THE EARTHWORMS (ANNELIDA, CLITELLATA, OLIGOCHAETA) OF ILLINOIS:

INTRODUCTION AND ANNOTATED CHECKLIST OF SPECIES

–  Families Acanthodrilidae, Glossoscolecidae, Komarekionidae, Lumbricidae, Megascolecidae, and Sparganophilidae –

Compiled by Mark J. Wetzel 1 and John W. Reynolds 2
[ page update: 2019-VII-01 ]


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.  E-Mail: mjwetzel{AT}illinois.edu

2  Oligochaetology Laboratory, 9-1250 Weber Street East, Kitchener, Ontario N2A 4E1 CANADA note: new postal address as of November 2016].  
         Tel.: (519) 896-4728; E-mail: john.reynolds1{AT}sympatico.ca


Introduction. Reynolds and Wetzel (2004) reported 161 species of earthworms representing 37 genera in 10 families known to occur in North America north of Mexico; of these, 45 are considered introduced (Reynolds and Wetzel 2004). An update of their paper (Reynolds and Wetzel, 2008) expanded their scope of North American earthworm distributions to include Mexico, Puerto Rico, Hawaii, and Bermuda, therein documenting the occurrence of 253 species representing 59 genera in 10 families, and increasing the number of introductions to 67. 

In their second update, Reynolds and Wetzel (2012) reported 46 new state or provincial records for 27 native species, and 46 new state or provincial records for 29 introduced species.  New North American records for two native species (both known only from Mexico and described as new to science in 2009), one introduced species, and new records of species from habitats in the continental U.S. were also highlighted.  To summarize, 256 species of earthworms representing 59 genera in 10 families are recognized; of these 188 species are considered native to North America, while 68 are considered to have been introduced.  [A third update is now in preparation,  to be published in late 2019]. 

There are at least 14 introduced species in North America that are known only from greenhouses and other indoor cultures in one or more states from which they have been reported; however, it is important to emphasize that some of these species have also been reported in 'nature', particularly from habitats located in some of the more southern/warmer areas of the continent.

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.  [An update to our first paper on earthworms of Illinois is now in preparation, to be published in late 2019]. 

During our review of previously unidentified specimens for inclusion in our 2011 paper, 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.  

Collections of megadriles in Illinois by Wetzel and Reynolds in 2015 and 2016 resulted in the first report of two additional species for the state:  Bimastos welchi from Nachusa Grasslands in Lee County (April 2015) – a native species, and Microscolex dubius from Pulaski County (June 2016) – a European introduction.  

To date, 21 of the 40 species in Illinois are considered introductions, while 19 are considered native to North America. Four of the 21 introduced species in Illinois -- Pontodrilus bermudensis, Pontoscolex corethrurus, Amynthas diffringens, and Amynthas hawayensis -- have thus far been collected only from greenhouses or other indoor cultures (Reynolds and Wetzel 2004, 2008, 2011). Amynthas agrestis, another introduction to North America, likely occurs in Illinois, but specimens (preferably mature individuals) attributed to this genus need to be identified by qualified megadrile taxonomists prior to listing as occurring in any state or region.  

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, and are commonly collected from substrates and riparian margins of streams, ponds, marshes, bogs, wetlands, and occasionally from lakes and ponds. Sparganophilus tamesis (Sparganophilus eiseni) [see Rota et al. 2016] 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. Additional 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) (Hirudinida, Haemopidae) - also occurs in Illinois (see Wetzel 1992, and Wetzel, Govedich, Moser, and Klemm, 2019).

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 accounts presented in the on-line second edition of the nomenclator, Nomenclatura Oligochaetologica - Editio Secunda (Reynolds and Wetzel, 2019). 



 Checklist of terrestrial megadrile Oligochaeta occurring in Illinois.

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.


