Printed in Canada ISSN 0380-9633

MEGADRILOGICAVolume 15, Number 11, November, 2012

THE STATUS OF TERRESTRIAL EARTHWORM (OLIGOCHAETA) SURVEYS INNORTH AMERICA AND SOME CARIBBEAN COUNTRIES.

John Warren Reynolds

Oligochaetology Laboratory, 18 Broadview Court, Kitchener, ON Canada N2A 2X8 andResearch Associate, New Brunswick Museum, Saint John, NB Canada E2K 1E5

(e-mail: john.reynolds1@sympatico. ca)

ABSTRACT

This paper traces the history of earthworm surveys in North America (Canada, Mexico and the UnitedStates) and some Caribbean countries (Belize, Bermuda, Cayman Islands, Dominica, Dominican Republic) beginningwith Altman, Olson and Causey up to the present time. There has been no earthworm survey in 11 of the United Statesand one state in Mexico. The three Canadian territories (NU, NW, YK)1 and two prairie provinces (MB, SK)2 havebeen sampled minimally. A recent paper currently recognized 256 species representing 59 genera in 10 families; ofthese, 188 species are considered native to North America, while 68 have been introduced.

Key words: North America, Annelida, Oligochaeta, earthworms, geographical distribution, surveys.

RÉSUMÉ

Cet article retrace l'histoire des inventaires de vers de terre en Amérique du nord (Canada, Mexique et États-Unis) et certains pays des Caraïbes (Belize, Bermudes, îles Caïmans, la Dominique et République dominicaine)commençant par Altman, Olson et Causey jusqu'à présent. Il y a eu aucun inventaire de vers de terre dans 11 des étatsaméricains et un état du Mexique. Les trois territoires canadiens (NU, NW, YK)1 et deux provinces des Prairies (MB,SK)2 ont seulement été échantillonnés de façon minimale. Une récente publication reconnait actuellement 256 espècesreprésentant 59 genres de 10 familles en Amérique du nord; 188 de ces espèces sont considérées comme indigènestandis que 68 ont été introduites.

Mots clé: Amérique du nord, annélides, Oligochaeta, vers de terre, répartition géographique, inventaires.

RESUMEN

En este trabajo se describe el conocimiento de lombrices de tierra en América del Norte (Canadá, Méxicoy Estados Unidos) y algunos paííses del Caribe (Belice, Bermuda, Islas Caimán, Dominica, República Dominicana)a partir de Altman, Olson y Causey hasta la actualidad. No ha habido estudio de lombrices en 12 de los EstadosUnidos y en un estado de México. En tres territorios canadienses (NU, NW, YK)1 y en dos provincias de praderas(MB, SK)2 se tomaron mínimamente muestras. Un estudio reconoció en la actualidad 256 especies que representan59 géneros incluídos en 10 familias; 188 especies son consideradas nativas de América del Norte, mientras que 68son introducidas.

Palabras clave: América del Norte, Annelida, Oligochaeta, lombrices de tierra, distribución geográfica,reconocimiento.

SAETAK

Ovaj rad prati istoriju istraivanja kišnih glista u Severnoj Americi (Kanada, Meksiko i SAD) i nekihkaripskih zemalja (Belize, Bermudi, Kajmanska Ostrva, Dominik, Dominicanska Republika) pocevši od Altmana,Olsona i Kouzija pa sve do dananašnjih dana. U 11 drava SAD-a i jednoj meksikoj dravi kišne gliste nisuistraivane. Tri kanadske oblasti (NU, NW, YK)1 i dve prerijske pokrajine (MB, SK)2 su bile minimalno uzorkovane.Sadašnjirad priznaje 256 vrsta koje predstavlju 59 rodova i 10 familija. Od njih, 188 vrsta se smatraju kao domaevrste Severne Amerike, dok je 68 introdukovano.

Key rei: Severna Amerika, Annelida, Oligochaeta, kišne gliste, geografska distribucija, istraivanja.

1 NU = Nunavut, NW = Northwest, YK = Yukon; 2 MB = Manitoba, SK = Saskatchewan.

The Status of Terrestrial Earthworm Surveys that have been conducted in North America, etc.228

ABSTRACTDocumentul acesta desemneazistoria cercetrilor viermilor de sol din America de Nord (Canada, Mexico

si Statele Unite ale Americii) si din câtevaâri Caribice (Belize, Bermuda, Insulele Cayman, Dominica, RepublicaDominican), ncepând cu Altman, Olsonşi Causey pânin timpul prezent. Nu s-a condus o cercetare a viermilorde sol n unsprezece state ale Statelor Unite ale Americiişi un stat din Mexico. Trei teritorii (NU, NW, YK)1şidouprovincii de prerie (MB, SK)2 din Canada au fost exemplificate minimal. Un document recent recunoaste 256de specii care reprezintâ 59 de clasificaii din 10 familii; din ele, 188 de specii sânt considerate native ale Americiide Nord, n timp ce 68 au fost introduse.Cuvinte principale: America de Nord, Annelida, Oligochaeta, viermi de sol, distibuie geografic, cercetri.

INTRODUCTIONThis paper presents a synoptic review of

historical and recent earthworm surveys that have beenconducted in North America and several Caribbeancountries. Distributional information for earthwormsin these areas, included in previous papers (e.g.,Reynolds 1995, 1998, 2004b), has been updated.

When I began earthworm studies in 1967, fewdistributional surveys for megadriles in North Americahad been published, although a significant number ofisolated records had been presented in several papers,most notably those of Garman (1888), Smith (1895a,1895b, 1896, 1897, 1900, 1912, 1915, 1917, 1928) andGates (1935). Three more substantive papers by Olson(1928, 1936, and 1940) summarized the results of hisearthworm surveys in Ohio, Missouri, and New York,respectively. Altman (1936) published a paper on theoligochaetes of the state of Washington, includingterrestrial as well as aquatic species. Two regionalearthworm surveys were published in 1942: Check listand bibliography of North American earthworms(Gates, 1942 – in which he summarized data from over100 papers published during the previous 121 years),and Earthworms from the North Eastern United States(Eaton, 1942 – in which he reported incidentalcollections from Maine to Pennsylvania). Causey(1952, 1953) published the results of his surveys ofearthworms in Arkansas.

Several doctoral dissertations included resultsof earthworm surveys in Michigan (Murchie, 1956),Illinois (Harman, 1960) and Louisiana (Tandy, 1969),although Tandy only included the genus Pheretima asit was known then.

In 1970, I changed direction from purelyecological studies on earthworms to a concentration intaxonomy and biogeography. I began in Tennessee,with the first of five parts on the earthworms of thatstate published in Reynolds et al. (1974). Whileworking with Dr. Gordon Gates, and with my doctoraladvisor Dr. Ed Clebsch and my wife, Wilma during theearly 1970s on the earthworms of Tennessee, I learnedfrom Gates that there were no published or unpublished

earthworm records for either Delaware or RhodeIsland. Information associated with previouslycollected specimens obtained from museums andindividuals, and those collected during field work inthese two states, resulted in the publication of Reynolds(1973a – earthworms of Delaware), and Reynolds(1973b – earthworms of Rhode Island). Thus beganmy study over 45 years ago on the distribution ofearthworms in North America and elsewhere. Theresults of these surveys can be seen in Figure 1 andTable 1, as well as contributions published with and bycolleagues and other researchers.

Over the years, my earthworm surveys haveextended beyond mainland North America, to includeBangladesh (Das and Reynolds, 2003, Reynolds,1994c, Reynolds et al., 1995); Belize (Reynolds andGuerra, 1994, Reynolds and Righi, 1994, Reynolds etal., 1995); Bermuda (Reynolds and Fragoso, 2004);Cayman Islands (Reeves and Reynolds, 2004);Contadora Island [Panama] (Reynolds and Reynolds,2002); Dakshina Kannada [India] (Siddaraju et al.,2010); Dominica (Reeves and Reynolds, 2006);Dominican Republic (Reynolds and Reynolds, 2001a,2004b, 2007); Dubai (Reynolds and Reynolds, 2010);Falkland Islands [= Ilas Malvinas] [UK] (Reynolds andJones, 2005); Gough Island [UK] (Reynolds, Jones,Gaston and Chown, 2002); Isle of Man [UK](Reynolds, 1996b); Marshall Islands (Reeves andReynolds, 2010); San Andrés [Colombia] Reynolds andReynolds, 2001b); South Africa (Reynolds andReinecke, 1977); South Korea (Hong and Reynolds,1997); Swaziland (Reynolds, 1993); Tamil Nadu[India] (Kathireswari et al., 2005); and Tristan daCunha and Nightingale Islands [UK] (Reynolds andHänel, 2005).

In recent papers (Reynolds and Wetzel, 2004,2008, 2012), we have recognized 256 speciesrepresenting 59 genera in 10 families; of these, 188species are considered native to North America, while68 have been introduced. At this time, I personallyknow of ten or more taxa from localities in the UnitedStates that have yet to be described, and that I considerto be native to North America.

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Fig. 1. Earthworm surveys in North America by various authors (map produced by the author, abbreviations are foundin Table 1).

