Abstract
A first comprehensive checklist of the Ocnerodrilidae in South America is provided. It includes 70 nominal taxa (species and subspecies), 31 of which were described as new by Gilberto Righi. Of this fauna, 86% is endemic to South America. Sixteen species are endemic in subtropical South America, 34 are endemic in tropical South America, and 10 are endemic in both tropical and subtropical South America. Ten species extend their distributions into Central America, North America, Africa, Europe, Asia, and/or Australia. Only two of these, Eukerria saltensis and Ocnerodrilus occidentalis, have become virtually pantropical, reaching also into the temperate region of the northern hemisphere. An updated distribution of each taxon is also given, with references for each locality. Full synonyms and detailed South American occurrences are provided.
Introduction
Ocnerodrilids are small worms, usually less than 40×1 mm, and mostly associated with freshwater. The Ocnerodrilidae form a homogeneous taxon (Jamieson Citation1971, Citation1974, Sims Citation1980, Sielfeld Citation2002).
Ocnerodrilid earthworms, according to the classical phylogeny, constitute the most recent offshot from the ancestral root‐genus of all megascolecoid forms. However, both digestive and vascular systems are, in several respects, less evolved than in the other megascolecoids (Gates Citation1979, p. 162). Sims (Citation1980) maintains them as a family of Megascolecoidea, while Reynolds and Cook (Citation1993) prefer to put them in their own superfamily Ocnerodriloidea. Jamieson (Citation1971, Citation1974), followed by Omodeo (Citation2000), considered them to represent the most primitive family among the Megascolecoidea. Recent molecular studies support this last point of view (Jamieson et al. Citation2002, Pop et al. Citation2005).
Ocnerodrilinae live in intertropical America and Africa, while Malabarinae are restricted to India. Paulistus has been considered an incertae sedis taxon within this family (Omodeo Citation2000, p. 179). Although an African origin of the group is assumed by many specialists, these worms have colonized so widely throughout the tropics that the original homes of some groups are unknown (Gates Citation1979, p. 162).
In this paper I provide an inventory of the ocnerodrilid megadriles cited to date in South America.
Abbreviations
I have used three‐letter abbreviations for South American countries (ARG, Argentina; BOL, Bolivia; BRA, Brazil; CHI, Chile; COL, Colombia; ECU, Ecuador; GUY, Guyana; PAR, Paraguay; PER, Peru; URU, Uruguay; VEN, Venezuela) and two‐letter abbreviations for sampled States in Brazil (AM, Amazonas; AP, Amapá; BA, Bahia; ES, Espírito Santo; GO, Goiás; MG, Minas Gerais; MS, Mato Grosso do Sul; MT, Mato Grosso; PA, Pará; PB, Paraíba; PE, Pernambuco; PR, Paraná; RJ, Rio de Janeiro; RO, Rondônia; RR, Roraima; RS, Rio Grande do Sul; SC, Santa Catarina; SE, Sergipe; SP, São Paulo).
Taxonomic accounts
Species marked with an asterisk (*) are only reported from South America. Geographical distributions are summarized in Tables – . Further known localities of species within each state in Brazil are given in the Appendix I.
Table I. Distribution of subtropical species of Ocnerodrilidae in South America (references for BRA include only the subtropical states of RS, SC, PR, and SP. Abbreviation: NEPac, North East Pacific).
Table II. Distribution of subtropical–tropical species of Ocnerodrilidae in South America (references for BRA are divided into South BRA, containing the states of RS, SC, PR and SP, and North BRA, including the remaining states north of SP. Abbreviations: Afr, Africa; Asi, Asia; CAm, Central America; pantr., pantropical; temp., temperate).
Table III. Distribution of tropical species of Ocnerodrilidae in South America (references for BRA include only the tropical states north of SP. Abbreviations: Afr, Africa; Asi, Asia; CAm, Central America; EAtl, East Atlantic).
LUMBRICINA De Blainville, 1830
Syn: Terricolae Örsted, 1850; Megadrili Benham, 1890; Diplotesticulata Yamaguchi, 1953; Crassiclitellata Jamieson, 1988
MEGASCOLECOIDEA Rosa, 1891
OCNERODRILIDAE Beddard, Citation1891
OCNERODRILINAE Beddard, Citation1891
Bauba Righi, 1980
Type: B. santosi Righi, 1980
1*. B. santosi Righi, 1980
Distr: BRA: SE: 1.
