INTRODUCTION AND ANNOTATED CHECKLIST OF SPECIES
Compiled by Mark J. Wetzel1 and John W. Reynolds 2
1 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.
Tel.: (217) 244-2108; E-Mail: mjwetzel{AT}illinois.edu
2 Oligochaetology Laboratory, 18 Broadview Court, Kitchener, Ontario, Canada N2A 2X8. Tel.: (519) 896-4728; E-mail: john.reynolds1{AT}sympatico.ca
Recently, Reynolds and Wetzel (2011) published a paper summarizing the current status of earthworms in Illinois, including a review of published records, previously unpublished records in project reports, field notes of researchers, specimens held in the Annelida collections of the INHS and the USNM (National Museum of Natural History -- Smithsonian Institution, Washington, DC), and recent collections of earthworms by the authors, colleagues, and the general public. Specimens and their associated data from over 360 collections were reviewed, increasing from 52 to 79 the number of counties in Illinois from which earthworms have been reported. During that survey, specimens representing three species previously unreported from the state -- Aporrectodea tuberculata, Murchieona muldali, and Diplocardia gracilis -- were discovered, increasing the diversity of earthworms occurring in Illinois to 38 species representing 18 genera in six families. Twenty of these species are considered introduced, while 18 are considered native to North America. Four species in Illinois -- Pontodrilus bermudensis, Pontoscolex corethrurus, Amynthas diffringens, and Amynthas hawayensis -- have been collected only from greenhouses or other indoor cultures (Reynolds and Wetzel 2004, 2008, 2011).
Several primarily terrestrial oligochaetes (e.g., Eisenia foetida and Eiseniella tetraedra - both in the Family Lumbricidae) have occasionally been collected from aquatic habitats. Species in the Family Sparganophilidae are considered limicolous, or mud-loving - commonly collected from substrates and riparian margins of streams, ponds, marshes, bogs, wetlands, and occasionally from lakes and ponds. Sparganophilus eiseni is the most widely distributed and commonly collected species in this Nearctic family, and to date is the only sparganophilid known to occur in Illinois.
Historical works focusing on one or more aspects of earthworms in Illinois include those by Garman (1888), the numerous studies of Smith (1895a,b, 1900, 1915, 1917, 1922, 1924, 1925, 1928), and those of Smith and Gittens (1915), and Harman (1960). More recently, Zaborski and Gittenger (2001) published a paper on worm circling in Amynthas hupeiensis (family Megascolecidae).
Pertinent references for the study of terrestrial Oligochaeta include Dindal (1990), Fend (2000, 2006), Fender (1985, 1994), Fender and McKey-Fender (1990), James (1990, 1994), Reynolds (1977a,b; 1978, 1980, 1994, 1995a,b,c; 2001, 2008a,b,c, and 2011), Reynolds and Reynolds (1992), Reynolds et al. (1974), Edwards (1998, 2004), and Reynolds and Wetzel (2004, 2008, 2011, 2012 - which include extensive bibliographies on terrestrial oligochaetes in Illinois and throughout North America].
Other annelids in Illinois. Wetzel (1992) published the first comprehensive review of aquatic Annelida in Illinois, which documented the occurrence of 130 species of aquatic worms representing 71 genera in 15 families for the state. Additonal species have since been identified from specimens collected in Illinois; this information will be updated on other websites.
Many genera and species in the family Enchytraeidae (microdrile oligochaetes) are commonly collected from terrestrial habitats as well as from freshwater systems (streams, springs, seeps, ponds, lakes) as well as from moist and muddy (limicolous) habitats.
In addition to terrestrial megadrile and microdrile oligochaetes, one semi-terrestrial leech - Haemopis terrestris (Forbes, 1890) (Hirudinea, Haemopidae) - also occurs in Illinois (see Wetzel 1992, and Klemm, Moser, and Wetzel 2012).