P H Y L U M - A N N E L I D A
SUBPHYLUM CLITELLATA
CLASS OLIGOCHAETA
-------------------------------------------------------------------------------------------------------
Subclass Metagynophora  
Order Opisthopora
Suborder Lumbricina
Superfamily Glossoscolecoidea
 
 
Family Glossoscolecidae Michaelsen, 1900
 
Genus Pontoscolex Schmarda 1861
# Pontoscolex corethrurus (Müller, 1856) - I -
 
Superfamily Lumbricoidea

Family Komarekionidae 
 
Genus Komarekiona Gates, 1974
Komarekiona eatoni Gates, 1974
 
Family Lumbricidae Rafinesque–Schmaltz 1815
 
Genus Allolobophora Eisen, 1873
Allolobophora chlorotica (Savigny, 1826) I -
 
Genus Aporrectodea Örley, 1885
Aporrectodea rosea (Savigny, 1826) - I -
Aporrectodea trapezoides (Dugès, 1828) - I -
Aporrectodea tuberculata (Eisen, 1874) I -
Aporrectodea turgida (Eisen, 1873) I -
 
Genus Bimastos H. F. Moore, 1893 
Bimastos beddardi (Michaelsen, 1891) 
Bimastos gieseleri (Ude, 1895) 
Bimastos heimburgeri (Smith, 1928) - type locality in Illinois [Piatt County]
Bimastos longicinctus (Smith & Gittens, 1915) - type locality in Illinois [Champaign County]
Bimastos parvus (Eisen, 1874) 
Bimastos tumidus (Eisen, 1874)
Bimastos welchi Smith, 1917 [ new Illinois record [ Lee Co., April 2015 ]
Bimastos zeteki (Smith & Gittens, 1915) - type locality in Illinois [Champaign County]
 
Genus Dendrobaena Eisen, 1873
Dendrobaena octaedra (Savigny, 1826) I -
 
Genus Dendrodrilus Omodeo, 1956
Dendrodrilus rubidus (Savigny, 1826) I -
 
Genus Eisenia Malm, 1877
Eisenia foetida Savigny, 1826 I -
Eisenia hortensis (Michaelsen, 1890) I -
 
Genus Eiseniella Michaelsen, 1900
Eiseniella tetraedra (Savigny, 1826 I -
 
Genus Eisenoides Gates, 1969 
Eisenoides carolinensis (Michaelsen, 1910)
 
Genus Lumbricus Linnaeus, 1758
Lumbricus rubellus Hoffmiester, 1843 I
Lumbricus terrestris Linnaeus, 1758 I -
 
Genus Murchieona Gates, 1978
Murchieonia muldali (Omodeo, 1956) I -
 
Genus Octolasion Örley, 1885
Octolasion tyrtaeum (Savigny, 1826) I -
 
Superfamily Megascolecoidea

Family Acanthodrilidae
 
Genus Diplocardia Garman, 1888
Diplocardia communis Garman, 1888 - type locality in Illinois [more specific locality information not included with published description]
Diplocardia conoyeri Murchie, 1961 - type locality in Illinois [St. Clair County]. Additional information presented in Stebbings (1962a, 1962b; citations below)]
Diplocardia gracilis Gates, 1943
Diplocardia ornata Gates, 1943
Diplocardia riparia Smith, 1895 - type locality in Illinois [Mason County]
[= Diplocardia prosenteris Macnab & McKey-Fender, 1955]
Diplocardia singularis (Ude, 1893) - type locality in Illinois [Vermilion County]
Diplocardia smithi Macnab & McKey-Fender, 1955
Diplocardia verrucosa Ude, 1895
 
Genus Microscolex Rosa, 1887
Microscolex dubius [ new Illinois record [ Pulaski Co., June 2016 ] I -
Microscolex phosphoreus (Dugès, 1837) I -
[= Microscolex hempeli Smith, 1896]
 
Genus Pontodrilus Perrier, 1874 
# Pontodrilus bermudensis (Beddard, 1891) I -
[= Perichaeta bermudensis Beddard, 1892]
 
Family Megascolecidae Rosa, 1891
 
Genus Amynthas Kinberg, 1867
# Amynthas diffringens (Baird, 1869) I -
# Amynthas hawayanus (Rosa, 1891) I -
Amynthas hupeiensis (Michaelsen, 1895) I -
 
Superfamily Sparganophiloidea

Family Sparganophilidae
 
Genus Sparganophilus Benham, 1892
Sparganophilus tamesis Benham, 1892
[= Sparganophilus eiseni Smith, 1895] - type locality in Illinois [Mason County]; see Rota et al. (2016).