The Status of Terrestrial Earthworm Surveys that have been conducted in North America, etc.230

Table 1. Regional Earthworm Surveys that have been conducted in extended North America.No. of Species Total No. of No. Sampled References

Region Total(native/exotic) Subdivisions 1 Subdivisions see Literature

Belize 9(2/7) 6 6 (Reynolds & Righi, 1994;Reynolds & Guerra, 1994;Reynolds et al., 1995)

Bermuda 15(1/14) 9 9 (Reynolds & Fragoso, 2004)

CanadaAlberta (AB) 11(1/11) 87 15 (Hilton & Reynolds,1983;

Scheu & McLean, 1993;Reynolds & Clapperton, 1996)

British Columbia (BC) 24(6/18) n/a n/a (Marshall & Fender, 2007)Manitoba (MB) 8(0/8) n/a n/a (Reynolds, 2000b)New Brunswick (NB) 15(1/14) 15 15 (Reynolds, 1996c, 2001b)Newfoundland &

Labrador (NL) 11(0/11) n/a n/a (Reynolds, 2000a)Nova Scotia (NS) 15(0/15) 18 18 (Reynolds, 1975d, 1976b,

2010a)Ontario (ON) 19(2/17) 54 54 (Reynolds, 1977a, 2011b,c;

Reynolds & Mayville, 1994;Reynolds & Reynolds, 1992a)

Prince Edward Island (PE) 11(0/11) 3 3 (Reynolds, 1975e)Quebec (PQ) 19(2/17) 78 74 (Reynolds, 1975a,b,c, 1976a,

2007, 2010a; Reynolds &Reynolds, 1992b)

Saskatchewan (SK) 6(0/6) n/a n/a (Reynolds & Khan, 1999)Northwest Territories (NW) 2(0/2) n/a n/a (Reynolds, 2003b; Reynolds &

Moore, 1996)Nunavut Territory (NU) 2(0/2) n/a n/a (Reynolds, 2003b, 2003c,

2004a)Yukon Territory (YK) 5(1/4) n/a n/a (Berman & Marusik, 1994;

Teale, 2007)

Cayman Islands 4(1/3) n/a n/a (Reeves & Reynolds, 2004)

Commonwealth of Dominica 6(0/6) 10 5 (Reeves & Reynolds, 2006)

Dominican Republic 10(5/5) 30 4 (Reynolds & Reynolds, 2001a,2004b, 2007)

Mexico 102(51/51) 32 31 (Fragoso, 2001, 2007; Fragoso& Reynolds, 1997; Fragoso &Rojas-Férnandez, 1994;Reynolds & Reynolds,2004a)

United StatesAlabama (AL) 28(16/12) 67 50 (Reynolds, 1994f)Alaska (AK) 11(0/11) 4 2 No survey; (Reynolds &

Wetzel, 2004, 2008, 2012)Arizona (AZ) 13(1/12) 15 11 (Reynolds, 2008m)Arkansas (AR) 35(18/17) 75 50 (Causey, 1952, 1953;

Reynolds, 2008a)California (CA) 37(13/24) 58 No survey; (Reynolds, 1999a;

Reynolds&Wetzel,2008,2012)Colorado (CO) 17(3/14) 63 51 (Damoff & Reynolds, 2004;

Reynolds,1999b; Reynolds &Damoff, 2011; Reynolds &Reeves, 2003)

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Table 1 (cont’d).Regional Earthworm Surveys that have been conducted in extended North America.No. of Species Total No. of No. Sampled References

Region Total(native/exotic) Subdivisions 1 Subdivisions see Literature

Connecticut (CT) 21(2/19) 8 8 (Reynolds, 1973cDelaware (DE) 13(4/9) 3 3 (Reynolds, 1973a)District of Columbia 9(0/9) 4 No survey; (Reynolds &

Wetzel, 2004, 2008, 2012)Florida (FL) 52(23/29) 68 58 (Reynolds, 1994d)Georgia (GA) 41(20/21) 163 96 (Reynolds, 2009c)Hawaii 33(1/32) 5 No survey; (Reynolds, 1999c;

Reynolds&Wetzel, 2008,2012)Idaho (ID) 12(1/11) 44 No survey; (Reynolds &

Wetzel, 2004, 2008, 2012)Illinois (IL) 37(17/20) 102 72 (Harman, 1960; Reynolds &

Wetzel, 2011)Indiana (IN) 38(11/27) 94 94 (Reynolds, 1994a)Iowa (IA) 10(1/9) 99 No survey; (Reynolds &

Wetzel, 2004, 2008, 2012)Kansas (KS) 16(11/5) 104 No survey; (Reynolds &

Wetzel, 2004, 2008, 2012)Kentucky (KY) 28(12/16) 120 60 (Reynolds, 2008g), + new

unpublished recordLouisiana (LA) 47(31/16) 64 64 (Reynolds, 2008e; Tandy,

1969)Maine (ME) 24(0/24) 16 16 (Reynolds, 1996a, 2008f)Maryland (MD) 32(10/22) 23 23 (Reynolds, 1974b)Massachusetts (MA) 21(3/18) 14 14 (Reynolds, 1977b)Michigan (MI) 21(6/15) 83 (Murchie, 1956; Snider, 1991)Minnesota (MN) 15(2/13) 87 51 (Reynolds, Linden & Hale,

2002)Mississippi (MS) 27(14/13) 82 63 (Reynolds, 1994e)Missouri (MO) 27(12/15) 115 55 (Olson, 1936; Reynolds,

2008h)Montana (MT) 13(1/12) 56 27 (Reynolds, 1972a, Reynolds,

2009a)Nebraska (NE) 14(5/9) 93 5 No survey; (Damoff, 2011;

Reynolds & Wetzel, 2004,2008, 2012)

Nevada (NV) 8(0/8) 17 5 No survey; (Reynolds &Wetzel, 2004, 2008, 2012)

New Hampshire (NH) 14(0/14) 10 10 (Reynolds, 2008k)New Jersey (NJ) 17(3/14) 21 21 (Reynolds, 2007c)New Mexico (NM) 11(3/8) 15 11 (Reynolds & Damoff, 2009)New York (NY) 29(4/25) 62 56 (Olson, 1940; Reynolds, 2008c)North Carolina (NC) 42(20/22) 100 80 (Reynolds, 1994b)North Dakota (ND) 18(0/18) 53 53 (Reynolds, 1978b; Utter et al.,

1995a, b)Ohio (OH) 25(12/13) 88 68 (Olson, 1928; Reynolds, 2007e)Oklahoma (OK) 21(14/7+32) 77 (Harman, 1954; Reynolds &

Damoff, 2010)Oregon (OR) 47(23/24) 36 Macnab & McKey-Fender,

1947; (Reynolds & Wetzel,2004, 2008, 2012)

Pennsylvania (PA) 28(8/20) 67 51 (Reynolds, 2008b)Puerto Rico 33(14/19) 8 8 No survey; (Reynolds &

Wetzel, 2004, 2008, 2012)Rhode Island (RI) 12(1/11) 5 5 (Reynolds, 1973b, 2002a)

The Status of Terrestrial Earthworm Surveys that have been conducted in North America, etc.232

Table 1 (cont’d).Regional Earthworm Surveys that have been conducted in extended North America.No. of Species Total No. of No. Sampled References

Region Total(native/exotic) Subdivisions 1. Subdivisions see Literature

South Carolina (SC) 25(4/21) 47 37 (Reynolds, 2001a; Reynolds &Reeves, 2004)

South Dakota (SD) 11(0/11) 67 37 (Reynolds, 2007a)Texas (East TX only) 57(27+63/19) 37 32 (Damoff & Reynolds, 2009)Tennessee (TN) 47(254/22) 95 95 (Damoff & Reynolds, 2013;

Reynolds, 1974a, 1977d,1977e, 1978a; Reynolds etal., 1974)

Utah (UT) 13(0/13) 29 26 (Reynolds, 2008i)Vermont (VT) 15(0/15) 14 14 (Reynolds, 2008d, 2012b)Virginia (VA) 37(15/22) 98 75 (Reynolds, 1994c)Washington ((WA) 31(16/15) 39 (Altman, 1936; Macnab &

McKey-Fender,1947;Reynolds& Wetzel, 2004, 2008, 2012)

West Virginia (WV) 18(4/14) 52 25 (Reynolds, 2007d)Wisconsin (WI) 13(2/11) 72 No survey; (Reynolds &

Wetzel, 2008, 2012)Wyoming (WY) 13(1/12) 23 22 (Reynolds, 2003a; Reynolds et

al., 2009)1 County, District, Municipality, Parish, State.2 3 Dilocardia n.sp., yet to be described.3 6 Diplocardia n.sp., yet to be described.4 1+ Diplocardia n.sp. yet to be described.

DISCUSSION

When I began my earthworm studies, only afew geographical earthworm surveys in the UnitedStates had been published – Olson, 1928 (Ohio), 1936(Missouri), 1940 (New York); Altman, 1936(Washington); Eaton, 1942 (northeastern U.S.);Causey, 1952 (Arkansas), and Murchie, 1956(Michigan). It was not my original plan to conductstate and provincial surveys, but rather to study theecology and soil relationships of earthworms.However, the lack of earthworm taxonomists and moreimportantly, the paucity of useful keys to species anddistributional studies, took me in a different direction.My extensive career working with the distribution andtaxonomy of megadriles began with my dissertation onthe earthworms occurring in the state of Tennessee(Reynolds et al., 1974; Reynolds, 1974a, 1977d, 1977e,1978a). It is important to note that inspiration wasprovided by my major professor at the University ofTennessee, Dr. Ed Clebsch, who spent the summer of1972 teaching at the University of Montana. Duringthat time, he made a number of collections ofearthworms, as little was known about the fauna in thatstate. Clebsch’s data formed the basis of my firstearthworm survey (Reynolds, 1972a) which was

expanded in Reynolds (2009a). Another importantmentor, Gordon Gates (at the time, on staff at the TallTimbers Research Station [TTRS], Tallahassee,Florida) informed me that several states, notablyDelaware and Rhode Island, had no known earthwormrecords. After graduation from the University ofTennessee, I was hired by TTRS with the freedom toconduct earthworm research anywhere, but particularlyin North America (see Crawford and Brueckheimer,2012). Thus began a 45 year venture of sampling inNorth America and numerous other locations in theWestern Hemisphere and beyond.