The list presented below has been compiled from records that have been published in the historical or recent scientific literature. Of note here are the species from the Walter J. Harman Terrestrial Annelida Collection, graciously donated to the Illinois Natural History Survey (INHS) Annelida Collection in Champaign by Dr. Harman in the late 1980s. Dr. Harman's PhD dissertation (University of Illinois at Urbana-Champaign, 1960) focused on the terrestrial oligochaetes in central Illinois. A short obituary for Dr. Harman is presented HERE; a more extensive obituary, including a list of Dr. Harman's publications, was presented in Wetzel (2006: Hydrobiologia vol. 564; full citation below).
The list below also accounts for recent nomenclatural and systematic changes. Many recent works discussing the phylogeny of aquatic Annelida also have been included in the section on classification below [but this section needs updating]. References noted in the text of this document are included in the 'Literature Cited' section which follows the checklist of species.
Classification of and phylogenetic relationships within the Annelida. Several classifications for and discussions regarding the phylogenetic relationships within the Annelida have been proposed over the last 30+ years. The reader is directed to Jamieson (1978, 1980, 1988), Timm (1981), Brinkhurst (1982, 1986, 1988, 1989, 1991a, 1991b, 1991c, 1992, 1994), Kasprzak (1982, 1984), Kathman and Brinkhurst (1998), Parker (1982), Holt (1986, 1989), Erséus (1987, 1990), Sawyer (1986a, 1986b, 1986c), Brinkhurst and Nemec (1987), Coates (1987, 1989), Nemec and Brinkhurst (1987), Brinkhurst and Gelder (1989), Martin et al. (2000), Gelder and Brinkhurst (2001), Purschke et al. (1993), Erséus (2005), Envall et al. (2006), Rouse and Pleijel (2007), Rousset et al. (2007, Schmelz and Timm (2008), and Govedich et al. (2010) for further discussion on the phylogenetic relationships of the annelid groups. Note that several recently published papers focus on the phylogenetic relationships of annelids; these citations will be added here soon.
Nomenclature. Nomenclature for all taxa on this page follows Reynolds and Cook (1976, 1981, 1989, 1993), and the second edition of Nomenclatura Oligochaetologica - currently being compiled by John W. Reynolds and Mark J. Wetzel.
Species preceeded by '#' represent collection records from greenhouses or other indoor cultures only; they have yet to be found occurring in 'nature' in Illinois.
Species followed by an ' I ' are considered introduced in North America.
A preliminary list of terrestrial microdrile oligochaetes in the family Enchytraeidae follows the list of megadriles (presented immediately below); several colleagues are assisting mjw in updating this list.
SUBPHYLUM CLITELLATA
CLASS OLIGOCHAETA
Family Glossoscolecidae
Family Lumbricidae
Family Megascolecidae
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Order Enchytraeida
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Note: The above list of enchytraeids in Illinois is incomplete / currently being updated.
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LITERATURE CITED and PERTINENT REFERENCES [incomplete at this time]
Brinkhurst, R.O. 1982. Evolution in the Annelida. Can. J. Zool. 60(5): 1043-1059.
Brinkhurst, R.O. 1986. Guide to the freshwater aquatic microdrile oligochaetes of North America. Can. Spec. Publ. Fish. Aquat. Sci. 84. vi + 259 pp.
Brinkhurst, R.O. 1991a. Ancestors. Mitt. Hamburg Mus. Inst. 88(1): 97-110.
Brinkhurst, R.O. 1992. Evolutionary relationships within the Clitellata. Soil. Biol. Biochem. 24(12): 1202-1205.
Brinkhurst, R.O. 1994. Evolutionary relationships within the Clitellata: an update. Megadrilogica 5(10: 109-112.
Brinkhurst, R.O., and A.F.L. Nemec. 1987. A comparison of phenetic and phylogenetic methods applied to the systematics of Oligochaeta. Hydrobiologia 155: 65-74.
Brinkhurst, R.O., and S.R. Gelder. 2001. Annelida: Oligochaeta, including Branchiobdellidae. Pages 431-463, In: J.H. Thorp and A.P. Covich (eds). Ecology and classification of North American freshwater invertebrates. Second Edition. Academic Press, San Diego, CA [superseded by Govedich et al. 2010, citation below].