Terrestrial microdrile oligochaetes
Order Enchytraeida

Family Enchytraeidae
 
Genus Fridericia Michaelsen, 1889
Fridericia agilis Smith, 1895
Fridericia firma Smith & Welch, 1913
Fridericia oconeensis Welch, 1914
Fridericia ratzeli (Eisen, 1904)
[ = Fridericia sima Welch, 1914, a junior synonym; see Schmelz, 2003]
Fridericia tenera Smith & Welch, 1913
 
Genus Henlea Michaelsen, 1889
Henlea ochracea (Eisen, 1878)
Henlea urbanensis Welch, 1914
Henlea welchi Bell, 1942
 
Genus Marionina Pfeffer, 1890
Marionina dirksi Bell, 1942
 
Genus Mesenchytraeus Eisen, 1878
Mesenchytraeus americanus Bell, 1942
 
--------------------------------------------------------------------------------------------------------------------------------
Note: The above list of enchytraeids in Illinois is incomplete / currently being updated.

 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.

M.J. Wetzel, M.J., F.R. Govedich, W.E. Moser, and D.J. Klemm. 2019. Classification and checklist of the leeches (Phylum Annelida: Class Clitellata: Subclass Hirudinida) occurring in North America north of Mexico.  World Wide Web URL: < https://www.inhs.illinois.edu/people/mjwetzel/fwleechesna >.

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.

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, cited below, are out of print; the second and third supplements, also 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. 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].

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 Mus. Monogr. Ser. (Nat. Sci.) No. 8. v + 37 pp. [this second supplement is 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 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.

Reynolds, J.W., E.E.C. Clebsch, and W.M. Reynolds. 1974. The earthworms of Tennessee (Oligochaeta). I. Lumbricidae. Continental North American Earthworms (Oligochaeta), no. 13. Bulletin of the Tall Timbers Research Station 17: 1-133.

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): 167-208.

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. 2014.  A checklist by counties of earthworms (Oligochaeta: Acanthodrilidae, Lumbricidae, Megascolecidae, and Sparganophilidae) in Michigan, USA.  Megadrilogica 17(5): 51-72. 

Reynolds, J.W. and M.J. Wetzel. 2018. Nomenclatura Oligochaetologica – A catalogue of names, descriptions and type specimens. Editio Secunda. URL: https://www.inhs.illinois.edu/people/mjwetzel/nomenoligo [accessed 01 June  2018]

Rota, E., S. Martinsson, M. Bartoli, A. Beylich, U. Graefe, A. Laini, M.J. Wetzel, and C. Erséus. 2016. Mitochondrial evidence supports a Nearctic origin for the spreading limicolous earthworm Sparganophilus tamesis Benham, 1892 (Clitellata, Sparganophilidae). Contributions to Zoology 85(1): 113-119. 

Rouse, G.W. and F. Pleijel. 2007. Annelida. Zootaxa 1668: 245-264.

Rousset, V., F. Pleijel, G.W. Rouse, C. Erséus, and M.E. Siddall. 2007. A molecular phylogeny of annelids. Cladistics 2007, 23:41-63.

Schmelz, R.M. 2003. Taxonomy of Fridericia (Oligochaetaa, Enchytraeidae). Revision of species with morphological and biochemical methods. Abhandlungen des Naturwissenschaftlichen Vereins in Hamburg (NF) 38. Goecke & Evers / Keltern-Weiler, D-75210 Keltern-Weiler, Germany. 415 pp. + 73 figs.

Schmelz, R.M., and T. Timm. 2008 [2007]. Advocating paraphyletic taxa in systematics of Clitellata. Pp. 99-108, In: Wang, H.-Z., M.J. Wetzel, A.M. Pinder, P.F.M. Verdonschot, and N. Arslan (eds.). Aquatic Oligochaete Biology X. Proceedings of the 10th International Symposium on Aquatic Oligochaeta, held at the Institute of Hydrobiology, Chinese Academy of Sciences in Wuhan, China, 16-21 October, 2006. Acta Hydrobiologica Sinica 31 (supplement). viii + 171 pp. ISSN 1000-3207.