While surveying Delaware and Rhode Island(Reynolds, 1973a and 1973b, respectively), I used theopportunity to also conduct preliminary surveys forearthworms in Connecticut (Reynolds, 1973c),Maryland (Reynolds, 1974b) and Massachusetts(Reynolds, 1977b). In 1973, we moved fromTennessee to Fredericton, New Brunswick. Aided bya travelling field laboratory provided by Tall TimbersResearch Station, we began detailed collections inNova Scotia (Reynolds, 1976b, revised and updated in2010a), Prince Edward Island (Reynolds, 1975e), theMagdalen Islands (Reynolds, 1975a), Cape Breton(Reynolds, 1975d), and the south shore of Quebec

MEGADRILOGICA 233

(Reynolds, 1975b, 1975c, 1976a). A survey of theearthworms occurring along the north shore of Quebecwas begun in 1976 while working on a summer projectat the Université de Québec á Trois Rivières – a surveyeventually completed and published many years later(Reynolds and Reynolds, 1992b). A revised andupdated survey of earthworms of the entire province ofQuebec was published in 2010 (Reynolds, 2010a).While still completing my doctoral studies at theUniversity of Tennessee, a survey of earthwormsoccurring in southern Ontario was undertaken in 1972–eventually published as a book by the Royal OntarioMuseum (Reynolds, 1977a). Several notes and paperson the distribution of megadriles in finite areas withinOntario have subsequently been published, e.g., thosefor Victoria Co. (Reynolds and Reynolds, 1992a),Rainy River District (Reynolds and Mayville, 1994),Guelph Arboretum(Reynolds, 2011b), and Pelee Island(Reynolds, 2011c).

During the early 1970s, I teamed up with Dr.David Cook of the Fisheries and Marine Service at theCanada Centre for Inland Waters [Burlington, Ontario],to compile Nomenclatura Oligochaetologica – Acatalogue of names, descriptions and type specimens ofthe Oligochaeta; the original volume was published in1976, which included entries for 24 families, 573genera, and 5753 species (Reynolds and Cook, 1976 –see Appendix, pp. 248-9). Subgenera were notincluded until the second supplement (Reynolds andCook, 1989). During 1975 and 1977, I published twopapers which included two new species ofSparganophilus – S. pearsei from North Carolina(Reynolds, 1975f), and S. tennesseensis fromTennessee (Reynolds, 1977d). These species havesubsequently been collected elsewhere by others.

In 1980, I published the first revision of theearthwormgenus Sparganophilus which included eightnew taxa (Reynolds, 1980).

During the 1980s, I redirected my primaryinterests to the study of law and political science, yetcontinued to amass collections of earthworms fromvarious localities with extensive assistance fromcolleagues and other researchers. I continued to workwith Dave Cook on supplements to the nomenclatorcatalogue, publishingNomenclatura OligochaetologicaSupplementum Primum (Reynolds and Cook, 1981) andNomenclatura Oligochaetologica SupplementumSecundum (Reynolds and Cook, 1989).

In 1991, I returned to academic work at SirSandford Fleming College, began collecting andobtaining specimens from numerous states andprovinces, and completed Nomenclatura Oligo-chaetologica Supplementum Tertium (Reynolds andCook, 1993).

Many years before, I had been given the JimJoyner collection of Indiana earthworms, and togetherwith specimens I collected while at Purdue Universityand afterwards, the Indiana survey was finallypublished (Reynolds, 1994a). The Indiana surveyprovided the opportunity to present fully the PostQuaternary Introduction Theory for megadrilemigration in North America. During the early throughmid-1990s, I had the opportunity to travel, collect, andpublish on earthworm surveys in Bangladesh (Das andReynolds, 2003; Reynolds, 1994c; Reynolds, Julka andKhan, 1995) and Belize (Reynolds and Righi, 1994,Reynolds and Guerra, 1994; Reynolds, Fragoso andGuerra, 1995). I also finalized collection data gatheredover the years (alone and with the aid of colleagues,primarily from Tall Timbers Research Station) forsurveys in North Carolina (Reynolds, 1994b), Virginia(Reynolds, 1994c), Florida (Reynolds, 1994d),Mississippi (Reynolds, 1994e) and Alabama (1994f).

In 1995, Dr. Jill Clapperton invited me toAlberta to conduct a training session focussing onearthworm collection, processing, and identification atAgriculture Canada in Lethbridge. During that visit,we took the opportunity to travel and samplethroughout the province; we subsequentlypublished theresults of our survey in Reynolds and Clapperton(1996), including the description of a new species,Aporrectodea bowcrowensis. This paper augmented anearlier report (Hilton and Reynolds, 1983) ofearthworms in Alberta.

I also had the opportunity to work with YongHong on an introductory paper on the earthworms ofKorea (Hong and Reynolds, 1997).

Soon after the publication of a paper onearthworms of southeastern Mexico (Fragoso andRojas-Férnandez, 1994 – in which two new genera andfive new species in the family Megascolecidae weredescribed), I was invited by Fragoso to summarizeearthworm collections obtained from Dr. Clebschduring one of his earlier (1973) summer surveys whenvisiting that country. Our subsequent publication ofClebsch’scollections (Fragoso and Reynolds, 1997)

The Status of Terrestrial Earthworm Surveys that have been conducted in North America, etc.234

included the description of two new species to science:Dichogaster michoacana and D. jalischensis. Oursecond paper together was published seven years later(Reynolds and Fragoso, 2004).

Three small notes were published in 1999(California, Colorado, Hawaii (Reynolds, 1999a,1999b, and 1999c, respectively), plus Saskatchewan(Reynolds and Khan, 1999).

With samples from colleagues and personsinterested in earthworms, surveys were published forNewfoundland and Labrador and Manitoba (both in2000a and b, respectively). In 2001, my first paper onearthworms in South Carolina was published(Reynolds, 2001a). The publication of this paper ledme to a wonderful collaboration with Dr. Will K.Reeves, first expanding on the distribution ofearthworms in South Carolina (Reynolds and Reeves,2004), then to surveys in other states and far beyond:Wyoming revisited (Reynolds et al., 2009), CaymanIslands (Reeves and Reynolds, 2004), Colorado(Reynolds and Reeves, 2003), Dominica (Reeves andReynolds, 2006), and the Marshall Islands (Reeves andReynolds, 2010).

The year 2001 was also the beginning of mycollaboration with my daughter, Debbie, on theearthworms in various Caribbean and Central Americanlocations: Dominican Republic No. 1 (Reynolds andReynolds, 2001a), San Andrès, Colombia (Reynoldsand Reynolds, 2001b), Contadora Island, Panama(Reynolds and Reynolds, 2002), Nuevo Léon, México(Reynolds and Reynolds, 2004a), Dominican RepublicNo. 2 (Reynolds and Reynolds, 2004b), Honduras(Reynolds and Reynolds, 2006), and DominicanRepublic No. 3 (Reynolds and Reynolds, 2007 – allpublished in Spanish. An earthworm survey of NewBrunswick was also published in 2001 (Reynolds,2001b).

With the help of two colleagues, DennisLinden and Cindy Hale, we published the earthwormsof Minnesota (Reynolds, Linden and Hale, 2002). In2003, I published the first paper on the earthworms ofWyoming (Reynolds, 2003a); and the first paper ofearthworms from the new Canadian territory ofNunavut (Reynolds, 2003b). Specimens in the museumat Clemson University (Clemson, South Carolina)provided a second species from Nunavut (Reynolds,2003c).

There are still very few collection recordsfrom the northern Canadian territories: NorthwestTerritories (Reynolds and Moore, 1996; Reynolds2003b), Nunavut (Reynolds, 2003b, 2003c, 2004a) andYukon (Berman and Marusik, 1994; Teale, 2007).

In collaboration with other colleagues, I wasfortunate to work with on earthworm surveys in theSouth Atlantic: Gough Island (Reynolds, Jones, Gastonand Chown, 2002), Tristan da Cunha and NightingaleIslands (Reynolds and Hänel, 2005), and the FalklandIslands (= Ilas Malvinas) (Reynolds and Jones, 2005).

In 1996, the Bermuda Zoo, in conjunctionwith the government of Bermuda [Contribution No. 83,Bermuda Biodiversity Project, Bermuda AquariumMuseum and Zoo], contracted me to conduct a surveyof Bermudan earthworms as part of their biodiversityproject. Due to changes in my employment, that surveywas not published until eight years later (Reynolds andFragoso, 2004). As published in that paper, this surveyresulted in the collection of specimens that allowed usto establish a neotype for Onychochaeta windlei,subsequently deposited in the New BrunswickMuseum, Saint John.

In 2006, after nine years with SchneiderNational Carriers Inc., I retired to devote myself fulltime to earthworm research. During my years withSchneider, I had the help of numerous colleagues therein obtaining samples from various locations throughoutNorth America. In 2007, I returned to surveys in NorthAmerica, with the publication of the earthworms ofSouth Dakota (Reynolds, 2007a), New Jersey(Reynolds, 2007c), West Virginia (Reynolds, 2007d)and an updated survey for Ohio (Reynolds, 2007e).

The year 2008 was extremely busy, with thepublication of twelve earthworm surveys: Arkansasrevisited (Reynolds, 2008a), an update to Causey(1952, 1953); Pennsylvania (Reynolds, 2008b); NewYork revisited (Reynolds, 2008c), an update to Olson(1940); Vermont (Reynolds, 2008d and subsequently2012b); Louisiana revisited (Reynolds, 2008e), as an allspecies update to Tandy (1969, which only includedPheretima species); Maine (Reynolds, 2008f –expanding Reynolds, 1996a); Kentucky (Reynolds,2008g); Missouri revisited (Reynolds, 2008h), anupdate to Olson (1936); Utah (Reynolds, 2008i); NewHampshire (Reynolds, 2008k); and Arizona (Reynolds,2008m). Reynolds (2008j) presented an update of thefamily Sparganophilidae in North America.

MEGADRILOGICA 235

Dr. George Damoff enlisted my assistance ona paper summarizing the distribution of earthworms inColorado (Damoff and Reynolds, 2004). This led to astrong friendship and subsequent collaboration onseveral surveys and their publications presentingdistributional information for earthworms in NewMexico (Reynolds and Damoff, 2009), east Texas(Damoff and Reynolds, 2009), Oklahoma (Reynoldsand Damoff, 2010) and then a second paper on theearthworms of Colorado (Reynolds and Damoff, 2011),which included numerous collection records suppliedby Dr. Whitney Cranshaw of Colorado StateUniversity, Fort Collins.

