Enchytraeids of North America

Clitellate Annelida Occurring in the United States and Canada – Order Enchytraeida

Families Enchytraeidae and Propappidae

Compiled by Kathryn A. Coates, Jan M. Locke, Brenda M. Healy*^†, and Mark J. Wetzel

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*Our dear friend and colleague, Brenda Healy, passed away on 5 March 2006 in Ireland. A memoriam celebrating her life was published in the proceedings of our 10th International Symposium on Aquatic Oligochaeta (Acta Hydrobiologica Sinica, vol. 31: 1-9) in mid-2008, and is also available as a PDF download. We miss her very much.

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Introduction

This electronic document provides a current checklist of North American species of Enchytraeidae and Propappidae (Annelida, Clitellata, Oligochaeta); marine, estuarine, and freshwater species are included. The list is not intended to be a taxonomic revision; nomenclature presented herein reflects the majority opinion of that published in the scientific literature. The scope of this list was initiated in 1989 by two of the authors (Coates and Wetzel) through their association with and guidance from the Committees on Common and Scientific Names of Aquatic Invertebrates sponsored by the American Fisheries Society (AFS-CNAI) and the North American Benthological Society (NABS), now the Society for Freshwater Science.

Objective

This list reflects a thorough and continuing survey of the historical and current literature, reflected in the modification date at the end of this electronic document. In the future, we hope to resolve errors and omissions that have inevitably occurred, then publish this document in the peer-reviewed literature. Your comments and contributions are welcome.

Taxonomic Coverage

Currently, more than 830 species of annelids representing 27 families of oligochaetous Clitellata, Aphanoneura, Branchiobdellae, Acanthobdellae, and Hirudinida are recognized as occurring in the U.S. and Canada; these include both native and introduced species. Certainly, continued systematic studies will add new species, identify previously recognized species as synonyms, and contribute to revisions within the family.

Area of Coverage

This list includes all species enchytraeids and propappids from the United States and Canada that live in (1) fresh waters, including wetlands, (2) marine waters from shoreline habitats out to a depth of 200 m on the continental shelf or a distance of 125 km offshore on the continental shelf where the depth is less than 200 m, (3) estuaries, and (4) terrestrial habitats (e.g., gardens, woodlands, mountains, deserts, and caves). Annelids from the Arctic Ocean and the northern Gulf of Mexico, south, to the mouth of the Rio Grande are included. Grainger’s (1971) definition of the Arctic Ocean is followed, which, to the west, includes the waters north of the Bering Strait, and to the east includes all waters west of the eastern end of Hudson Strait (including Hudson Bay, Ungava Bay, Frobisher Bay, and Cumberland Sound) and waters north of the Arctic Circle in Davis Strait. Annelids known only from offshore islands (e.g., Greenland, Iceland, Bermuda, the Bahamas, and the West Indies) are excluded. A lack of studies and/or the absence of a good compilation of literature on the annelids of many regions in the area may have resulted in the exclusion of numerous taxa. Species known from the periphery of the area of coverage, e.g., Mexico (including Baja California), are likely to be found to have populations extending into the study area.

For the eastern Pacific region, published records of annelids from waters along Canadian and U.S. shores and coastal islands, from the Aleutian Islands to the Mexico – United States boundary, are listed. Hawai’i and the Pacific Trust Territories are excluded from the study area for two major reasons: (1) their faunas are of Indo-Pacific origin whereas the northeast Pacific Coast fauna is largely indigenous or Holarctic, and (2) the preceding AFS endeavors excluded Hawai’i and the Trust Territories from its study area. Annelids occurring in Hawai’i may be included in later editions.

Common Names

The AFS-CNAI acknowledges the need to recognize common names that reflect broad current usage; adopt appropriate names from the rich and colorful vernacular names that exist; and develop descriptive names when desirable, based on AFS guidelines and principles. The Committee expects that the selected vernacular names and newly designated names of this list will be widely used after publication.