Coates, K.A. 1989. Phylogeny and origins of Enchytraeidae. Hydrobiologia 180: 17-33.
Dindal, D.L., editor. 1990. Soil biology guide. John Wiley & Sons, Inc., New York. xviii 1349 pages.
Erséus, C. 1987. Phylogenetic analysis of the aquatic Oligochaeta under the principle of parsimony. Hydrobiologia 155: 75-89.
Erséus, C. 2005. Phylogeny of oligochaetous Clitellata. Hydrobiologia 535: 357-372.
Edwards, C.A. Editor. 1998. Earthworm ecology. St. Lucie Press [CRC Press LLC], Boca Raton, FL. vi + 389.
Edwards, C.A. Editor. 2004. Earthworm ecology, Second Edition. CRC Press LLC, Boca Raton, FL. 441 pp.
Envall, I., M. Kállersjó, and C. Erséus. 2006. Molecular evidence for the non-monophyletic status of Naidinae (Annelida, Clitellata, Tubificidae). Molecular Phylogenetics and Evolution 40: 570-584.
Fend, S.V. 2010. Megadrile Oligochaeta. Pp. 149-166, In: Wetzel, M.J., S.V. Fend, K.A. Coates, R.D. Kathman, and S.R. Gelder. 2010. (full citation, below).
Fender, W. M. 1985. Earthworms of the western United States. I. Lumbricidae. Megadrilogica 4: 93-129.
Fender, W. M. 1994. Native earthworms of the Pacific Northwest: an ecological overview. in P. F. Hendrix, editor. Ecology and biogeography of earthworms in North America. Lewis Publishing, Boca Raton, Florida.
Fender, W. M., and McKey-Fender, D. 1990. Oligochaeta: Megascolecidae and other earthworms from western North America. Pages 357-378, in D. L. Dindal, editor. Soil biology guide. John Wiley & Sons, Inc., New York.
Garman, H. 1888. On the anatomy and histology of a new earthworm (Diplocardia communis, gen. et sp. nov.). Bulletin of the Illinois State Laboratory of Natural History 3: 47-77.
Gates, G.E. 1937. Notes on some species of Drawida and Pheretima with descriptions of three new species of Pheretima. Bull. Mus. Comp. Zool., Harvard 80: 305-335.
Gates, G.E. 1942. Check list and bibliography of North American earthworms. AmericanMidl.Nat. 27(1): 86-108.
Gates, G.E. 1943. On some American and Oriental earthworms. Ohio J. Sci. 43(2): 87-98.
Gates, G.E. 1954. Exotic earthworms in the United States. Bull. Mus. Comp. Zool., Harvard 111(6): 219-258.
Gates, G.E. 1956. Notes on American earthworms of the family Lumbricidae. III-IV. Bull. Mus. Comp. Zool., Harvard 115(1): 1-46.
Gates, G.E. 1960. Earthworms of North American caves. Natl. Speleol. Bull. 21(2): 77-84.
Gates, G.E. 1963. Miscellanea Megadrilogica. VII. Greenhouse earthworms. Proc. Biol. Soc. Washington 76: 9-18.
Gates, G.E. 1967. On the earthworm fauna of the Great American desert and adjacent areas. Great Basin Nat. 27(3): 142-176.
Gates, G.E. 1972a. Contributions to North American Earthworms (Annelida: Oligochaeta) No. 3. Toward a revision of the family Lumbricidae. IV. The trapezoides species group. Bull. Tall Timbers Res. Stn. No. 12: 1-146.
Gates, G.E. 1972b. Contributions to North American Earthworms (Annelida). On American earthworm genera. I. Eisenoides (Lumbricidae). Bull. Tall Timbers Res. Stn. No. 13: 1-17.
Gates, G.E. 1972c. Burmese earthworms. An introduction to the systematics of megadrile oligochaetes with special references to south-east Asia. Trans. American Philos. Soc. 62(7): 1-326.