The last single state paper to date focussing onthe earthworm fauna of the American southeast waspublished for Georgia (Reynolds, 2009c), with manyspecimens provided by Dr. Will Reeves resulting fromhis collections in counties in that state with no previousrecords.

The most recent single state survey ofearthworms in the U.S. was published by Reynolds andWetzel (2011) for Illinois, augmenting WalterHarman’s (1960) doctoral dissertation on theearthworms of central Illinois counties, and includinga lengthy history of oligochaete research in the state.This Illinois survey represented over 15 years of closecollaboration with Mark Wetzel of the Illinois NaturalHistory Survey on numerous earthworm projects,including papers on North American distribution(Reynolds and Wetzel, 2004, 2008, 2012), the secondedition of Nomenclatura Oligochaetologica, andediting assistance on numerous papers published in thejournal, Megadrilogica, and elsewhere.

During the past two years, I have merged theindividual state and provincial surveys into regionalpapers, including maps of all the counties, to create abetter understanding of the distribution of earthwormsand how they might relate to physiography, soils andvegetation; these include the American Northeast, fromMaine to West Virginia (Reynolds, 2010b); the AtlanticMaritime Ecosystem in Canada, covering NewBrunswick, Nova Scotia, Prince Edward Island,Southern Quebec, and the Magdalen Islands (Reynolds,2010c); American Southeast, from Virginia to eastTexas (Reynolds, 2011a); American Midwest, fromOhio to Minnesota (Reynolds, 2011d); and AmericanNorth-Central, including the Dakotas, Montana,Wyoming, Colorado, and Utah (Reynolds, 2012a).

At this time, there are no provinces, territories,or states in Canada or the United States which do nothave several earthworm species recorded from them.For Mexico, only the state of Coahuila is without arecorded earthworm – in spite of our collecting effortsin that state (with daughter, Debbie, and wife, Wilma).More collecting during the wet months of the yearmight be successful, even though Coahila is a very aridstate. There are no detailed or wide rangingcollections, i.e. surveys, from the following U.S. states,districts, or territories: Alaska, California, District ofColumbia, Hawaii, Idaho, Iowa, Kansas, Nebraska,Nevada, Puerto Rico, and Wisconsin. Although thereare several collections of earthworms that have beenreported from the northern Canadian territories, theyare from extremely limited areas; fortunately, there areseveral younger colleagues who are now picking up thechallenge to obtain specimens from these under-sampled areas. I also have been assisting graduatestudents in several countries with their own faunisticstudies focussing on oligochaetes. As much as I wouldlike to travel and collect in these other regions, I thinkit will be up to others to fill in the missing areas on themap (Fig. 1).

Current Projects.

When I visited the Illinois Natural HistorySurvey in December 2011 to finalize our paper onearthworms of Illinois with Mark Wetzel (Reynoldsand Wetzel, 2011), we made plans for the Spring of2012 to conduct surveys in 27 counties in that statefrom which no records of specimens were available;unfortunately, the lack of precipitation during thespring and summer months forced us to postpone thatfield work until 2013.

Although my survey work in North Americamay be limited in the future, there are other challengesin oligochaetology which require my attention.

In 1974, Dr. Gordon Gates gave me a box of3x5 cards on which information for originaldescriptions of oligochaete taxa had been recorded.This simple ‘catalog’ of records was the beginning ofNomenclatura Oligochaetologica (Reynolds and Cook,1976). For several years I worked with David Cook onthe initial book, establishing a format with a six-language introduction, and expanding on theinformation from Gates to be presented for each taxonentry [account], e.g.: name of the taxon (genus, species,subspecies), authority name(s), year of published

The Status of Terrestrial Earthworm Surveys that have been conducted in North America, etc.236

description, name of publication (for journals, thevolume and issue, and for books, title, chapter, edition,as applicable), page of that publication on which thedescription begins, and the institution(s) and accessionnumbers for the holotype and paratypes in thoseinstitutions. Indexes for the authors, full titles ofpublications, and contact information for the museumsand other places where type material had beendeposited were also included in this first nomenclator.When I presented Dr. Gates with his copy he wasimpressed with what we had done with his box of cards(see Appendix, pp. 248-9). During the followingdecades David Cook and I worked on publishing threesupplements (Reynolds and Cook, 1981, 1989 and1993). After David Cook retired, I invited MarkWetzel to be the new co-author for the fourthsupplement. Recently, we decided to redirect ourefforts into the compilation of a second edition of thenomenclator – bringing together the accounts in theoriginal book with those presented in the threesupplements, plus those published since 1993 thatwould have comprised the fourth supplement. I wouldexpect this project to be completed within one yearfrom this publication.

ACKNOWLEDGEMENTS

The author wishes to thank his colleagues atSchneider National Carriers, as well as colleagues fromother organizations with whomI have collaborated overthe years – for their assistance in obtaining additionalsamples from North America and other locations, andin co-authoring publications discussed above.Appreciation is also extended to Linda Ward and othersat the United States National Museum – SmithsonianInstitution, who have arranged access to material attheir institution for examination over the years. Overthe years, my family has been very supportive,understanding and in some cases, extremely helpful inco-authoring a number of papers: my wife Wilma -three papers, eldest daughter Kristin - one paper,middle daughter Debbie - ten papers, and youngestdaughter Jennifer - one paper. I also acknowledge thehelp of Dr. Catalina de Mischis, Universidad Nacionalde Córdoba, Argentina, Dr. Jean-Marc Gagnon,Canadian Museum of Nature, Ottawa, and Dr. MirjanaStojanovi, University of Kragujevac, Serbia forspecial assistance. My gratitude extends to Wilma M.Reynolds of the Oligochaetology Laboratory, Mark J.Wetzel, Illinois Natural History Survey, and GeorgeDamoff, Texas for comments and suggestions duringtheir reviews of this paper.

EARTHWORM SURVEY REFERENCES

BELIZE

Reynolds, J.W. and C.A. Guerra. 1994. Two speciesof earthworms newly reported from Belize,C.A. (Oligochaeta: Glossoscolecidae andMegascolecidae). Megadrilogica 5(11): 122-124.

Reynolds, J.W. and G. Righi. 1994. On some earth-worms from the Belize, C.A. with thedescription of a new species (Oligochaeta:Acanthodrilidae, Glossoscolecidae and Octo-chaetidae). Megadrilogica 5(9): 97-106.

Reynolds, J.W., C. Fragoso and C.A. Guerra. 1995.New earthworm records from Belize, C.A.(Oligochaeta: Acanthodrilidae, Eudrilidae,Glossoscolecidae, Megascolecidae, Ocnero-drilidae and Octochaetidae). Megadrilogica6(7): 63-70.

BERMUDA

Reynolds, J.W. and C. Fragoso. 2004. The earth-worms (Oligochaeta: Acanthodrilidae,Eudrilidae, Glossoscolecidae, Lumbricidae,Megascolecidae and Ocnerodrilidae) ofBermuda. Megadrilogica 10(4):17-26.

CANADA

Alberta

Hilton, D.F.J. and J.W. Reynolds. 1983. The earth-worm (Oligochaeta: Lumbricidae) fauna ofthe Cypress Hills Provincial Park, Alberta.Megadrilogica 4(4): 91-92.

Reynolds, J.W. and M.J. Clapperton. 1996. Newearthworm records for Alberta (Oligochaeta:Lumbricidae) including the description of anew Canadian species. Megadrilogica 6(8):73-82.

Scheu, S. and M.-A. McLean. 1993. The earthworm(Lumbricidae) distribution in Alberta(Canada). Megadrilogica 4(11): 175-180.

MEGADRILOGICA 237

British Columbia

Marshall, V.G. and W.M. Fender. 2007. Native andintroduced earthworms (Oligochaeta) ofBritish Columbia, Canada. Megadrilogica11(4): 29-52.

Manitoba

Reynolds, J.W. 2000b. A contribution to our know-ledge of the earthworm fauna of Manitoba,Canada (Oligochaeta, Lumbricidae). Mega-drilogica 8(3): 9-12.

New Brunswick

Reynolds, J.W. 1976c. A preliminary checklist anddistribution of the earthworms of NewBrunswick. New Brunswick Nat. 7(2): 16-17.

Reynolds, J.W. 2001b. The earthworms of NewBrunswick(Oligochaeta: Lumbricidae) Mega-drilogica 8(8): 37-47.

Newfoundland and Labrador

Reynolds, J.W. 2000a. A contribution to our know-ledge of the earthworm fauna of Newfound-land and Labrador, Canada (Oligochaeta,Lumbricidae). Megadrilogica 8(2): 5-8.

Northwest Territories

Reynolds, J.W. 2003b. First earthworm record fromNunavut, Canada and second from the North-west Territories. Megadrilogica 9(6): 40.

Reynolds, J.W. and S.M. Moore. 1996. Note on thefirst earthworm record from the NorthwestTerritories, Canada. Megadrilogica 6(10): 96.

Nova Scotia

Reynolds, J.W. 1975d. Boiteagan (Oligochaeta:Lumbricidae) Cheap Breatunn. Megadril-ogica 2(6): 1-7.

Reynolds, J.W. 1976b. The distribution and ecologyof the earthworms of Nova Scotia. Mega-drilogica 2(8): 1-8.

Reynolds, J.W. 2010a. The earthworms (Oligochaeta:Lumbricidae) of Nova Scotia, revisited.Megadrilogica 14(6): 77-100.

Nunavut

Reynolds, J.W. 2003b. First earthworm record fromNunavut, Canada and second from theNorthwest Territories. Megadrilogica 9(6):40.

Reynolds, J.W. 2003c. A second earthworm species(Lumbricidae) from Nunavut, Canada. Mega-drilogica 9(8): 52.

Reynolds, J.W. 2004a. More earthworm (Lumbric-idae) collections from Nunavut, Canada.Megadrilogica 10(3): 13-14.