The AFS-CNAI Annelida Subcommittee recognizes that common names of annelids are used primarily by commercial wormers and amateur collectors, for aquarium specimens, in ecological studies, in government listings of threatened and endangered species, and in popular writing. Common names of annelids are uncommon and usually absent from scientific writings, however, especially those focusing on freshwater species. The Annelida Subcommittee has taken the position that standardized and uniform common names should always be referenced with appropriate scientific names, not used as substitutes for them. Proper identification and recording of some species are important because of their high economic value. The worm and leech bait industries alone have a total value of several million dollars in southern Ontario, Canada. The presence of different common names across separate areas of a species’ geographic range creates communication difficulties. Similarly, a single common name used in several places for diverse species can cause confusion. Therefore, because common names have been assigned to only a few species to date, only common names recognized for some of the higher taxa will be included in this electronic posting (e.g., Family Enchytraeidae – Potworms). The Annelida Subcommittee will not be proposing common names for annelid species that do not presently have one or more common names established in the literature.

Plan of the List

The list of annelids is presented in a natural or phyletic sequence of classes and orders, as this natural sequence is currently understood, with families, genera, and species within each order arranged alphabetically. The higher classification of Annelida is complex, controversial, and frequently revised. The classification adopted here is conservative – it reflects only the consistently supported parts of the phylogeny of annelids (Timm, 1981; Brinkhurst, 1982; Kasprzak, 1984; Jamieson, 1988; Holt, 1989; Gelder 1996; other recent papers to be added here) and is not resolved where relationships are unsupported by data; alphabetical listings predominate. Undoubtedly, the hierarchical status of groups will change in the future – resulting from ongoing studies of the relationships among clitellates as well as those between all clitellate and aclitellate groups of annelids.

Authors and dates of establishment for scientific names of species are included in this list. These are commonly needed by persons who may not have ready access to the original literature. Use of the authority’s names reflects a current interpretation of the International Code of Zoological Nomenclature (1999). In line with that code, the name(s) of the author(s) and date of publication follow the specific name; if the species was originally described in another genus, the name of the author and date appear in parentheses. Some synonyms have been noted [= name] of North American names synonymized with European taxa. Frequently, the North American names are re-erected subsequent to detailed comparative studies of North American specimens.

In 2003, Rüdiger Schmelz published an extensive revision of the species in the genus Fridericia, so you are encouraged to secure a copy of that book for a current perspective on the species in this genus (taxonomy, nomenclature, morphology, biochemistry, valid species, nomina dubia, species transferred to other genera, an extensive reference section, and numerous figures).

Occurrences of species by province and state will not be provided here; an annotated list of species, including common names, is currently being prepared for publication in a print medium.

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The following list is incomplete at this time; additional species will be added soon.

