Classification and Checklist of the Leeches
(Phylum Annelida: Class Clitellata: Subclass Hirudinida)
occurring in North America north of Mexico

Compiled by Donald J. Klemm1, William E. Moser2, and Mark J. Wetzel3

1 Retired [formerly, U.S. Environmental Protection Agency, Office of Research and Development, National Exposure Research Laboratory, Ecological Exposure Research Division, Ecosystems Research Branch (MS-642), 26 W. Martin Luther King Drive, Cincinnati, Ohio USA.] E-Mail: Don's new email address will be posted here soon.

2 Smithsonian Institution, National Museum of Natural History, Department of Invertebrate Zoology, Museum Support Center, MRC 534, 4210 Silver Hill Road, Suitland, MD 20746 USA. Phone: (301) 238-1761; Fax: (301) 238-1833; E-mail: moserw{AT}

3 Illinois Natural History Survey, Prairie Research Institute at the University of Illinois at Urbana-Champaign, Forbes Natural History Building, 1816 S. Oak Street, Champaign, Illinois 61820 USA. Phone: (217) 244-2108; FAX: (217) 265-4678; E-Mail: mjwetzel{AT}

Introduction. Freshwater leeches (segmented worms) are important components of benthic communities in nearly every freshwater ecosystem. They are widely distributed, and some groups are found in great abundance. Several of the Hirudinea groups have been used for monitoring and detecting changes in water quality and physical habitats. The habitat and water quality requirements as well as the pollution tolerance of some species of freshwater leeches have been documented in the literature by a few investigators. Practical taxonomic keys are now available to species, but many benthic water quality assessment studies still do not treat the leeches adequately because the investigators lack the knowledge and experience in using these keys. The inadequate treatment by some investigators represents a loss of valuable ecological data and information for use in bioassessment of the quality of water resources, water pollution, or other changes in aquatic ecosystems resulting from natural causes or anthropogenic activities of man. We present here a current classification and list of freshwater leeches occurring in North America north of Mexico.

Some of the recent distributional information for leeches 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, is 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. K.A. Coates and Wetzel served on this Committee from 1989 through 2005 (as co-chairs of the clitellate Annelid Subcommittee), and (in collaboration with many colleagues, including Don Klemm, Bill Moser, John Reynolds, Steve Fend, R. Deedee Kathman, and Stuart Gelder) had been preparing the non-clitellate Annelida volume for this series. Dr. Kristian Fauchald (Smithsonian Institution, National Museum of Natural History) had been chairing the polychaete annelid subcommittee). Unfortunately, the AFS CNAI committee has been dormant for several years following the retirement of our enthusiastic and tireless CNAI committee chair, Dr. Turgeon; we are awaiting new direction and support from AFS administrators so we may complete the Annelida volume.

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 leeches and other annelid groups; our efforts to maintain 'currency' in this checklist is dependent upon collaboration between and corroboration among our fellow aquatic biologists.

The checklist. Currently, three orders, eight families, 19 genera, and 81 nominal species of leeches are known to occur in North America north of Mexico. 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 leeches 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 leech specialists, 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 (e.g., geographical, physical, biological, water quality).

Sources of information for species included in this checklist. The freshwater leech fauna of North America has been documented in several major publications (e.g., Davies 1991; Davies and Govedich 2001; Govedich et al. 2010; Klemm 1985, 1990, 1991, 1995; Sawyer 1972, 1986a,b,c), literature cited therein, and more recent publications (some of which are included in the literature cited/references section at the bottom of this website). Please consult these publications for additional (and usually more specific) distributional information for selected taxa. Citations for newly published and otherwise pertinent publications focusing on leech biology, ecology, distribution, taxonomy, and systematics in North America are added to this website periodically.