Ontario

Reynolds, J.W. 1977a. The earthworms (Lumbricidaeand Sparganophilidae) of Ontario. Life Sci.Misc. Publ., Roy. Ont. Mus. xi + 141 pp.

Reynolds, J.W. 2011b. Earthworms of the Universityof Guelph Arboretum. Megadrilogica 15(2):15-20.

Reynolds, J.W. 2011c. The earthworms (Oligochaeta:Lumbricidae) of Pelee Island, Ontario,Canada. Megadrilogica 15(3): 23-33.

Reynolds, J.W. and P.N. Mayville. 1994. Newearthworm records from Rainy River Districtin north western Ontario (Oligochaeta:Lumbricidae). Megadrilogica 6(2): 13-16.

Reynolds, J.W. and J.W. Reynolds. 1992a. AdditionalOntario earthworm county records (Lumbric-idae). Megadrilogica 4(8): 144.

Prince Edward Island

Jordan, G.A., J.W. Reynolds and A.J. Burnett. 1976.Computer plotting and analysis of earthwormpopulation distribution in Prince EdwardIsland. Megadrilogica 2(10): 1-7.

Reynolds, J.W. 1975e. The earthworms (Oligochaeta:Lumbricidae) of Prince Edward Island.Megadrilogica 2(7): 4-10.

The Status of Terrestrial Earthworm Surveys that have been conducted in North America, etc.238

Quebec

Reynolds, J.W. 1975a. Les lombricidés (Oligochaeta)des Îles-de-la-Madeleine. Megadrilogica 2(3):1-8.

Reynolds, J.W. 1975b. Les lombricidés (Oligochaeta)de la Gaspésie, Québec. Megadrilogica 2(4):4-9.

Reynolds, J.W. 1975c. Les lombricidés (Oligochaeta)de l'Île d'Orléans, Québec. Megadrilogica2(5): 8-11.

Reynolds, J.W. 1976a. Catalogue et clé d'identifica-tion des lombricidés du Québec. Naturalistecanadien 103(1): 21-27.

Reynolds, J.W. 2007. First earthworm records fromWentworth Township, Argenteuil County,Quebec. [Premiers signalements de lombricsdans le canton de Wentworth, comté d’Argen-teuil (Québec)] Megadrilogica 11(5): 58-62.

Reynolds, J.W. 2010a. The earthworms (Oligochaeta:Lumbricidae and Sparganophilidae) ofQuebec, Canada, revisited. Megadrilogica14(1): 1-46.

Reynolds, J.W. and K.W. Reynolds. 1992b. Les versde terre (Oligochaeta: Lumbricidae etSparganophilidae) sur la rive nord du Saint-Laurent (Québec). Megadrilogica 4(9): 145-161.

Saskatchewan

Reynolds, J.W. and M.N. Khan. 1999. A contributionto our knowledge of the earthworm fauna ofSaskatchewan, Canada. Megadrilogica 7(12):81-82.

Yukon

Berman, D.I. and Y.M. Marusik. 1994. On Bimastosparvus (Oligochaeta: Lumbricidae) fromYukon Territory (Canada), with discussion ofdistribution of the earthworms in north-western North America and northeasternSiberia. Megadrilogica 5(10): 113-116.

Teale, C.L. 2007. A preliminary survey of theOligochaete fauna of the Yukon Territory,Canada. Megadrilogica 11(1): 3-7.

CAYMAN ISLANDS

Reeves, W.K. and J.W. Reynolds. 2004. Earthworms(Oligochaeta: Glossoscolecidae, Megascolec-idae, and Octochaetidae) and terrestrialpolychaetes (Polychaeta: Nereididae) ofGrand Cayman Island, with notes on theirnatural enemies. Megadrilogica 19(6): 39-41.

DOMINICA

Reeves, W.K. and J.W. Reynolds. 2006. A note onsome earthworms (Annelida: Oligochaeta)from the Commonwealth of Dominica.Megadrilogica 10(11): 91-93.

DOMINICAN REPUBLIC

Reynolds, J.W. and D.W. Reynolds. 2001a. Nuevosdatos de lombrices de tierra (Oligochaeta) dela República Dominicana. Megadrilogica8(5): 17-19.

Reynolds, J.W. and D.W. Reynolds. 2004b. Nuevosregistros de lombrices de tierra (Oligochaeta:Glossoscolecidae) de la República Domini-cana. Megadrilogica 10(3): 14-16.

Reynolds, J.W. and D.W. Reynolds. 2007. Primerosexpedientes de la lombrices de tierra (Oligo-chaeta) de la provincia de Espaillat, RepúblicaDominicana. Megadrilogica 11(2): 19-20.

MEXICO

Fragoso, C. 2001. Las lombrices de tierra de México(Annelida, Oligochaeta): diversidad, ecologíay manejo. Acta Zool. Mexicana (n.s.),Número especial 1: 131-171.

Fragoso, C. 2007. Diversidad y patrones biogeo-gráficos de las lombrices de tierra de México(Oligochaeta, Annelida). Pp. 108-124, In:Brown, G.G. and C. Fragoso (eds.). Min-hocas na América Latina: Biodiversidade eEcologia. Londrina: Embrapa Soja, 545 pp.

MEGADRILOGICA 239

Fragoso, C. and P. Rojas-Férnandez. 1994. Earth-worms from southeastern Mexico, newacanthodriline genera and species (Mega-scolecidae, Oligochaeta). Megadrilogica 6(1):1-12.

Fragoso, C. and J.W. Reynolds. 1997. On someearthworms from central and southeasternMexican mountains, including two newspecies of the genus Dichogaster (Dicho-gastrini). Megadrilogica 7(2): 9-20.

Reynolds, J.W. and D.W. Reynolds. 2004a. Nuevosregistros de las Grandes Carreras VI-VIIMundiales de Lombrices de tierra desdeNuevo León (México). Megadrilogica 10(2):5-11.

UNITED STATES OF AMERICA

Alaska - no formal earthworm survey, only incidentalrecords (Reynolds and Wetzel, 2004, 2008).

Alabama

Reynolds, J.W. 1994f. Earthworms of Alabama(Oligochaeta: Acanthodrilidae, Eudrilidae,Lumbricidae, Megascolecidae and Spargano-philidae). Megadrilogica 6(4): 35-46.

Arizona

Reynolds, J.W. 2008m. The earthworms (Oligo-chaeta: Acanthodrilidae, Lumbricidae, andOcnerodrilidae) of Arizona, USA. Mega-drilogica 12(11): 155-166.

Arkansas

Causey, D. 1952. The earthworms of Arkansas. Proc.Arkansas Acad. Sci. 5: 31-42.

Causey, D. 1953. Additional records for Arkansasearthworms. Proc. Arkansas Acad. Sci. 6: 47-48.

Reynolds, J.W. 2008a. The earthworms (Oligochaeta:Acanthodrilidae, Lumbricidae, Megascolec-idae and Sparganophilidae) of Arkansas,USA, revisited. Megadrilogica 11 (11): 115-130.

California - no formal earthworm survey, onlyincidental records (Reynolds, 1999a;Reynolds and Wetzel, 2004, 2008, 2012).

Colorado

Damoff, G.A. and J.W. Reynolds. 2004. Additionalearthworms (Oligochaeta: Lumbricidae) fromColorado, USA. Megadrilogica 10(5) 29-37.

Reynolds, J.W. 1999b. A contribution to our know-ledge of the earthworm fauna of Colorado.Megadrilogica 7(10): 71-72.

Reynolds, J.W. and G.A. Damoff. 2011. More earth-worms (Oligochaeta: Lumbricidae andSparganophilidae) from Colorado, USA.Megadrilogica 14(8): 159-172.

Reynolds, J.W. and W.R. Reeves. 2003. The earth-worms (Oligochaeta: Lumbricidae) of Color-ado, USA. Megadrilogica 9(7): 41-48.

Connecticut

Reynolds, J.W. 1973c. The earthworms of Connecti-cut (Oligochaeta: Lumbricidae, Megascolec-idae and Sparganophilidae). Megadrilogica1(7): 1-6.

Delaware

Reynolds, J.W. 1973a. The earthworms of Delaware(Oligochaeta: Acanthodrilidae and Lumbric-idae). Megadrilogica 1(5): 1-4.

District of Columbia - no formal earthworm survey,only incidental records (Reynolds and Wetzel,2004, 2008, 2012).

Florida

Reynolds, J.W. 1994d. Earthworms of Florida (Oligo-chaeta: Acanthodrilidae, Eudrilidae, Glosso-scolecidae, Lumbricidae, Megascolecidae,Ocnerodrilidae, Octochaetidae and Spar-ganophilidae). Megadrilogica 5(12): 125-141.

Georgia

Reynolds, J.W. 2009c. The earthworms (Oligochaeta:Acanthodrilidae, Eudrilidae, Lumbricidae,Megascoecidae, Ocnerodrilidae, and Spar-ganophilidae) of Georgia, USA. Megadril-ogica 13(7): 85-111.

The Status of Terrestrial Earthworm Surveys that have been conducted in North America, etc.240

Hawaii - no formal earthworm survey, only incidentalrecords (Reynolds, 1999c; Reynolds andWetzel, 2004, 2008, 2012).

Idaho - no formal earthworm survey, only incidentalrecords (Gates, 1967b; Reynolds and Wetzel,2004, 2008, 2012).

Illinois

Harman, W.J. 1960. Studies on the taxonomy andmusculature of the earthworms of centralIllinois. Ph.D. dissertation, Champaign, IL:Univ. of Illinois, 107 pp.

Reynolds, J.W. and M.J. Wetzel. 2011. The earth-worms (Oligochaeta: Acanthodrilidae,Komarekionidae, Lumbricidae, Megascolec-idae and Sparganophilidae.) of Illinois, USA.Megadrilogica 15(4): 35-67.