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Phylum Annelida

• Class Clitellata
  • Order Enchytraeida
    • Family Enchytraeidae – Potworms
      • Genus Achaeta (Vejdovský, 1877)
        • Achaeta camerani (Cognetti, 1899)
        • Achaeta eiseni (Vejdovský, 1877)
        • Achaeta silvatica (Nurminen, 1973)
      • Genus Barbidrilus (Loden & Locy, 1980)
        • Barbidrilus paucisetus (Loden & Locy, 1980)
      • Genus Bryodrilus (Ude, 1892)
        • Bryodrilus arctica (Bell, 1962)
        • Bryodrilus ehlersi (Ude, 1892)
        • Bryodrilus novaescotiae (Bell, 1962)
        • Bryodrilus parvus (Nurminen, 1970)
        • Bryodrilus parvus kananaskis (Dash, 1970)
      • Genus Buchholzia (Michaelsen, 1886)
        • Buchholzia appendiculata (Buchholz, 1862)
        • Buchholzia fallax (Michaelsen, 1887)
      • Genus Cernosvitoviella (Nielsen & Christensen, 1959)
        • Cernosvitoviella atrata (Bretscher, 1903)
        • Cernosvitoviella celere (Nurminen, 1973)
        • Cernosvitoviella christenseni (Dash, 1970)
        • Cernosvitoviella immota (Knöllner, 1935)
        • Cernosvitoviella pusilla (Nurminen, 1973)
        • Cernosvitoviella tatrensis (Kowalewski, 1916)
      • Genus Cognettia (Nielsen & Christensen, 1959) (Invalidated)(This genus is now a junior synonym of Chamaedrilus)**
      • Genus Chamaedrilus (Friend, 1913)
        • Chamaedrilus floridae (Healy, 1996)
        • Chamaedrilus glandulosa (Michaelsen, 1888)
        • Chamaedrilus sphagnetorum (Vejdovský, 1878)
      • Genus Enchytraeus (Henle, 1837)
        • Enchytraeus albidus (Henle, 1837)
        • Enchytraeus buchholzi (Vejdovský, 1879)
        • Enchytraeus capitiatus (von Bulow, 1957)
        • Enchytraeus citrinus (Eisen, 1904) (incertae sedis)
        • Enchytraeus fragmentosus (Bell, 1959) (incertae sedis)
        • Enchytraeus gillettensis (Welch, 1914)
        • Enchytraeus kincaidi (Eisen, 1904)
        • Enchytraeus metlakatlensis (Eisen, 1904) (incertae sedis)
        • Enchytraeus minutus (Nielsen & Christensen, 1961)
        • Enchytraeus multiannulatoides (Altman, 1936)
        • Enchytraeus multiannulatus (Altman, 1936)
        • Enchytraeus rupus (Coates, 1980)
        • Enchytraeus saxicola (Eisen, 1904) (incertae sedis)
        • Enchytraeus syracussus (Dash & Mitchell, 1981)
      • Genus Fridericia (Michaelsen, 1889)
        • Fridericia agilis (Smith, 1895)
        • Fridericia agricola (Moore, 1895)
        • Fridericia berkeleyensis (Bell, 1936)
        • Fridericia bulboides (Nielsen & Christensen, 1959)
        • Fridericia bulbosa (Rosa, 1887) (nomen dubium) (= F. alba (Moore, 1895) (nomen dubium)) (= F. parva (Moore, 1895) (nomen dubium))
        • Fridericia douglasensis (Welch, 1914)
        • Fridericia firma (Smith & Welch, 1913)
        • Fridericia fuchsi (Eisen, 1904) (nomen dubium)
        • Fridericia galba (Hoffmeister, 1843)
        • Fridericia harrimani (Eisen, 1904) (nomen dubium)
        • Fridericia johnsoni (Eisen, 1904) (nomen dubium)
        • Fridericia kalffi (Nurminen, 1973) (nomen dubium)
        • Fridericia longa (Moore, 1895) (nomen dubium)
        • Fridericia losangelensis (Bell, 1936)
        • Fridericia macgregori (Eisen, 1904)
        • Fridericia oconeensis (Welch, 1914)
        • Fridericia paroniana (Issel, 1904) (= Fridericia vancouverensis (Dash, 1983))
        • Fridericia perrieri (Vejdovský, 1878)
        • Fridericia popofiana (Eisen, 1904) (nomen dubium)
        • Fridericia pretoriana (Stephenson, 1930) (= Fridericia caprensis (Bell, 1947) (augm. (Nielsen & Christensen, 1959))
        • Fridericia ratzeli (Eisen, 1872) (= F. californica (Eisen, 1904)) (= F. canadensis (Dash, 1972)) (= F. macgregori (Eisen, 1904)) (= F. sacculata (Bell, 1936)) (= F. tenera (Smith & Welch, 1913)) (= F. whatcomae (Altman, 1936))