Classification and checklist of the leeches
occurring in North America North of Mexico

- - Phylum Annelida - -

Class Clitellata
Subclass Hirudinida

Order Acanthobdellida Livanow, 1905

Family Acanthobdellidae Livanow, 1905

    Genus Acanthobdella Grube, 1851
    Acanthobdella peledina Grube, 1851
Order Arhynchobdellida R. Blanchard, 1894

Family Haemopidae Sawyer, 1986

    Genus Haemopis Savigny, 1822
    Haemopis grandis (Verrill, 1874)
    Haemopis plumbea Moore, 1912
    Haemopis kingi Mathers, 1954
    Haemopis lateromaculata Mathers, 1963
    Haemopis marmorata (Say, 1824)
    Haemopis ottorum Wirchansky & Shain, 2010
    Haemopis septagon Sawyer & Shelley, 1976
    Haemopis terrestris (Forbes, 1890)

Family Hirudinidae Whitman, 1886

    Genus Hirudo Linnaeus, 1758
    Hirudo medicinalis Linnaeus, 1758 - I -
Family Macrobdellidae Richardson, 1969

    Genus Macrobdella Verrill, 1872
    Macrobdella decora (Say, 1824)
    Macrobdella ditetra Moore, 1953
    Macrobdella diplotertia Meyer, 1975
    Macrobdella sestertia Whitman, 1886
    Genus Philobdella Verrill, 1874
    Philobdella floridana (Verrill, 1874)
    Philobdella gracilis Moore, 1901

Family Erpobdellidae R. Blanchard, 1894

    Genus Erpobdella de Blainville, 1818
    Erpobdella [= Dina] anoculata (Moore, 1898)
    Erpobdella [= Mooreobdella] bucera Moore, 1949
    Erpobdella [= Dina] dubia (Moore & Meyer, 1951)
    Erpobdella [= Mooreobdella] fervida (Verrill, 1871)
    Erpobdella lahontana Hovingh & Klemm, 2000
    Erpobdella [= Mooreobdella] melanostoma (Sawyer & Shelley, 1976)
    Erpobdella [= Mooreobdella] microstoma (Moore, 1901)
    Erpobdella [= Nephelopsis] obscura (Verrill, 1872)
    Erpobdella [= Dina] parva (Moore, 1912)
    Erpobdella punctata (Leidy, 1870)
    Erpobdella coastalis Sawyer & Shelley, 1976
    Erpobdella [= Mooreobdella] tetragon (Sawyer & Shelley, 1976)

    Genus Motobdella Govedich, Blinn, Keim, & Davies, 1998
    Motobdella [= Erpobdella] montezuma (Davies, Singhal, and Blinn, 1985)
    Motobdella sedonensis Govedich, Blinn, Keim, & Davies, 1998

Family Salifidae Johansson, 1910

    Genus Barbronia Johansson, 1918
    Barbronia weberi R. Blanchard, 1897 - I - (See Rutter and Klemm, 2001)
Order Rhynchobdellida R. Blanchard, 1894