Belladrilus Righi, 1984
Type: B. jimi Righi, 1984
Belladrilus (Belladrilus) Righi, 1984
Type: B. jimi Righi, 1984
2*. B. (B.) arua Righi, 1984
Distr: BRA: MT: 2.
3*. B. (B.) jimi Righi, 1984
Distr: ARG: Buenos Aires Prov: Buenos Aires (Burela & Cazzaniga Citation2001, p. 49); Córdoba: Sierras Chicas (De Mischis Citation1999, p. 19); Corrientes. BRA: MS: 1; MT: 1.
4*. B. (B.) otarion Righi, Citation1995
Distr: BRA: SP: 1, 2.
5*. B. (B.) pocaju Righi, 1984
Distr: BRA: MS: 2; MT: 3.
6*. B. (B.) vaucheri Zicsi, Citation1995
Distr: BOL: Gran Chaco Prov: Entre Ríos; Tarija Dep. (Zicsi Citation1995a, p. 604).
Belladrilus (Santomesia) Righi, 1984
Type: Belladrilus emilianii Righi, 1984
7*. B. (Santomesia) auka Righi & De Mischis, Citation1999
Distr: ARG: Córdoba Prov: Parque Siquiman Dep. (Righi & De Mischis Citation1999, p. 78).
8*. B. (Santomesia) emilianii Righi, 1984
Distr: ARG (Righi Citation1984a, p. 20).
Brunodrilus Righi, 1971
Type: B. angeloi Righi, 1971
9*. B. angeloi Righi, 1971
Distr: BRA: MG: 1.
Dariodrilus Righi, Ayres & Bittencourt, Citation1978
Type: D. ferrarius Righi, Ayres & Bittencourt, Citation1978
10*. D. ferrarius Righi, Ayres & Bittencourt, Citation1978
Distr: BRA: AM: 1.
Eukerria Michaelsen, Citation1935 (Pro: Kerria)
Syn: Kerria Beddard, Citation1892 (Type: Kerria halophila Beddard, Citation1892)
11.* E. asilis (Righi, 1968)
Syn: Kerria asilis Righi, 1968.
Distr: BRA: PE: 1; AP: 1.
12*. E. asuncionis (Rosa, Citation1895)
Syn: Kerria asuncionis Rosa, Citation1895.
Distr: ARG: Santa Fé Prov. (Ljungström et al. Citation1973, p. 240). PAR: Asunción (Rosa Citation1895, p. 2; Rosa Citation1896, p. 89). BOL: Vaca Diez Prov: Beni Dep: Guayaremerín: Mamoré river (Zicsi Citation1995a, p. 604).
13*. E. cuca (Righi, 1984)
Syn: Kerria cuca Righi, 1984
Distr: BRA: MT: 4.
14*. E. eiseniana (Rosa, Citation1895)
Syn: Kerria eiseniana Rosa, Citation1895; Kerria hortensis Stephenson, Citation1931; E. hortensis (Stephenson, Citation1931).
Distr: ARG: Northwest Provs (Gavrilov Citation1979: p. 109); Jujuy Prov: San Lorenzo (Cognetti Citation1902, p. 3); Córdoba: Sierras Chicas (De Mischis Citation1999, p. 19); Santa Fé Prov: La Capital, Santo Tomé and Garay (Ljungström Citation1971, p. 39; Ljungström & Emiliani Citation1971, p. 21; Emiliani et al. Citation1973, p. 50); Corrientes Prov: Bella Vista (Gavrilov Citation1979, p. 109). PAR: Central Dep: Asunción; Concepción Dep: Apa river; Arnamby Dep: Bella Vista (Rosa Citation1895, p. 2; Rosa Citation1896, p. 89); Presidente Hayes: Makthalawaiya (Stephenson Citation1931, p. 314). BOL: Chiquitos Prov: Santa Cruz Dep: Puerto Suarez (Righi Citation1990, p. 64); Vaca Diez Prov: Beni Dep: Guayaramerín (Zicsi Citation1995a, p. 603). VEN: Bolívar (Lavelle & Lapied Citation2003, p. 424). BRA: RS: 1; PR: 1–3; SP: 1, 3; MS: 3; MT: 5–8; RO: 1, 2; RR: 1, 2.
15*. E. emete Righi & Guerra, Citation1985
Distr: BRA: PR: 1, 2; SP: 1; MT: 9–12.