        • Fridericia santaebarbarae (Eisen, 1904) (nomen dubium)
        • Fridericia santaerosae (Eisen, 1904) (nomen dubium)
        • Fridericia silvestris (Leidy, 1882) (nomen dubium)
        • Fridericia sima (Welch, 1914)
        • Fridericia striata (Levinsen, 1884) (Fridericia syracussa (Dash & Mitchell, 1981)) (nov comb. Enchytraeus syracussus)
      • Genus Grania (Southern, 1913)
        • Grania americana (Kennedy, 1966)
        • Grania atlantica (Coates & Erséus, 1985)
        • Grania bermudensis (Lasserre & Erséus, 1976)
        • Grania hylae (Locke & Coates, 1999)
        • Grania incerta (Coates & Erséus, 1980)
        • Grania laxartus (Locke & Coates, 1999)
        • Grania levis (Coates & Erséus, 1985)
        • Grania longiducta (Erséus, & Lasserre, 1976)
        • Grania monospermatheca (Erséus, & Lasserre, 1976)
        • Grania paucispina (Eisen, 1904)
        • Grania reducta (Coates & Erséus, 1985)
      • Genus Guaranidrilus (Cernosvitov, 1937)
        • Guaranidrilus europeus (Healy, 1979)
        • Guaranidrilus oregonensis (Coates & Diaz, 1988)
      • Genus Hemienchytraeus (Cernosvitov, 1934)
        • Hemienchytraeus bifurcatus (Nielsen & Christensen, 1959)
        • Hemienchytraeus stephensoni (Cognetti, 1927)
      • Genus Henlea (Michaelsen, 1989)
        • Henlea californica (Eisen, 1904)
        • Henlea californica monticola (Eisen, 1904)
        • Henlea ehrhorni (Eisen, 1904)
        • Henlea eiseni (Bell, 1942)
        • Henlea glabra (Altman, 1936)
        • Henlea helenae (Eisen, 1904)
        • Henlea heleotropha (Stephenson, 1922) (augm. (Nielsen & Christensen, 1959))
        • Henlea moderatoidea (Altman, 1936)
        • Henlea nasuta (Eisen, 1878)
        • Henlea ochracea (Eisen, 1878) (= H. arctica (Welch, 1919)) (= H. moderata (Welch, 1914)) (= H. tubulifera (Welch, 1914))
        • Henlea perpusilla (Friend, 1911) (augm. (Cernosvitov, 1937))
        • Henlea scharffi (Southern, 1910)
        • Henlea udei (Eisen, 1904) (augm. (Holmquist, 1968)) (= Bryohenlea nuda (Bell, 1962))
        • Henlea urbanensis (Welch, 1914)
        • Henlea ventriculosa (d’Udekem, 1854)
        • Henlea welchi (Bell, 1942)
        • Henlea yukonensis (Tynen & Coates, 1991)
      • Genus Lumbricillus (Ørsted, 1844)
        • Lumbricillus annulatus (Eisen, 1904)
        • Lumbricillus arenarius (Michaelsen, 1889) (sensu Knöllner, 1935)
        • Lumbricillus belli (Tynen, 1969)
        • Lumbricillus charae (Tynen, 1970)
        • Lumbricillis curtus (Coates, 1981)
        • Lumbricillus franciscanus (Eisen, 1904) (incertae sedis)
        • Lumbricillus franciscanus borealis (Eisen, 1904) (incertae sedis)
        • Lumbricillus franciscanus unalaskae (Eisen, 1904) (incertae sedis)
        • Lumbricillus kalatdlitus (Nurminen, 1970)
        • Lumbricillus lineatus (O.F. Müller, 1774) (= L. agilis (Moore, 1905))
        • Lumbricillus merriami (Eisen, 1904) (incertae sedis)
        • Lumbricillus merriami elongatus (Eisen, 1904)
        • Lumbricillus mirabilis (Tynen, 1969)
        • Lumbricillus pagenstecheri (Ratzel, 1869) (= L. ritteri (Eisen, 1904)) (= L. georgensis (Tynen, 1969))
        • Lumbricillus qualicumensis (Tynen, 1969)
        • Lumbricillus rivalis (Levinsen, 1883) (augm. (Ditlevesen, 1904)) (= L. rutilus (Welch, 1914))
        • Lumbricillis rupertensis (Coates, 1981)
        • Lumbricillus santaeclarae (Eisen, 1904)
        • Lumbricillis tsimpseanis (Coates, 1981)
        • Lumbricillis tuba (Stephenson, 1911)
      • Genus Marionina (Michaelsen, in Pfeffer, 1890)
        • Marionina achaeta (Lasserre, 1964)
        • Marionina alaskae (Eisen, 1904) (incertae sedis)
        • Marionina appendiculata (Nielsen & Christensen, 1959)