Gates, G.E. 1973a. Contributions to North American Earthworms (Annelida) No. 6. Contributions to a revision of the earthworm family Glossoscolecidae. I. Pontoscolex corethrurus (Müller, 1857). Bull. Tall Timbers Res. Stn. No. 14: 1-12.
Gates, G.E. 1973b. Contributions to North American Earthworms (Annelida) No. 8. The earthworm genus Octolasion in America. Bull. Tall Timbers Res. Stn. No. 14: 29-50.
Gates, G.E. 1974a. Contribution to a revision of the Lumbricidae. X. Dendrobaena octaedra (Savigny) 1826, with special reference to the importance of its parthenogenetic polymorphism for the classification of earthworms. Bull. Tall Timbers Res. Stn. No. 15, pp. 15-57.
Gates, G.E. 1974b. Contributions to a revision of the family Lumbricidae. XI. Eisenia rosea (Savigny, 1826). Bull. Tall Timbers Res. Stn., No. 16: 9-30.
Gates, G.E. 1974c. On a new species of earthworm in a southern portion of the United States. Bull. Tall Timbers Res. Stn., No. 15: 1-13.
Gates, G.E. 1977a. More on the earthworm genus Diplocardia. Megadrilogica 3(1): 1-48.
Gates, G.E. 1977b. Contributions to a revision of the earthworm family Lumbricidae. XX. The genus Eiseniella in North America. Megadrilogica 3(5): 71-79.
Gates, G.E. 1978. The earthworm genus Lumbricus in North America. Megadrilogica 3(6): 81-116.
Gates, G.E. 1982. Farewell to North American Megadriles. Megadrilogica 4(1-2): 12-77.
Govedich, F.R., B.A. Bain, W.E. Moser, S.R. Gelder, R.W. Davies, and R.O. Brinkhurst. 2010. Annelida (Clitellata) Oligochaeta, Branchiobdellida, Hirudinida, and Acanthobdellida. Pages 385-436, In: J.H. Thorp and A.P. Covich (eds). Ecology and classification of North American freshwater invertebrates. Third Edition xiv + 1,021 pp. Academic Press / Elsevier, San Diego, CA. ISBN 978-0-12-374855-3. [www.elsevier.com] USD$ 139.95 / 93.95 Euro.
Harman, W.J. 1960. Studies on the taxonomy and musculature of the earthworms of central Illinois. University of Illinois at Urbana-Champaign. unpublished Ph.D. thesis. v + 107 pp. [UIUC Library Cat. No. 595.16 H227s]
Heimburger, H.V. 1915. Notes on Indiana earthworms. Proc. Indiana Acad. Sci. 25: 281-285.
Holt, P.C. 1989. Comments on the classification of the Clitellata. Hydrobiologia 180: 1-5.
James, S.W. 1990. Diplocardia kansensis, a new earthworm from Kansas, with redescriptions of Diplocardia riparia Smith and D. fuscula Gates (Annelida: Oligochaeta: Megascolecidae). Proceedings of the Biological Society of Washington 103:179-186.
James, S.W. 1990. Oligochaeta: Megascolecidae and other earthworms from southern and midwestern North America. Pages 379-386, in D. L. Dindal, editor. Soil biology guide. John Wiley & Sons, Inc., New York.
James, S. W. 1994. Systematics, biogeography and ecology of earthworms from eastern, central, southern and southwestern USA. Pages 29-51, in P. F. Hendrix, editor. Ecology and biogeography of earthworms in North America. Lewis Publishing, Boca Raton, Florida.
Jamieson, B.G.M. 1978. Phylogenetic and phenetic systematics of the opisthoporous Oligochaeta (Annelida: Clitellata). Evol. Theor. 3(4): 195-233.
Jamieson, B.G.M. 1980. Preliminary discussion of an Hennigian analysis of the phylogeny and systematics of opisthoporous oligochaetes. Rev. Ecol. Biol. Sol 17(2): 261-275.
Jamieson, B.G.M. 1988. On the phylogeny and higher classification of the Oligochaeta. Cladistics 4: 367-410.