Family Glossiphoniidae Vaillant, 1890

    Genus Actinobdella Moore, 1901
    Actinobdella annectens Moore, 1906
    Actinobdella inequiannulata Moore, 1901
    Genus Alboglossiphonia Lukin, 1976
    Alboglossiphonia heteroclita (Linnaeus, 1761)
    Genus Batracobdella Viguier, 1879
    Batracobdella paludosa (Carena, 1824)
    Genus Glossiphonia Johnson, 1816
    Glossiphonia complanata (Linnaeus, 1758)
    Glossiphonia [= Boreobdella] verrucata (Fr. Müller, 1844)
    Genus Helobdella R. Blanchard, 1896
    Helobdella austinensis Kutschera, Langguth, Kuo, D.A. Weisblat & M. Shankland, 2013.
    Helobdella bowermani Moser, Fend, Richardson, Hammond, Lazo-Wasem, Govedich, & Gullo, 2013
    Helobdella californica Kutschera, 1988
    Helobdella [= Gloiobdella] elongata (Castle, 1900)
    Helobdella fusca (Castle, 1900)
    Helobdella lineata (Verrill, 1874)
    Helobdella modesta (Verrill, 1872)
    Helobdella papillata Moore, 1952 1
      [commonly misidentified as H. triserialis; see footnote]
    Helobdella robusta Shankland, Bissen & Weisblat, 1992
    Helobdella stagnalis (Linnaeus, 1758)
    Helobdella transversa Sawyer, 1972
    Genus Marvinmeyeria Soos, 1969
    Marvinmeyeria lucida (Moore, 1954)
    Genus Placobdella R. Blanchard, 1893
    Placobdella ali Hughes & Siddall, 2007
    Placobdella appalachiensis Moser & Hopkins, 2014
    Placobdella [= Oligobdella] biannulata (Moore, 1900)
    Placobdella burresonae Siddall & Bowerman, 2006
    Placobdella [= Desserobdella] cryptobranchii (Johnson & Klemm, 1977)
    Placobdella hollensis (Whitman, 1892)
    Placobdella [= Desserobdella] michiganensis (Sawyer, 1972)
    Placobdella kwetlumye Oceguera-Figueroa, Kvist, Watson, Sankar, Overstreet & Siddall, 2010
    Placobdella montifera (Moore, 1906)
    Placobdella multilineata Moore, 1953
    Placobdella nuchalis Sawyer & Shelley, 1976
    Placobdella ornata (Verrill, 1872)
    Placobdella papillifera (Verrill, 1872)
    Placobdella parasitica (Say, 1824)
    Placobdella pediculata Hemingway, 1908
    Placobdella [= Desserobdella] phalera (Graf, 1899)
    Placobdella [= Desserobdella] picta (Verrill, 1872)
    Placobdella sophieae Oceguera-Figueroa, Kvist, Watson, Sankar, Overstreet & Siddall, 2010
    Placobdella translucens Sawyer & Shelley, 1976
    Genus Theromyzon Phillippi, 1867
    Theromyzon bifarium Oosthuizen & Davies, 1993
    Theromyzon maculosum (Rathe, 1862)
    Theromyzon rude (Baird, 1863)
    Theromyzon tessulatum (O.F. Müller, 1774)
    Theromyzon trizonare Davies & Oosthuizen, 1993
Family Piscicolidae Johnston, 1865

    Genus Cystobranchus Diesing, 1859
    Cystobranchus [= Gonimosobdella] klemmi (Williams & Burreson, 2005)
    Cystobranchus mammillatus (Malm, 1863)
    Cystobranchus meyeri Hayunga & Grey, 1976
    Cystobranchus salmositicus (Meyer, 1946) [= Piscicola salmositica]
    Cystobranchus verrilli Meyer, 1940
    Cystobranchus virginicus Hoffmann, 1964
    Genus Myzobdella Leidy, 1851 [= Illinobdella Meyer, 1940]
    Myzobdella lugubris Leidy, 1851
    Myzobdella [= Piscicolaria] reducta (Meyer, 1940)
    Genus Piscicola de Blainville, 1818
    Piscicola geometra (Linnaeus, 1761)
    Piscicola milneri (Verrill, 1871)
    Piscicola punctata (Verrill, 1871)


- I - = Introduced

1 In previous keys (Klemm 1982, 1985, Sawyer 1986b), specimens would fall under the description of Helobdella triserialis (E. Blanchard, 1849). Based upon the original species descriptions and the results of a phylogenetic analysis, Siddall and Borda (2003) differentiated Helobdella papillata (three irregular rows of papillae irrespective of size and pigmentation) from Helobdella triserialis (cephalic transverse banding). Helobdella papillata -- as it is now defined -- is common and widely distributed throughout eastern North America. Helobdella triserialis occurs in South America {Modified from Moser et al. 2006.}

Literature Cited and Selected References

Barta, J.R., and R.T. Sawyer. 1990. Definition of a new genus of glossiphoniid leech and a redescription of the type species, Clepsine picta Verrill, 1972. Canadian Journal of Zoology 68: 1942-1950.