        • Marionina argentea (Michaelsen, 1889)
        • Marionina canadensis (Dash, 1970)
        • Marionina charlottensis (Coates, 1980)
        • Marionina craggi (Dash, 1970)
        • Marionina diazi (Coates & Erséus, 1985)
        • Marionina dirksi (Bell, 1942)
        • Marionina eiseni (Cernosvitov, 1937) (species inquirende)
        • Marionina forbesae (Smith & Welch, 1913) (incertae sedis)
        • Marionina glandulifera (Jansson, 1960)
        • Marionina klaskisharum (Coates, 1983)
        • Marionina neroutsensis (Coates, 1980)
        • Marionina nevisensis (Righi & Kanner, 1979)
        • Marionina paludis (Healy, 1994)
        • Marionina preclitellochaeta (Nielsen & Christensen, 1963)
        • Marionina sjaelandica (Nielsen & Christensen, 1961)
        • Marionina southerni (Cernosvitov, 1937)
        • Marionina spartinae (Healy, 1994)
        • Marionina spicula (Leuckart, 1847)
        • Marionina subterranea (Knöllner, 1935)
        • Marionina tubifera (Nielsen & Christensen, 1959)
        • Marionina vancouverensis (Coates, 1980)
        • Marionina waltersi (Healy, 1994)
        • Marionina welchi (Lasserre, 1971)
      • Genus Mesenchytraeus (Eisen, 1878)
        • Mesenchytraeus altus (Welch, 1917)
        • Mesenchytraeus americanus (Bell, 1942)
        • Mesenchytraeus antaeus (Rota & Brinkhurst, 2000)
        • Mesenchytraeus arcticus (Bell, 1962)
        • Mesenchytraeus argentatus (Nurminen, 1973)
        • Mesenchytraeus armatus (Levinsen, 1884)
        • Mesenchytraeus armatus kananaskis (Dash, 1970)
        • Mesenchytraeus atriaphorus (Altman, 1936)
        • Mesenchytraeus beringensis (Eisen, 1904)
        • Mesenchytraeus chromophorus (Altman, 1936)
        • Mesenchytraeus diplobulbosus (Bell, 1949)
        • Mesenchytraeus eastwoodi (Eisen, 1904)
        • Mesenchytraeus fontinalis (Eisen, 1904)
        • Mesenchytraeus franciscanus (Eisen, 1904)
        • Mesenchytraeus fuscus (Eisen, 1904)
        • Mesenchytraeus fuscus inermis (Eisen, 1904)
        • Mesenchytraeus gelidus (Welch, 1916)
        • Mesenchytraeus grandis (Eisen, 1904)
        • Mesenchytraeus hamiltoni (Healy, 1996)
        • Mesenchytraeus harrimani (Eisen, 1904)
        • Mesenchytraeus johanseni (Welch, 1919)
        • Mesenchytraeus kincaidi (Eisen, 1904)
        • Mesenchytraeus macnabi (Bell, 1942)
        • Mesenchytraeus maculatus (Eisen, 1904)
        • Mesenchytraeus magnus (Altman, 1936)
        • Mesenchytraeus minimus (Altman, 1936)
        • Mesenchytraeus monothecatus (Bell, 1945)
        • Mesenchytraeus nanus (Eisen, 1904)
        • Mesenchytraeus obscurus (Eisen, 1904)
        • Mesenchytraeus orcae (Eisen, 1904)
        • Mesenchytraeus pedatus (Eisen, 1904)
        • Mesenchytraeus penicillus (Eisen, 1904)
        • Mesenchytraeus setchelli (Eisen, 1904)
        • Mesenchytraeus solifugus (Emery, 1898) (ice worm, glacier ice worm)
        • Mesenchytraeus unalaskae (Eisen, 1904)
        • Mesenchytraeus vegae (Eisen, 1904)
      • Genus Randidrilus (Coates & Erséus, 1985)
        • Randidrilus codensis (Lasserre, 1971)
        • Randidrilus quadrithecatus (Coates & Erséus, 1985)
      • Genus Stephensoniella (Cernosvitov, 1934)
        • Stephensoniella marina (Moore, 1902)
        • Stephensoniella sterreri (Lasserre & Erséus, 1976)
        • Stephensoniella trevori (Coates, 1980)
    • Family Propappidae
      • Genus Propappus (Michaelsen, 1905)
        • Propappus volki (Michaelsen, 1922) (Considered introduced to North America)

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**Colleagues 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, Cognettia 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.

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Literature Cited (Incomplete)