Joyner, J.W. 1960. Earthworms of the Upper Whitewater Valley (East-Central) Indiana. Proc. IndianaAcad. Sci. 69: 313-319.
Kasprzak, K. 1982. Problems of the origin of oligochaetes (Annelida: Oligochaeta). Prezgl. Zool. 26: 145-160. (English translation, Can. Transl. Fish. Aquat. Sci. No. 4996).
Kasprzak, K. 1984. The previous and contemporary conceptions on phylogeny and systematic classifications of Oligochaeta (Annelida). Ann. Zool. (Polska Akad. Nauk Inst. Zool.) 38: 205-223.
Klemm, D.J., W.E. Moser, and M.J. Wetzel. 2012. Classification and checklist of the leeches (Phylum Annelida: Class Clitellata: Subclass Hirudinida) occurring in North America north of Mexico. 4 January. World Wide Web URL: < http://www.inhs.uiuc.edu/~mjwetzel/FWLeechesNA.html >.
Martin, P., I. Kaygorodova, D. Yu. Sherbakov, and E. Verheyen. 2000. Rapidly evolving lineages impede the resolution of phylogenetic relationships among Clitellata (Annelida). Mol. Phyl. Evol. 15: 355-368.
Minnich, J. 1977. The earthworm book. Rodale Press, Emmaus, PA. xii + 372 pp.
Murchie, W.R. 1961. A new diplocardian earthworm from Illinois (Oligochaeta: Megascolecidae). Ohio J. Sci. 61(6): 367-371.
Olson, H.W. 1936. Earthworms of Missouri. Ohio J. Sci. 36(2): 102-113.
Parker, S.P. 1982. Synopsis and classification of living organisms. McGraw Hill, New York. 1,232 pp.
Peck, S.B., and J.L. Lewis. 1978. Zoogeography and evolution of the subterranean invertebrate faunas of Illinois and southeastern Missouri. National Speleological Society Bulletin 40(2): 39-63.
Reynolds, J.W. 1977a. The earthworms of Tennessee (Oligochaeta). III. Komarekionidae, with notes on distribution and biology. Megadrilogica 3: 65-69.
Reynolds, J.W. 1977b. The earthworms (Lumbricidae and Sparganophilidae) of Ontario. Life Sciences Miscellaneous Publications, Royal Ontario Museum, Toronto, Ontario, Canada. ix + 141 pp.
Reynolds, J.W. 1978. The earthworms of Tennessee (Oligochaeta). IV. Megascolecidae, with notes on distribution, biology and a key to the species in the state. Megadrilogica 3: 117-129.
Reynolds, J.W. 1980. The earthworm family Sparganophilidae (Annelida, Oligochaeta) in North America. Megadrilogica 3: 189-204.
Reynolds, J. W. 1994. The status of exotic earthworm systematics and biogeography in North America. Pages 1-27, in P. F. Hendrix, editor. Ecology and biogeography of earthworms in North America. Lewis Publishing, Boca Raton, Florida.
Reynolds, J.W. 1994. The distribution of the earthworms (Oligochaeta) of Indiana: a case for the Post Quaternary Introduction Theory of megadrile migration in North America. Megadrilogica 5(3): 13-32.
Reynolds, J.W. 1995a. The distribution of earthworms (Annelida, Oligochaeta) in North America. Pages 133-153, in P.C. Mishra, N. Behera, B.K. Senapati, and B.C. Guru, (editors). Advances in Ecology and Environmental Sciences. Ashish Publishing House, New Delhi. 651 pp.
Reynolds, J.W. 1995b. Earthworms of the world. Global Biodiversity 4(1): 11-16.
Reynolds, J.W. 1995c. The status of exotic earthworm systematics and biogeography in North America. Pp. 1-17, In P.F. Hendrix, Editor. Ecology and biogeography fo earthworms in North America. Lewis Publ., Boca Raton, FL. 244 pp.