Borda, E., and M.E. Siddall. 2004a. Arhynchobdellida (Annelida: Oligochaeta: Hirudinida): phylogenetic relationships and evolution. Molecular Phylogenetics and Evolution, 30: 213-225.

Borda, E., and M.E. Siddall. 2004b. Review of the evolution of life history strategies and and phylogeny of the Hirudinida (Annelida: Oligochaeta). Lauterbornia, 52: 5-25.

Burreson, E.M., and D.W. Williams. 2005. Resdescription of Cystobranchus virginicus Hoffman, 1964, and Cystobranchus salmositicus (Meyer, 1946) (Hirudinida: Piscicolidae) from freshwater fishes in North America. Comparative Parasitology 72(2): 157-165.

Davies, R.W., R.N. Singhal, and D.W. Blinn. 1985. Erpobdella montezuma (Hirudinoidea: Erpobdellidae), a new species of freshwater leech from North America. Canadian Journal of Zoology 63: 065-969.

Davies, R.W. 1991. Annelida: Leeches, Polychaetes, and Acanthobdellids. Pp. 437-479, In: James H. Thorp and Alan P. Covich (eds.). Ecology and classification of North American freshwater invertebrates. Academic Press, NY.

Davies, R.W., and F.R. Govedich. 2001. Chapter 13. Annelida: Euhirudinea and Acanthobdellidae. Pp. 465-504, In: James H. Thorp and Alan P. Covich (eds.). Ecology and Classification of North American freshwater invertebrates. Second Edition. Academic Press, NY. [superseded by Govedich et al. 2010, citation below].

Davies, R.W., F.R. Govedich, and W.E. Moser. 2008. Leech Parasites of Birds. Pp. 501-514, In: Parasitic Diseases of Wild Birds. C.T. Atkinsen, N.J. Thomas, and D.B. Hunter (eds). Wiley-Blackwell.

Davies, R.W., and J.H. Oosthuizen. 1993. A new species of duck leech from North America formerly confused with Theromyzon rude (Rhynchobdellida: Glossiphoniidae). Canadian Journal of Zoology 71: 770-775.

Govedich, F.R., D. Blinn, P. Keim, and R.W. Davies. 1998. Phylogenetic relationships of three genera of Erpobdellidae (Hirudinoidea), with a description of a new genus, Motobdella, and species, Motobdella sedonensis. Canadian Journal of Zoology 76: 2164-2171.

Govedich, F.R., B.A. Bain, W.E. Moser, S.R. Gelder, R.W. Davies, and R.O. Brinkhurst. 2010. Annelida (Clitellata) Oligochaeta, Branchiobdellida, Hirudinida, and Acanthobdellida. Pages 385-436, In: J.H. Thorp and A.P. Covich (eds). Ecology and classification of North American freshwater invertebrates. Third Edition xiv + 1,021 pp. Academic Press / Elsevier, San Diego, CA. ISBN 978-0-12-374855-3. [] USD$ 139.95 / 93.95 Euro.

Hopkins W.A., W.E. Moser, D.W. Garst, D.J. Richardson, C.I. Hammond, and E.A. Lazo-Wasem. 2014. Morphological and molecular characterization of a new species of leech (Glossiphoniidae, Hirudinida): Implications for the health of its imperiled amphibian host (Cryptobranchus alleganiensis). ZooKeys 378: 83–101. doi: 10.3897/zookeys.378.6545.

Hovingh, P. 1993. Zoogeography and paleozoology of leeches, molluscs, and amphibians in western Bonneville Basin. Journal of Paleolimnology 9: 41-54.

Hovingh, P. 2004. Erpobdella (Dina) parva complex (Annelida: Hirudinea: Arhynchobdellida: Erpobdellidae): additional description of Erpobdella parva, E. dubia, and E. lahontana and taxonomic revision. Hydrobiologia 517: 89-105.

