Loden, 1979) 11
Varichaetadrilus minutus (Brinkhurst, 1965)
Varichaetadrilus pacificus (Brinkhurst, 1981)
Subfamily Telmatodrilinae Eisen, 1879
Telmatodrilus vejdovskyi Eisen, 1879
Alexandrovia onegensis Hrabě, 1962
Bothrioneurum vejdovskyanum Štolc, 1888
Branchiura sowerbyi Beddard, 1892 – i –
Rhizodrilus lacteus F. Smith, 1900
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 ]
[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]
Crustipellis tribranchiata (Harman, 1970) 6 (a)
Trieminentia corderoi (Harman, 1970) 6 (b)
Opistocysta flagellum (Leidy, 1880) – species inquirenda 6 (c)
Parvidrilus strayeri Erséus, 1999
– – – – – Order Enchytraeida – – – – –
This listing includes genera and species collected from terrestrial, limicolous, semi-aquatic, and aquatic habitats.
Family Enchytraeidae Vejdovský, 1879
Achaeta spp.
Barbidrilus paucisetus Loden & Locy, 1980
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
Cernosvitoviella pusilla Nurminen, 1973
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
Hemifridericia spp.
Genus Henlea Michaelsen, 1889
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
Marionina sp. 20 MN, WI
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
Allolobophora chlorotica (Savigny, 1826)
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
Parergodrilus heideri Reisinger, 1925 20 MN, WI
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 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, 1828, not 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.
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Hartmann G.L. 1821. Beyträge zur Geschichte der Fadenwürmer, nebst Beschreibung einer bisher mit ihnen verwechselten Art von Regenwurm, Lumbricus Gordioides. Der Schweizerischen Gesellschaft für die gesammten Naturwissenschaften vorgelesen den 28 Junius 1819 von G. L. Hartmann, Neue Alpina: eine Schrift der Schweizerischen Naturgeschichte Alpen- und Landwirthschaft gewiedmet. Winterthur 1821: 1: 32–50.
Healy, B. 1989. Preliminary report on the Enchytraeidae (Oligochaeta) of West Florida. – Hydrobiologia 180: 47–56. ^
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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.
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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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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.
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Martin, P., S. Fend, S. Martinsson, M. Klinth, T. Torii, and C. Erséus. 2024. Towards an integrative revision of Haplotaxidae (Annelida: Clitellata) Zoological Journal of the Linnean Society 202, zlae141 [38 pp].^
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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).
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 http://fada.biodiversity.be/group/show/12 [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.
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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. https://doi.org/10.1016/j.pedobi.2013.09.006
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.
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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 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.
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Reeves, W.K., J.W. Reynolds and M.J. Wetzel. 2024. Sparganophilus (Annelida, Oligochaeta, Sparganophilidae) in North America. Sparganophilus (Annelida, Oligochaeta, Sparganophilidae) in North America. Megadrilogica 28(5): 53–80. April. [paper in English; abstracts in English, French, Spanish, German; with 18 colour plates].
Reynolds, J.W. 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.
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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].
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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.
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