Wetzel, M.J. and J.W. Reynolds. 2012. The earth-worms (Annelida, Clitellata, Oligochaeta) ofIllinois – Introduction and annotated checklistof species, World Wide Web URL:[http://www.inhs.uiuc.edu/~mjwetzel/IllinoisEarthworms.html]

Indiana

Reynolds, J.W. 1994a. The distribution of the earth-worms (Oligochaeta) of Indiana: a case for thePost Quaternary Introduction Theory ofmegadrile migration in North America.Megadrilogica 5(3): 13-32.

Iowa - no formal earthworm survey, only incidentalrecords (Reynolds and Wetzel, 2004, 2008,2012).

Kansas - no formal earthworm survey, only incidentalrecords (James, 1984, 1988; Reynolds andWetzel, 2004, 2008, 2012).

Kentucky

Reynolds, J.W. 2008g. The earthworms (Oligochaeta:Acanthodrilidae, Komarekionidae, Lumbric-idae, Megascolecidae, and Sparganophilidae)of Kentucky, USA. Megadrilogica 12(5):73-88.

Louisiana

Reynolds, J.W. 2008e. The earthworms (Oligochaeta:Acanthodrilidae, Lumbricidae, Lutodrilidae,Megascolecidae, and Sparganophilidae) ofLouisiana, USA, revisited. Megadrilogica12(3): 31-60.

Tandy, R.E. 1969. The earthworm genus PheretimaKinberg, 1866, in Louisiana. Ph.D.dissertation, Baton Rouge: Louisiana StateUniv., 155 pp.

Maine

Reynolds, J.W. 1996a. Earthworm diversity in Maine.pp. 135-139, In: Gawler, S.C., J.J. Albright,P.D. Vickery and F.C. Smith (eds.).Biological Diversity in Maine. Appendices.Augusta: Maine Natural Areas Program, Dept.Conserv., 170 pp.

Reynolds, J.W. 2008f. The earthworms (Oligochaeta:Acanthodrilidae, Lumbricidae, and Mega-scolecidae) of Maine, USA. Megadrilogica12(4): 61-72.

Maryland

Reynolds, J.W. 1974b. The earthworms of Maryland(Oligochaeta: Acanthodrilidae, Lumbricidae,Megascolecidae and Sparganophilidae.Megadrilogica 1(11): 1-12.

Massachusetts

Reynolds, J.W. 1977b. The earthworms of Mass-achusetts (Oligochaeta: Lumbricidae, Mega-scolecidae and Sparganophilidae). Mega-drilogica 3(2): 49-54.

Michigan

Murchie, W.R. 1956. Survey of Michigan earthwormfauna. Pap. Michigan Acad. Sci. Arts. Lett.41: 53-72.

Snider, R.M. 1991. Checklist and distribution ofMichigan earthworms. Michigan Academi-cian 24: 105-114.

MEGADRILOGICA 241

Minnesota

Reynolds, J.W., D.R. Linden and C.M. Hale. 2002.The earthworms of Minnesota (Oligochaeta:Acanthodrilidae, Lumbricidae and Mega-scolecidae). Megadrilogica 8(12): 85-99.

Mississippi

Reynolds, J.W. 1994e. Earthworms of Mississippi(Oligochaeta: Acanthodrilidae, Lumbricidae,Megascolecidae, Ocnerodrilidae and Spar-ganophilidae). Megadrilogica 6(3): 21-33.

Missouri

Olson, H.W. 1936. Earthworms of Missouri. Ohio J.Sci. 36(2): 102-113.

Reynolds, J.W. 2008h. The earthworms (Oligochaeta:Acanthodrilidae, Lumbricidae, Megascolec-idae, and Sparganophilidae) of Missouri,USA, revisited. Megadrilogica 12(6): 89-104.

Montana

Reynolds, J.W. 1972a. A contribution to the earth-worm fauna of Montana. Proc. MontanaAcad. Sci. 32(1): 6-13.

Reynolds, J.W. 2009a. The earthworms (Oligochaeta:Lumbricidae) of Montana, USA revisited.Megadrilogica 13(2): 9-20.

Nebraska - no formal earthworm survey, onlyincidental records (Damoff, 2011; Reynoldsand Wetzel, 2004, 2008, 2012).

Nevada - no formal earthworm survey, only incidentalrecords.

New Hampshire

Reynolds, J.W. 2008k. The earthworms (Oligochaeta:Lumbricidae) of New Hampshire, USA.Megadrilogica 12(10): 145-153.

New Jersey

Reynolds, J.W. 2007c. The earthworms (Oligochaeta:Lumbricidae, Megascolecidae and Spar-ganophilidae) of New Jersey, USA. Mega-drilogica 11(6): 63-72.

New Mexico

Reynolds, J.W. and G.A. Damoff. 2009. The earth-worms (Oligochaeta: Acanthodrilidae andLumbricidae) of New Mexico, USA. Mega-drilogica 13(5): 37-52.

New York

Olson, H.W. 1940. Earthworms of New York state.Amer. Mus. Nov., No. 1090, 9 pp.

Reynolds, J.W. 2008c. The earthworms (Oligochaeta:Lumbricidae, Megascolecidae, and Spargano-philidae) of New York, USA, revisited.Megadrilogica 12(1): 1-17.

North Carolina

Reynolds, J.W. 1994b. Earthworms of North Carolina(Oligochaeta: Acanthodrilidae, Komarekion-idae, Lumbricidae, Megascolecidae, Ocnero-drilidae and Sparganophilidae). Megadrilog-ica 5(6): 53-72.

North Dakota

Reynolds, J.W. 1978b. A contribution to our know-ledge of the earthworm fauna of NorthDakota. Megadrilogica 3(8): 148-149.

Utter, R.A., E.J. Deibert, and D.P. Schwert. 1995a.Survey of North Dakota earthworms(Lumbricidae). Proceedings of the NorthDakota Academy of Science 49: 48.

Utter, R.A., E.J. Deibert and D.P. Schwert. 1995b.Earthworms of North Dakota. Jamestown,ND: Northern Prairie Wildlife Res. Cent.Home Page. World Wide Web URL:http://www.npwrc.usgs.gov/resource/inverts/wormsnd/ [version 24 August 2006; accessed16 November 2012]

Ohio

Olson, H.W. 1928. The earthworms of Ohio, with astudy of their distribution in relation tohydrogen-ion concentration, moisture andorganic matter content in the soil. Bull. OhioBiol. Surv. 4(2), Bull. 17: 47-90.

The Status of Terrestrial Earthworm Surveys that have been conducted in North America, etc.242

Reynolds, J.W. 2007e. The earthworms (Oligochaeta:Acanthodrilidae, Lumbricidae and Spargano-philidae) of Ohio, USA, revisited. Mega-drilogica 11(9): 91-105.

Oklahoma

Harman, W.J. 1954. On some earthworms fromsouthern Oklahoma. Proc. Oklahoma Acad.Sci. 35: 51-55.

Reynolds, J.W. and G.A. Damoff. 2010. The earth-worms (Oligochaeta: Acanthodrilidae,Lumbricidae, Megascolecidae and Spargano-philidae) of Oklahoma, USA. Megadrilogica13(12): 173-193.

Oregon

Macnab, J.A. and D. McKey-Fender. 1947. An intro-duction to Oregon earthworms with additionsto the Washington list. Northwest Sci. 21(2):69-75.

Pennsylvania

Reynolds, J.W. 2008b. The earthworms (Oligochaeta:Acanthodrilidae, Lumbricidae, Megascolec-idae and Sparganophilidae) of Pennsylvania,USA. Megadrilogica 11 (12): 131-146.

Puerto Rico

see Reynolds and Wetzel (2004, 2008, 2012).

Rhode Island

Reynolds, J.W. 1973b. The earthworms of RhodeIsland (Oligochaeta: Lumbricidae). Mega-drilogica 1(6): 1-4.

Reynolds, J.W. 2002a. Additional earthworm (Oligo-chaeta: Lumbricidae and Megascolecidae)records from Rhode Island, USA. Mega-drilogica 9(4): 21-27.

South Carolina

Reynolds, J.W. 2001a. The earthworms of SouthCarolina (Oligochaeta: Acanthodrilidae,Lumbricidae, Megascolecidae, Ocnerodrilidaeand Sparganophilidae). Megadrilogica 8(7):25-36.

Reynolds, J.W. and W.K. Reeves. 2004. Additionalearthworm records (Oligochaeta: Lumbric-idae, Megascolecidae and Sparganophilidae)from South Carolina. Megadrilogica 9(12):99-110.

South Dakota

Reynolds, J.W. 2007a. The earthworms (Oligochaeta:Lumbricidae) of South Dakota, USA.Megadrilogica 10(12): 95-105.

Texas

Damoff, G.A. and J.W. Reynolds. 2009. The earth-worms (Oligochaeta: Acanthodrilidae,Eudrilidae, Lumbricidae, Megascolecidae,Ocnerodrilidae, and Sparganophilidae) of eastTexas, USA. Megadrilogica 13(8): 113-140.

Tennessee

Damoff, G.A. and J.W. Reynolds, 2013. The earth-worms of Tennessee (Oligochaeta). V.Acanthodrilidae (in prep.)

Reynolds, J.W. 1974a. Checklist, distribution and keyto the Lumbricidae in Tennessee. J. Tennes-see Acad. Sci. 49(1): 16-20.

Reynolds, J.W. 1977d. The earthworms of Tennessee(Oligochaeta). II. Sparganophilidae, with thedescription of a new species. Megadrilogica3(3): 61-64.

Reynolds, J.W. 1977e. The earthworms of Tennessee(Oligochaeta). III. Komarekionidae, withnotes on distribution and biology. Mega-drilogica 3(4): 65-69.

Reynolds, J.W. 1978a. The earthworms of Tennessee(Oligochaeta). IV. Megascolecidae, withnotes on distribution, biology and a key to thespecies in the state. Megadrilogica 3(7):117-129.

Reynolds, J.W., E.E.C. Clebsch and W.M. Reynolds.1974. The earthworms of Tennessee (Oligo-chaeta). I. Lumbricidae. Bull. Tall TimbersRes. Stn., No. 17, vii + 133 pp.