Hovingh, P., and D.J. Klemm. 2000. Erpobdella lahontana (Annelida: Hirudinea: Arhynchobdellida: Erpobdellidae), a new species of freshwater leech from North America. Proceedings of the Biological Society of Washington 113(1): 155-161.

Hovingh, P., W.H. Clark, and J. Keebaugh. 2008. Leeches of the Snake River in Idaho and Oregon: paleodrainage implications of Mooreobdella microstoma. Western North American Naturalist 68(2): 210-224.

Hughes, J.L., and M.E. Siddall. 2007. A new species of leech from the New York metropolitan area. American Museum Novitates 3578: 1-6.

Klemm, D.J. 1972. The leeches (Annelida: Hirudinea) of Michigan. Michigan Academician 4(4): 405-444.

Klemm, D.J. 1977. A review of the leeches (Annelida: Hirudinea) in the Great Lakes region. Michigan Academician 9(4): 397-418.

Klemm, D.J. 1982. Leeches (Annelida: Hirudinea) of North America. Environmental Monitoring and Support Laboratory, U.S. Environmental Protection Agency, Cincinnati, OH 45268. EPA-600/3-82-025. 177 pp.

Klemm, D.J. 1985. Freshwater leeches. Pp. 70-173, In: D.J. Klemm (ed.). A guide to the freshwater Annelida (Polychaeta, Naidid and Tubificid Oligochaeta, and Hirudinea) of North America. Kendall/Hunt Publ. Co., Dubuque, Iowa.

Klemm, D.J. 1990. Hirudinea. Pp. 398-415, In: B.L. Peckarsky, P.R. Fraissinet, M.A. Penton, and D.J. Conklin, Jr. (eds.). Freshwater macroinvertebrates of northeastern North America. Cornell University Press, Ithaca, NY.

Klemm, D.J. 1991. Taxonomy and pollution ecology of the great Lakes Region leeches (Annelida: Hirudinea). Michigan Academician 24: 37-103.

Klemm, D.J. 1995. Identification guide to the freshwater leeches (Annelida: Hirudinea) of Florida and other southern states. Bureau of Surface Water Management, Florida Department of Environmental Protection, 2600 Blair Stone Road, Tallahassee, FL 32399. v + 82 pp. [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 ''Leeches.pdf'' to begin the download; this file is 3.1 MB in size, so will be a very slow download if you are using a dial-up internet connection].

Klemm, D.J., B.A. Daniels, W.E. Moser, and R.J.G. Lester. 2003. Biology of the leech Actinobdella inequiannulata Moore, 1901 (Annelida: Hirudinea: Rhynchobdellida: Glossiphoniidae), parasitic on the white sucker, Catostomus commersoni Lacepède, 1803, and the longnose sucker, Catostomus catostomus Forster, 1773, in Algonquin Provincial Park, Ontario, Canada. Comparative Parasitology 70: 120-127.

Klemm, D.J., D.G. Huggins, and M.J. Wetzel. 1979. Kansas leeches (Annelida: Hirudinea) with notes on distribution and ecology. Technical Publications of the State Biological Survey of Kansas 8: 38-46.

Klemm, D.J., P.A. Lewis, F. Fulk, and J.M. Lazorchak. 1990. Macroinvertebrate field and laboratory methods for evaluating the biological integrity of surface waters. EPA/600/4-90/030. Environmental Monitoring systems Laboratory, U.S. Environmental Protection Agency, Cincinnati, OH 45268.

Kutschera, U. 1988. A new leech species from North America, Helobdella californica nov. sp. (Hirudinea: Glossiphoniidae). Zoologischer Anzeiger 220: 173-178.

Kutschera, U. 2011. The Golden Gate Leech Helobdella californica (Hirudinea: Glossiphoniidae): Occurrence and DNA-based taxonomy of a species restricted to San Francisco. International Review of Hydrobiology 96(3): 286-295.