16*. E. garmani argentinae Jamieson, Citation1970.
Distr: ARG: Misiones (Jamieson Citation1970, p. 133). BOL: Vaca Diez Prov: Beni Dep: Guayaramerín (Zicsi Citation1995a, p. 605). BRA: RS: 1, 2.
17*. E. garmani garmani (Rosa, Citation1895)
Syn: Kerria garmani Rosa, Citation1895.
Distr: PAR: Central Dep. (Rosa Citation1895, p. 1; Rosa Citation1896, p. 89). BOL: Vaca Diez Prov: Beni Prov: Guayaramerín (Zicsi Citation1995a, p. 603).
18*. E. guamais (Righi, 1971)
Syn: Kerria guamais Righi, 1971
Distr: BRA: PA: 1, 2; AM: 1, 2; RO: 3.
19*. E. halophila (Beddard, Citation1892)
Syn: Kerria halophila Beddard, Citation1892.
Distr: CHI (Righi Citation1990, p. 64). ARG: Santa Fé Prov. (Ljungström & Emiliani Citation1971, p. 21; Emiliani et al. Citation1971, p. 19; Emiliani et al. Citation1973, p. 50; Ljungström et al. Citation1973, p. 236; Ljungström et al. Citation1975, p. 12). BOL: Gran Chaco Prov: Tarija Dep: tributaries of Pilcomayo river (Beddard Citation1892, p. 355).
20. E. kuekenthali (Michaelsen, Citation1908)
Syn: Kerria kükenthali Michaelsen, Citation1908; E. selangorensis Stephenson, Citation1931; E. peguana Gates, Citation1942.
Distr: BRA: AM: 3.
Furth. Distr: West Indies (Michaelsen Citation1908, p. 24), Asia (Gates Citation1942, p. 67), and Christmas Island (Michaelsen Citation1935, p. 100).
21*. E. limosa (Stephenson, Citation1931)
Syn: Kerria limosa Stephenson, Citation1931.
Distr: PAR: Makthalawaiya (Stephenson Citation1931, p. 312).
22. E. mcdonaldi (Eisén, 1893)
Syn: Kerria mcdonaldi Eisén, 1893.
Distr: ARG: Córdoba Prov: Cosquin (Cognetti Citation1901, p. 2).
Furth. Distr: West Coast of North America (Michaelsen Citation1900a, p. 372; Gavrilov Citation1979, p. 109).
23*. E. mucu Righi, Citation1988
Distr: BRA: RO: 4.
24*. E. papillifera (Rosa, 1885)
Syn: Kerria papillifera Rosa, 1885.
Distr: PAR: Central region (Rosa Citation1895, p. 1; Rosa Citation1896, p. 89).
25*. E. pascuorum (Stephenson, Citation1931)
Syn: Kerria pascuorum Stephenson, Citation1931.
Distr: PAR: Makthalawaiya (Stephenson Citation1931, p. 316).
26*. E. rosae (Beddard, Citation1895)
Syn: Kerria rosae Beddard, Citation1895
Distr: ARG: Buenos Aires (Beddard Citation1895, p. 210; Michaelsen Citation1900b, p. 25).
27. E. saltensis (Beddard, Citation1895)
Syn: Kerria saltensis Beddard, Citation1895; Acanthodrilus sydneyensis Sweet, 1900; Kerria gunningi Michaelsen, 1913; Kerria nichollsi Jackson, 1931; E. saltensis var. bellavistensis Gavrilov, Citation1952.
Distr: CHI: Juan Fernandez, Coquimbo, Valparaiso, Quillta (Michaelsen Citation1898, p. 471), and El Salto (Beddard Citation1895, p. 225). ARG: Aranco and Sanagasta Dep: La Rioja Prov: Velasco and Famatina mountain range; La Rioja (De Mischis & Gleiser Citation1999, p. 61); Córdoba: Sierras Chicas (De Mischis Citation1999, p. 19); Catamarca Prov: Lãs Pirquitas Ville: Del Valle river (De Mischis & Righi Citation1999, p. 63); San Luis Prov. (De Mischis & Brigada Citation1985, p. 134); Tucumán Prov. (Gavrilov Citation1952, p. 673); Santa Fé Prov; Córdoba Prov. (Ljungström et al. Citation1973, p. 240; Ljungström et al. Citation1975, p. 12). BOL: Larecoche Prov: Between Teoponte and Alcoche; Nor Yungas Prov: La Paz Dep; Tarija Prov. (Zicsi Citation1995a, p. 604). BRA: 1; RS: 1, 3; SC: 1; PR: 2, 3; SP: 1, 4; MG: 2.