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Utah

Reynolds, J.W. 2008i. The earthworms (Oligochaeta:Lumbricidae and Megascolecidae) of Utah,USA. Megadrilogica 12(7): 105-115.

Vermont

Reynolds, J.W. 2008d. The earthworms (Oligochaeta:Lumbricidae) of Vermont, USA. Megadril-ogica 12(2): 19-27.

Reynolds, J.W. 2012b. New earthworm (Oligochaeta:Megascolecidae) records from Vermont,USA. Megadrilogica 15(7): 187-190.

Virginia

Reynolds, J.W. 1994c. Earthworms of Virginia(Oligochaeta: Acanthodrilidae, Komarek-ionidae, Lumbricidae, Megascolecidae andSparganophilidae). Megadrilogica 5(8): 77-94.

Washington

Altman, L.A. 1936. Oligochaeta of Washington.Univ. Washington Publ. Biol. 4: 1-137.

Macnab, J.A. and D. McKey-Fender. 1947. An intro-duction to Oregon earthworms with additionsto the Washington list. Northwest Sci. 21(2):69-75.

West Virginia

Reynolds, J.W. 2007d. The earthworms (Oligochaeta:Acanthodrilidae, Lumbricidae and Mega-scolecidae) of West Virginia, USA. Mega-drilogica 11(7): 73-84.

Wisconsin - no formal earthworm survey, onlyincidental records (Reynolds and Wetzel,2004, 2008, 2012).

Wyoming

Reynolds, J.W. 2003a. The earthworms (Oligochaeta:Lumbricidae) of Wyoming, USA. Megadril-ogica 9(6): 33-39.

Reynolds, J.W., W.K. Reeves and R.M. Spence. 2009.The earthworms (Oligochaeta: Lumbricidae)of Wyoming, USA, revisited. Megadrilogica13(4): 25-35.

OTHER REFERENCES

Bhatti, H.K. 1965. Earthworms of Swarthmore, Penn-sylvania, and vicinity. Proc. PennsylvaniaAcad. Sci. 39: 8-24.

Crawford, R.L. and W.R. Brueckheimer. 2012. Thelegacy of a Red Hills Plantation: Tall TimbersResearch Station and Land Conservancy.Gainesville, FL: University Press of Florida,384 pp.

Das, B.C. and J.W. Reynolds. 2003. A checklist anddistribution of the freshwater and terrestrialAnnelida (Oligochaeta, Hirudinea and Poly-chaeta) of Bangladesh. Megadrilogica 9(10):61-70.

Damoff, G.A. 2011. Earthworms (Oligochaeta:Acanthodrilidae and Lumbricidae) fromNebraska, USA. Megadrilogica 14(8): 172-174.

Eaton, T.H., Jr. 1942. Earthworms from the NorthEastern United States. J. Washington Acad.Sci. 32(8): 242-249.

Fender, W.M. 1985. Earthworms of the westernUnited States. Part I. Lumbricidae. Mega-drilogica 4(5): 93-129.

Fragoso, C., S.W. James and S. Borges. 1995. Nativeearthworms of the north Neotropical Region:current status and controversies. Pp. 67-115,In: Hendrix, P.F. (ed.). Ecology and Bio-geography of Earthworms in North America.Boca Raton, FL: Lewis Publ., 244 pp.

Garman, H. 1888. On the anatomy and histology of anew earthworm (Diplocardia communis gen.et sp. nov.). Bull. Illinois St. Lab. Hist. 3: 47-77.

Gates, G.E. 1935. The earthworms of New England.Proc. New England Zool. Club, Cambridge15: 41-44.

The Status of Terrestrial Earthworm Surveys that have been conducted in North America, etc.244

Gates, G.E. 1942. Check list and bibliography ofNorth American earthworms. American Midl.Nat. 27(1): 86-108.

Gates, G.E. 1955. Notes on several species of theearthworm genus Diplocardia Garman, 1888.Bull. Mus. Comp. Zool., Harvard 113: 229-259.

Gates, G.E. 1967a. On two little known species of theearthworm genus Diplocardia. Great BasinNat. 27: 1-10.

Gates, G.E. 1967b. On the earthworm fauna of thegreat American desert and adjacent areas.Great Basin Nat. 27: 142-176.

Gates, G.E. 1972a. Burmese earthworms. An intro-duction to the systematics of megadrile oligo-chaetes with special references to southeastAsia. Trans. American Philos. Soc. 62(7): 1-326.

Gates, G.E. 1972b. Contributions to North AmericanEarthworms (Annelida: Oligochaeta) No. 3.Toward a revision of the family Lumbricidae.IV. The trapezoides species group. Bull. TallTimbers Res. Stn. No. 12: 1-146.

Gates, G.E. 1974. Contributions to a revision of thefamily Lumbricidae. XI. Eisenia rosea(Savigny, 1826). Bull. Tall Timbers Res. Stn.,No. 16: 9-30.

Gates, G.E. 1977. More on the earthworm genusDiplocardia. Megadrilogica 3(1): 1-48.

Gates, G.E. 1979. Contributions to a revision of theearthworm family Lumbricidae. XXIII. Thegenus Dendrodrilus Omodeo, 1956, in NorthAmerica. Megadrilogica 3(9): 151-162.

Görres, J.H. and R.D.S. Melnichuk. 2012. Asianinvasive earthworms of genus AmynthasKinberg in Vermont. Northeastern Nat. 19(2):313-322.

Greiner, H.G., A.M.T. Stonehouse and S.D. Tiegs.2011. Cold tolerance among compostingearthworm species to evaluate invasionpotential. American Midl. Nat. 166: 349-357.

Hong, Y. and J.W. Reynolds. 1997. Brief historicalreview and checklist of Korean earthworms.Megadrilogica 6(11): 103-104.

James, S.W. 1984. New records of earthworms fromKansas (Oligochaeta: Acanthodrilidae,Lumbricidae, Megascolecidae). The PrairieNaturalist 16: 91-95.

James, S.W. 1988. Diplocardia hulberti and D.rugosa, new earthworms (Annelida: Oligo-chaeta: Megascolecidae) from Kansas. Proc.Biol. Soc. Washington 101(2): 300-307.

Kathireswari, P., J.M. Julka and J.W. Reynolds. 2005.Checklist of Oligochaeta of Tamil Nadu,India. Megdrilogica 10(8): 57-68.

McAlpine, D.F. and J.W. Reynolds. 1977. TerrestrialOligochaeta of some New Brunswick caveswith remarks on their ecology. Can. Field-Nat. 91(4): 360-366.

McAlpine, D.F., J.W. Reynolds, T.J. Fletcher, J.L.Trecartin and D.L. Sabine. 2001. Spargano-philus eiseni (Oligochaeta, Sparganophilidae)in New Brunswick, an addition to the earth-worm fauna of Maritime Canada.Megadrilogica 8(10): 53-56.

Moore, J.-D. and J.W. Reynolds. 2003. Premièrsmentions de vers de terre (Oligochaeta:Lumbricidae) dans deux écosystèmesforestiers du bouclier canadien, Québec,Canada. Megadrilogica 9(9): 53-60.

Moore, J.-D., R. Ouimet and J.W. Reynolds. 2009.Premièrs mentions de vers de terre dans troisécosystèmes forestiers du Bouclier canadien,Québec, Canada. Nat. can. 133(1): 31-37.

Murchie, W.R. 1961. A new diplocardian fromIllinois (Oligochaeta: Megascolecidae). OhioJ. Sci. 61(6): 367-371.

Murchie, W.R. 1962. A new subspecies of Diplo-cardian earthworm from Oklahoma (Oligo-chaeta: Megascolecidae). Ohio J. Sci. 62(4):185-190.

Murchie, W.R. 1963. Description of a new diplo-cardian earthworm, Diplocardia longiseta(Oligochaeta: Megascolecidae). Ohio J. Sci.63(1): 15-18.

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Reeves, W.K. and J.W. Reynolds. 2010. The firstearthworm (Oligochaeta: Megascolecidae)record from the Republic of the MarshallIslands. Megadrilogica 14(5): 75-76.

Reynolds, J.W. 1975f. Sparganophilus pearsei n. sp.(Oligochaeta: Sparganophilidae) a nearcticearthworm from western North Carolina.Megadrilogica 2(2): 9-11.

Reynolds, J.W. 1976. Un aperçu vers de terre dans lesforêts nord-américanes, leurs activitiés et leursrépartition. Megadrilogica 2(9): 1-11.

Reynolds, J.W. 1980. The earthworm family Spar-ganophilidae (Annelida, Oligochaeta) in NorthAmerica. Megadrilogica 3(12): 189-204.

Reynolds, J.W. 1993. On some earthworms fromSwaziland (Oligochaeta: Glossoscolecidae,Megascolecidae, Microchaetidae and Octo-chaetidae). Megadrilogica 5(1):1-8.

Reynolds, J.W. 1994f. The distribution of earthworms(Annelida, Oligochaeta) in North America.Pp. 133-153. In: Mishra, P.C., N. Behera,B.K. Senapati and B.C. Guru (eds.), Advancesin Ecologyand Environmental Sciences. NewDelhi: Ashish Publishing House, 651 pp.

Reynolds, J.W. 1994g. Earthworms of the world.Global Biodiversity 4(1): 11-16.

Reynolds, J.W. 1994c. The earthworms of Bangla-desh (Oligochaeta: Megascolecidae, Monili-gastridae and Octochaetidae). Megadrilogica5(4): 33-44.

Reynolds, J.W. 1995. The status of exotic earthwormsystematics and biogeography in NorthAmerica. Pp. 1-27 In Hendrix, P.F. (ed.),Ecology and Biogeography of Earthworms inNorth America. Boca Raton, FL: Lewis Publ.,244 pp.

Reynolds, J.W. 1996b. On some cave earthworms(Oligochaeta: Lumbricidae) from the Isle ofMan, U.K. Megadrilogica 6(9): 89-90.