Kutschera, U., H. Langguth, D.-H. Kuo, D.A. Weisblat, and M. Shankland. 2013. Description of a new leech species from North America, Helobdella austinensis n. sp. (Hirudinea: Glossiphoniidae), with observations on its feeding behaviour. Zoosyst. Evol. 89(2): 239-246.

Light, J.E., and M.E. Siddall. 1999. Phylogeny of the leech family Glossiphoniidae based on mitochondrial gene sequences and morphological data. Journal of Parasitology, 85: 815-823.

McCoy, J.C., E.L. Failey, S.J. Price, and M.E. Dorcas. 2007. An assessment of leech parasitism on semi-aquatic turtles in the Western Piedmont of North Carolina. Southeastern Naturalist 6(2): 191-202.

Madill, J., and P. Hovingh. 2007. Freshwater leech (Annelida: Hirudinida) distribution in the Canadian Province of Newfoundland and Labrador and adjacent regions: check-list, new records, new pigmentation forms, and Pleistocene refugia. Zootaxa 1657: 1-21.

Moser, W.E., R.W. Van Devender, and D.J. Klemm. 2005. Life history and distribution of the leech Oligobdella biannulata (Moore, 1900) (Euhirudinea: Glossiphoniidae). Comparative Parasitology 72(1): 17-21.

Moser, W.E., S.V. Fend, D.J. Richardson, C.A. Hammond, E.A. Lazo-Wasem, F.R. Govedich, and B.S. Gullo. 2013. A new species of Helobdella (Hirudinida: Glossiphoniidae) from Oregon, USA. Zootaxa 3718(3): 287-294.

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Moser, W.E., D.J. Klemm, D.J. Richardson, B.A. Wheeler, S.A. Trauth, and B.A. Daniels. 2006. Leeches (Annelida: Hirudinida) of northern Arkansas. Journal of the Arkansas Academy of Science 60: 84-95.

Moser, W.E., D.J. Richardson, C.I. Hammond, and E. Lazo-Wasem. 2011. Molecular characterization of Helobdella modesta (Verrill, 1872) (Hirudinida: Glossiphoniidae) from its type locality, West River and Whitneyville Lake, New Haven County, Connecticut, USA. Zootaxa 2834: 65-68.

Moser, W.E., D.J. Richardson, C.I. Hammond, and E. Lazo-Wasem. 2012. Molecular characterization of Glossiphonia elegans (Verrill, 1872) (Glossiphoniidae: Hirudinida) from its type locality, West River, New Haven County, Connecticut, USA. Zootaxa 3195: 57-60.

Moser, W.E., D.J. Richardson, B.A. Wheeler, K.J. Irwin, B.A. Daniels, S.E. Trauth, and D.J. Klemm. 2008. Placobdella cryptobranchii (Rhynchobdellida: Glossiphoniidae) on Cryptobranchus alleganiensis bishopi (Ozark Hellbender) in Arkansas and Missouri. Comparative Parasitology 75(1): 98-101.

Oceguera-Figueroa, A., S. Kvist, S.C. Watson, D.F. Sankar, A.M. Overstreet, and M.E. Siddall. 2010. Leech collections from Washington State, with the description of two new species of Placobdella (Annelida: Glossiphoniidae). American Museum Novitates 3701: 1-14.

Oosthuizen, J.H., and R.W. Davies. 1992. Redescription of the duck leech, Theromyzon rude (Baird, 1869) (Rhynchobdellida: Glossiphoniidae). Canadian Journal of Zoology 70: 2028-2033.

Oosthuizen, J.H., and R.W. Davies. 1993. A new species of Theromyzon (Rhynchobdellida: Glossiphoniidae), with a review of the genus in North America. Canadian Journal of Zoology 71: 1311-1318.

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Richardson, D.J., W.E. Moser, C.I. Hammond, A.C. Shevchenko, and E. Lazo-Wasem. 2010. New geographic distribution records and host specificity of Placobdella ali (Hirudinida: Glossiphoniidae). Comparative Parasitology 77(2): 202-206.