Sims, R.W., and E.G. Easton. 1972. A numerical revision of the earthworm genus Pheretima auct. (Megascolecidae: Oligochaeta) with the recognition of new genera and an appendix on the earthworms collected by the Royal Society North Borneo Expedition. Biological Journal of the Linnaean Society 4(3): 169-268, with 4 figures.

Smith, F. 1895a. A preliminary account of two new Oligochaeta from Illinois. Bull. Illinois St. Lab. Nat. Hist. 4(5): 142-147.

Smith, F. 1895b. VIII. Notes on species of North American Oligochaeta. Bull. Illinois St. Lab Nat. Hist. 4(8): 285-297.

Smith, F. 1900. Notes on species of North American Oligochaeta. III. List of species found in Illinois, and descriptions of Illinois Tubificidae. Bull. Illinois St. Lab. Nat. Hist. 5(10): 441-458 + plates 39, 40.

Smith, F. 1915. Two new varieties of earthworms with a key to described species in Illinois. Bull. Illinois St. Lab. Nat. Hist. 10(8): 551-559 + 1 plate.

Smith, F. 1917. North American earthworms of the family Lumbricidae in the collections of the United States National Museum. Proc. United States Nat. Mus. 52(2174): 157-182.

Smith, F. 1922. Abnormal earthworm specimens, Helodrilus subrubicundus and H. tenuis. Trans. American Micros. Soc. 41(3): 153-154.

Smith, F. 1924. The calciferous glands of Lumbricidae and Diplocardia. Illinois Biol. Monogr. 9(1): 7-76.

Smith, F. 1925. Abnormal earthworms of the species Helodrilus subrubicundus (Eisen). Trans. American Micros. Soc. 44(2): 90-96.

Smith, F. 1928. An account of changes in the earthworm fauna of Illinois and a description of one new species. Illinois Nat. Hist. Surv. Bull. 17(10): 347- 362.

Smith, F. and E.M. Gittins. 1915. Two new species of Lumbricidae from Illinois. Bull. Illinois St. Lab. Nat. Hist. 10(7): 545-550.

Stebbings, J.H. 1962b. Endemic-Exotic Earthworm Competition in the American Midwest. Nature 196(4857): 905-906. December 1.

Stebbings, J.H. 1962a. Additions to the Illinois and Missouri Earthworm Fauna. The American Midland Naturalist 67(2: 504-505. April.

Struck, T.H., N. Schult, T. Kusen, E. Hickman, C. Bleidorn, D. McHugh, and K.M. Halanych. 2007. Annelid phylogeny and the status of Sipuncula and Echiura. BMC Evolutionary Biology 7: 57 (11 pp.).

Timm, T. 1981. On the origin and evolution of aquatic Oligochaeta. Eesti NSV Tead. Akad. Toim. Biologia 30(3): 174-181.

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. 2018. Classification and checklist of the freshwater oligochaetes occurring in North America north of Mexico. World Wide Web URL: https://www.inhs.illinois.edu/people/mjwetzel/fwoligonachecklist. 01 June.

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.



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Suggested citation for this electronic web page: Wetzel, M.J., and J.W. Reynolds. 2019. The earthworms (Annelida, Clitellata, Oligochaeta) of Illinois - Introduction and annotated checklist of species - Families Acanthodrilidae, Glossoscolecidae, Komarekionidae, Lumbricidae, Megascolecidae, and Sparganophilidae. World Wide Web URL: [https://www.inhs.illinois.edu/people/mjwetzel/illinoisearthworms]. 01 July 2019.


 

Copyright 1995-2011; 2012-2019, by Mark J. Wetzel {Research Scientist, and Curator / Collections Manager of the INHS Annelida Collection --
Illinois Natural History Survey (INHS), Prairie Research Institute at the University of Illinois at Urbana-Champaign, Champaign, Illinois USA}
unless otherwise noted; All Rights Reserved.This website contains original, copyrighted material; it is being provided here as a professional courtesy, exclusively for your private, non-commercial use. Reference to or redistribution of any part of the information contained herein - whether it be through oral, printed, electronic, or other tangible medium of expression - shall acknowledge the Illinois Natural History Survey (INHS), The Oligochaetology Lab, and any other agency or organization noted herein for their support of this research, and shall cite this website as the source of information. Unless otherwise noted, all photographs are © M.J. Wetzel (INHS). 

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