² Department of Invertebrate Zoology MRC 183, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0163 USA. Phone: (202) 633-0669; FAX: (202) 357-3043; E-Mail: moserw{AT}si.edu

³ Illinois Natural History Survey, University of Illinois Institute of Natural Resource Sustainability, Forbes Natural History Building, 1816 S. Oak Street, Champaign, Illinois 61820 USA. Phone: (217) 244-2108; FAX: (217) 333-4949; E-Mail: mjwetzel{AT}illinois.edu

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 have served on this Committee since 1989, and (in collaboration with many colleagues, including Don Klemm and Bill Moser) have been preparing the Annelida volume for this series.

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, 77 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 (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; 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. Additional citations for publications focusing on leech biology, ecology, distribution, taxonomy, and systematics will be added to this website soon.

Genus Acanthobdella Grube, 1851
Acanthobdella peledina Grube, 1851

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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 septagon Sawyer & Shelley, 1976

Haemopis terrestris (Forbes, 1890)

Genus Hirudo Linnaeus, 1758

Hirudo medicinalis Linnaeus, 1758 ^{- I -}

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

Genus Erpobdella Blainville, 1918

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

Genus Barbronia Johansson, 1918
Barbronia weberi Blanchard, 1897 ^{- I -} (See Rutter and Klemm, 2001)

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Genus Actinobdella Moore, 1901
Actinobdella annectens Moore, 1906
Actinobdella inequiannulata Moore, 1901
Actinobdella [= Placobdella] pediculata (Hemingway, 1908)

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 Blanchard, 1896
Helobdella californica Kutschera, 1988
Helobdella [= Gloiobdella] elongata (Castle, 1900)
Helobdella fusca (Castle, 1900)
Helobdella lineata (Verrill, 1874)
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 Blanchard, 1893
Placobdella ali Hughes & Siddall, 2007
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 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 [= Desserobdella] phalera (Graf, 1899)
Placobdella [= Desserobdella] picta (Verrill, 1872)
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

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 Blainville, 1818
Piscicola geometra (Linnaeus, 1761)
Piscicola milneri (Verrill, 1871)
Piscicola punctata (Verrill, 1871)

Footnotes

^{- 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 H. papillata (three irregular rows of papillae irrespective of size and pigmentation) from H. 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.

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.

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. Packarsky, 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.

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. Macroinvertebrae 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.

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., 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, 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.

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.

Readel, A.M., C.A. Phillips, and M.J. Wetzel. 2008. Leech parasitism in a turtle assemblage: effects of host and environmental characteristics. Copeia 2008(1): 227-233.

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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.

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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.

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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.

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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. 1982. Kansas leeches (Annelida: Hirudinea) with notes on distribution and ecology. II. Technical Publications of the State Biological Survey of Kansas 12: 105-111.

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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.

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Klemm, D.J., W.E. Moser, and M.J. Wetzel. 2009. Classification and checklist of the leeches (Phylum Annelida: Class Clitellata: Subclass Hirudinida) occurring in North America north of Mexico. World Wide Web URL:
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