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Rutter, R.P., and D.J. Klemm. 2001. The presence of an Asian leech, Barbronia weberi, in a home aquarium in south Florida (Hirudinea: Salifidae). Florida Scientist 64(3): 216-218.

Sawyer, R.T. 1972. North American freshwater leeches, exclusive of the Piscicolidae with a key to all species. Illinois Biological Monographs 46(1): 1-154.

Sawyer, R.T. 1974. Leeches (Annelida: Hirudinea). Pp. 81-142, In: C.W. Hart, Jr., and S.L.H. Fuller (eds.). Pollution ecology of freshwater invertebrates. Academic Press, Inc., New York.

Sawyer, R.T. 1986a. Leech biology and behavior, Volume I. Anatomy, physiology, and behaviour. Clarendon Press, Oxford. ix + pp. 1-417 + index (33 pp.).

Sawyer, R.T. 1986b. Leech biology and behavior, Volume II. Feeding biology, ecology, and systematics. Clarendon Press, Oxford. xiv + pp. 418-793 + index (33 pp.).

Sawyer, R.T. 1986c. Leech biology and behavior, Volume III. Bibliography. Clarendon Press, Oxford. x + pp. 799-1,065.

Shankland, M., S.T. Bissen, and D.A. Weisblat. 1992. Description of the Californian leech Helobdella robusta sp. nov., and comparison with Helobdella triserialis on the basis of morphology, embryology, and experimental breeding. Canadian Journal of Zoology 70: 1258-1263.

Siddall, M.E. 2002. Phylogeny of the leech family Erpobdellidae (Hirudinida: Oligochaeta). Invertebrate Systematics 16: 1-6.

Siddall, M.E., K. Apakupakul, E.M. Burreson, K.A. Coates, C. Erséus, and S.R. Gelder. 2001. Validating Livanow: molecular data agree that leeches, branchiobdellidans, and Acanthobdella peledina forma monophyletic group of oligochaetes. Molecular Phylogenetics and Evolution 21: 346-351.

Siddall, M.E., and E. Borda. 2003. Phylogeny and revision of the leech genus Helobdella (Glossiphoniidae) based on mitochondrial gene sequences and morphological data and a special consideration of the triserialis complex. Zoologica Scripta 32(1): 23-33.

Siddall, M.E., and J. Bowerman. 2006. A new species of glosssiphoniid leech from Rana pretiosa (Amphibia: Ranidae) in Oregon. Journal of Parasitology 92(4): 855-857.

Siddall, M.E., R.B. Budinoff, and E. Borda. 2005. Phylogenetic evaluation of systematics and biogeography of the leech family Glossiphonidae. Invertebrate Systematics 19: 105-112.

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

Williams, J.I., and E.M. Burreson. 2005. Gonimosobdella klemmi n. gen. sp. (Hirudinida: Piscicolidae) from cyprinid fishes in Arkansas, Illinois, and Missouri, U.S.A. Comparative Parasitology 72 (2): 166-172.

Williams, J.I., and E.M. Burreson. 2006. Phylogeny of the fish leeches (Oligochaeta, Hirudinida, Piscicolidae) based on nuclear and mitochondrial genes and morphology. Zoologica Scripta 35(6): 627-639.

Wirchansky, B.A., and D.H. Shain. 2010. A new species of Haemopis (Annelida: Hirudinea): Evolution of North American terrestrial leeches. Molecular Phylogenetics and Evolution 54: 226-234.

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Suggested citation for this electronic web page: Klemm, D.J., W.E. Moser, and M.J. Wetzel. 2015. Classification and checklist of the leeches (Phylum Annelida: Class Clitellata: Subclass Hirudinida) occurring in North America north of Mexico. 24 January. World Wide Web URL: < >.

page update: 24 January 2015.
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