Reynolds, J.W. 2001. Sparganophilidae -- are terrestrial oligochaetologists missing the habitat in North America? Megadrilogica 8(11): 82-84.
Reynolds, J.W. 2008a. The earthworms (Oligochaeta: Acanthodrilidae, Komarekionidae, Lumbricidae, Megascolecidae, and Sparganophilidae) of Kentucky, USA. Megadrilogica 12(5): 73-88.
Reynolds, J.W. 2008b. The earthworms (Oligochaeta: Acanthodrilidae, Lumbricidae, Megascolecidae, and Sparganophilidae) of Missouri,USA, revisited. Megadrilogica 12(6): 89-104.
Reynolds, J.W. 2008c. Sparganophilidae (Annelida, Oligochaeta) distribution in North America and elsewhere, revisited. Megadrilogica 12(9): 125-143.
Reynolds, J.W. 2011. The earthworms (Oligochaeta: Acanthodrilidae, Glossoscolecidae, Komarekionidae, Lumbricidae, Megascolecidae and Sparganophilidae) of the Midwestern United States. Megadrilogica 15(5): 69-139.
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Reynolds, J.W., and D.G. Cook. 1981. Nomenclatura oligochaetologica. Supplementum primum. A catalogue of names, descriptions and type specimens of the Oligochaeta. [published by the University of New Brunswick] Fredericton, New Brunswick. v + 39 pp. [out of print].
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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 Mus. Monogr. Ser. (Nat. Sci.) No. 9. vi + 33 pp. [this third supplement is available, at no charge, from the New Brunswick Museum, 277 Douglas Avenue, Saint John, NB, Canada E2K 1E5.].
Reynolds, J.W., and K.W. Reynolds. 1992. Les vers de terre (Oligochaeta: Lumbricidae et Sparganophilidae) sur la rive nord du Saint-Laurent (Québec). Megadrilogica 4: 145-161.
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Reynolds, J.W., and M.J. Wetzel. 2011. The earthworms (Oligochaeta: Acanthodrilidae, Glossoscolecidae, Komarekionidae, Lumbricidae, Megascolecidae and Sparganophilidae) of Illinois, USA. Megadrilogica 15(5): 35-67.
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. Nomenclatura oligochaetologica. Second Edition. A catalogue of names, descriptions and type specimens of the Oligochaeta. In prep.
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Verdonschot, P.F.M., H. Wang, A. Pinder, and R. Nijboer (eds). 2006. Aquatic Oligochaete Biology IX. Selected papers from the 9th Symposium on Aquatic Oligochaeta, 6-10 October 2003, Wageningen, The Netherlands. Developments in Hydrobiology 186. Published by Springer, Dordrecht, The Netherlands. ix + 222 pp. ISBN 1-4020-4781-9.
Wetzel, M.J. 1992. Aquatic Annelida of Illinois: Introduction and checklist of species. Trans. Illinois St. Acad. Sci. 85(1 and 2): 87-101.
Wetzel, M.J. 2006. In memoriam: Professor Walter James Harman, PhD (1938-2002). Hydrobiologia 564: 1-4. [also, same citation, pages, in Verdonschot et al. 2006. cited above].
Wetzel, M.J., S.V. Fend, K.A. Coates, R.D. Kathman, and S.R. Gelder. 2010. Taxonomy, systematics, and ecology of aquatic Oligochaeta and Branchiobdellidae (Annelida, Clitellata) of North America. A workbook. 1 November. vi + 280 pp. + 4 color plates.
Wetzel, M.J., R.D. Kathman, S.V. Fend, and K.A. Coates. 2012. Classification and checklist of the freshwater oligochaetes occurring in North America north of Mexico. 4 January.
World Wide Web URL: http://www.inhs.uiuc.edu/~mjwetzel/FWOligoNAChklst.html.
Zaborski, E.R., and L.A.S. Gittenger. 2001. Amynthas hupeiensis (Michaelsen, 1895) (Oligochaeta: Megascolecidae) in Illinois, USA, with observations on worm circling. Megadrilogica 8(4): 13-16.