Reynolds, J.W. 1998. The status of earthwormbiogeography, diversity and taxonomy inNorth America revisited with glimpses into

the future. Pp. 15-34 In: Edwards, C.A. (ed.),Earthworm Ecology. Delray Beach, FL: St.Lucie Press, 389 pp.

Reynolds, J.W. 1999a. New earthworm records forthe state of California, USA. Megadrilogica7(8): 54.

Reynolds, J.W. 1999c. A contribution to our know-ledge of the earthworm fauna of Hawaii,USA. Megadrilogica 7(11): 79-80.

Reynolds, J.W. 2004b. The status of earthwormbiogeography, diversity, and taxonomy inNorth America revisited with glimpses intothe future. Pp. 63-74. In: EarthwormEcology, 2nd edition. Edwards, C.A. (ed.). St.Lucia, FL: CRC Press, 441 pp.

Reynolds, J.W. 2008j. Sparganophilidae (Annelida,Oligochaeta) distribution in North Americaand elsewhere, revisited. Megadrilogica12(9):105-120.

Reynolds, J.W. 2010b. The earthworms (Oligochaeta:Acanthodrilidae, Lumbricidae, Megascolec-idae and Sparganophilidae) of northeasternUnited States, revisited. Megadrilogica 14(7):101-157.

Reynolds, J.W. 2010c. Earthworms (Oligochaeta:Lumbricidae, Sparganophilidae) of theAtlantic Maritime Ecozone. Chapter 13, Pp.225-281. In: Assessment of Species Diversityin the Atlantic Maritime Ecozone, D.F.McAlpine and I. Smith (eds.). Ottawa: NRCPress, xii + 785 pp.

Reynolds, J.W. 2011a. The earthworms (Oligochaeta:Acanthodrilidae, Eudrilidae, Glossoscolec-idae, Komarekionidae, Lumbricidae, Luto-drilidae, Ocnerodrilidae, Octochaetidae,Megascolecidae and Sparganophilidae) ofsoutheastern United States. Megadrilogica14(9-12): 175-318.

Reynolds, J.W. 2011d. The earthworms (Oligochaeta:Acanthodrilidae, Glossoscolecidae, Komarek-ionidae, Lumbricidae, Megascolecidae andSparganophilidae) of the midwestern UnitedStates. Megadrilogica 15(5): 69-139.

The Status of Terrestrial Earthworm Surveys that have been conducted in North America, etc.246

Reynolds, J.W. 2012a. The earthworms (Oligochaeta:Lumbricidae, Megascolecidae and Spargano-philidae) of the North-Central United States.Megadrilogica 15(7): 151-187.

Reynolds, J.W. and D.S. Christie. 1977. Additionalrecords of New Brunswick earthworms. NewBrunswick Nat. 8(3): 25.

Reynolds, J.W. and D.G. Cook. 1976. NomenclaturaOligochaetologica: a catalogue of the names,descriptions and type specimens of theOligochaeta. Fredericton: University of NewBrunswick, x + 217 pp.

Reynolds, J.W. and D.G. Cook. 1981. NomenclaturaOligochaetologica Supplementum Primum: acatalogue of names, descriptions and typespecimens of the Oligochaeta. Fredericton:Centennial Print and Litho Ltd., v + 39 pp.

Reynolds, J.W. and D.G. Cook. 1989. NomenclaturaOligochaetologica Supplementum Secundum:a catalogue of the names, descriptions andtype specimens of the Oligochaeta. NewBrunswick Museum, Monographic Series(Natural Science) No. 8, v + 37 pp.

Reynolds, J. W. and D. G. Cook. 1993. NomenclaturaOligochaetologica Supplementum Tertium: acatalogue of the names, descriptions and typespecimens of the Oligochaeta. New Bruns-wick Museum Monographic Series (NaturalScience) No. 9, vi + 33 pp.

Reynolds, J.W. and C. Hänel. 2005. The earthworms(Oligochaeta: Lumbricidae) of Tristan daCunha and Nightingale Islands, South AtlanticOcean. Megadrilogica 10(7): 47-56.

Reynolds, J.W. and A.G. Jones. 2005. The earth-worms (Oligochaeta: Acanthodrilidae,Glossoscolecidae, and Lumbricidae) of theFalkland Islands, South Atlantic Ocean.Megadrilogica 10(10): 75-86.

Reynolds, J.W. and A.J. Reinecke. 1977. A prelim-inary survey of the earthworms of the KrugerNational Park, South Africa (Oligochaeta:Glossoscolecidae, Megascolecidae and Octo-chaetidae). Wet. Bydraes, P.U. vir C.H.O.(B), No. 89, 19 pp.

Reynolds, J.W. and D.W. Reynolds. 2001b. Primerosdatos de lombrices de tierra (Oligochaeta) dela Isla de San Andrés, Colombia. Mega-drilogica 8(6): 21-23.

Reynolds, J.W. and D.W. Reynolds. 2002. Primerosdatos de lombrices de tierra (Oligochaeta) della Isla de Contadora, Panama. Megadrilogica9(1): 1-4.

Reynolds, J.W. and D.W. Reynolds. 2006. Sobrealgunas lombrices de tierra (Oligochaeta) deHonduras, C.A. Megadrilogica 10(11): 87-91.

Reynolds, J.W. and D.W. Reynolds. 2010. The GreatGlobal Earthworm Race goes to Dubai,U.A.E. Megadrilogica 14(2): 49-52.

Reynolds, J.W. and M.J. Wetzel. 2004. TerrestrialOligochaeta (Annelida: Clitellata) in NorthAmerica north of Mexico. Megadrilogica9(11): 71-98.

Reynolds, J.W. and M.J. Wetzel. 2008. Oligochaeta(Annelida: Clitellata) in North America,including Mexico, Puerto Rico, Hawaii, andBermuda. Megadrilogica 12(12): 157-208.

Reynolds, J.W. and M.J. Wetzel. 2012. TerrestrialOligochaeta (Annelida: Clitellata) in NorthAmerica, including Mexico, Puerto Rico,Hawaii, and Bermuda. III. Megadrilogica15(8): 191-211.

Reynolds, J.W. and. M.J. Wetzel. 2013. NomenclaturaOligochaetologica, 2nd ed. (in prep.)

Reynolds, J.W., J.M. Julka and M.N. Khan. 1995.Additional earthworm records fromBangladesh (Oligochaeta: Glossoscolecidae,Megascolecidae, Moniligastridae, Ocnero-drilidae and Octochaetidae). Megadrilogica6(6): 51-60.

Reynolds, J.W., A.G. Jones, K.J. Gaston and S.L.Chown. 2002. Earthworms (Oligochaeta:Lumbricidae) of Gough Island, SouthAtlantic. Megadrilogica 9(2): 5-15.

Rodriguez, A.R., R.S. Ostfeld, F. Keesing and J.W.Reynolds. 2012. The earthworms (Oligo-chaeta: Lumbricidae and Megascolecidae) ofDutchess County, New York, USA. Mega-drilogica 15(6): 141-150.

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Siddaraju, M., K.S. Sreepada and J.W. Reynolds.2010. Checklist of earthworms (Annelida:Oligochaeta) from Dakshina Kannada,Karnataka South West India. Megadrilogica14(5): 65-75.

Smith, F. 1895a. A preliminary account of two newOligochaeta from Illinois. Bull. Illinois St.Lab. Nat. Hist. 4(5): 138-148.

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

Smith, F. 1896. Notes on species of North AmericanOligochaeta II. Bull. Illinois St. Lab. Nat.Hist. 4(14): 396-413.

Smith, F. 1897. Upon an undescribed species ofMegascolides from the United States. Ameri-can Midl. Nat. 31: 202-204.

Smith, F. 1900. Notes on species of North AmericanOligochaeta. III. List of species found inIllinois, and descriptions of IllinoisTubificidae. Bull. Illinois St. Lab. Nat. Hist.5(10): 441-458.

Smith, F. 1912. Earthworms from Illinois. Trans.Illinois Acad. Sci. 5: 145-148.

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

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

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

The Status of Terrestrial Earthworm Surveys that have been conducted in North America, etc.248

APPENDIXBangor Daily News, Monday, June 27, 1977 17

A dream come true

Dr. Gordon E. Gates of Bangor(left) long ago saw the need for acatalog of all known scientific namesfor Oligochaeta, or worms, to aidresearchers. He started the project butwas unable to complete it because ofhis age. So in 1972, John W.

Reynolds, a colleague of Gates andassistant professor at the University ofNew Brunswick took over the project.The book now has been published; andReynolds was in Bangor this week toshow Gates the finished product.(NEWS Photo by Carroll Hall)

MEGADRILOGICA 249

APPENDIX (Continued)

Bangor resident’s workhas come to fruition

By Nancy RemsenOf the NEWS Staff

A Bangor scientist who spent yearsworking on a catalog of the scientificnames of Oligochaeta, or worms, buthad to give up the project because ofhis age, saw his dream come true thisweek.

The research guide which he beganwas completed by a colleague, John W.Reynolds, a professor at the Universityof New Brunswick, and David G.Cook, an English scientist who nowlives in Ottawa. Reynolds was inBangor this week to show Gates thecompleted book.

The idea of the book is unusual,Reynolds said, because it lists thescientific names of all known worms asof December 31at, 1975, who namedthe worm, in what journal the name anddescription were first published, and inwhat institution the specimen used for

naming and describing is now located.

Such data is valuable to researcherswho might be trying to verify theidentity of a specimen they have or tosomeone who might be reclassifying agroup of worms, Reynolds said.

And anyone who thinks he or she hasa new type of worm would have tomake sure the name proposed hasn’talready been given to another type ofworm, he said.

Also unusual about the book is thatthe introduction and explanation ofhow to use the book was written in sixlanguages, English, French, German,Portuguese, Russian and Chinese.“Anyonein the world can use thebook,” Reynolds said, sinceinternational scientists speak one ofthose languages in order to read otherscientists’ research.

Web Site for the Journal Megadrilogica:http://www.inhs.uiuc.edu/~mjwetzel/Megadrilogica.home.html