Furth. Distr: Central America (Römbke & Zicsi Citation2007), North America (Righi Citation1979, p. 121; Rodrigues Aragones Citation1999, p. 155), Europe (Rota Citation2004), Africa, Asia, including Japan (Blakemore et al. Citation2006, p. 173), and Australia (Blakemore Citation1999, p. 182).
Rem: Brown et al. (Citation2006, p. 358) consider that this widespread species may have originated in the Argentina–Paraguay region.
28*. E. santafesina Ljungström, Citation1972
Distr: ARG: Santa Fé Prov. (Ljungström Citation1972, p. 217).
29*. E. stagnalis (Kinberg, Citation1867)
Syn: Mandane stagnalis Kinberg, Citation1867; Kerria stagnalis (Kinberg, Citation1867); Acanthodrilus spegazzinii Rosa, Citation1890; Kerria spegazzinii (Rosa, Citation1890); E. spegazzinii (Rosa, Citation1890).
Distr: ARG: Buenos Aires (Rosa Citation1890, p. 516); Santa Fé Prov; Corrientes Prov. (Ljungström et al. Citation1973, p. 240; Ljungström et al. Citation1975, p. 13). URU: Montevideo (Kinberg Citation1867, p. 100): Prado: Miguelete stream; Rocha, Los Adobes, near Castillos lagune (Cordero Citation1931, p. 352). PAR: North region (Jamieson Citation1970, p. 133). BRA: 2: RS: 1, 2, 4; SP: 1, 5.
30*. E. subandina (Rosa, Citation1895)
Syn: Kerria subandina Rosa, Citation1895; Kerria borellii Cognetti, Citation1900; E. borellii (Cognetti, Citation1900).
Distr: ARG: Santa Fé Prov (Ljungström et al. Citation1975, p. 14); Salta Prov. (Rosa Citation1895, p. 2); Jujuy Prov: San Lorenzo (Cognetti Citation1902, p. 3); Tucumán Prov. (Gavrilov Citation1967a, p. 139): Tucumán (Gavrilov Citation1967b, p. 255). PAR (Römbke & Zicsi Citation2007). BOL: Quijarro Prov: Potosí Dep: Caiza; Gran Chaco Prov: Aguyrenda; Tarija Prov. (Cognetti Citation1902, p. 3). BRA: MT: 13, 14; RO: 5.
31*. E. taisa Righi, Citation1983
Distr: BRA: PA: 3.
32*. E. tucumana Cordero, Citation1942
Distr: ARG: Tucumán (Cordero Citation1942, p. 269).
33*. E. urna Righi, 1968
Distr: VEN: Bolivar (Lavelle & Lapied Citation2003, p. 424). BRA: BA: 1; PA: 4, 5; RO: 2; RR: 3; AM: 3.
34*. E. weyenberghi Cordero, Citation1942
Distr: ARG: Buenos Aires Prov: Oslas de Tigre (Cordero Citation1942, p. 269).
Exisdrilus Righi, Ayres & Bittencourt, Citation1978
Type: E. rarus Righi, Ayres & Bittencourt, Citation1978
35*. E. rarus Righi, Ayres & Bittencourt, Citation1978
Distr: BRA: AM: 4.
Gordiodrilus Beddard, Citation1892
Type: G. elegans Beddard, Citation1892
36. G. habessinus Michaelsen, 1913
Distr: BRA: SP: 1; MS: 3; MT: 6, 7, 12; RO: 2, 7, 8.
Furth. Distr: Africa (Righi Citation1990, p. 74).
37*. G. marcusi Righi, 1968
Distr: BRA: SP: 1, 4.
38. G. paski Stephenson, 1928
Syn: G. unicus Stephenson, 1928; G. peguanus Stephenson, 1928.
Distr: BRA: SP: 1, 4, 6, 7; GO: 1.
Furth. Distr: Central America, Africa, and Asia (Gavrilov Citation1979, p. 110).
Haplodrilus Eisén, Citation1900
Type: Ocnerodrilus borellii Rosa, Citation1895
Syn: Gatesia Jamieson, Citation1962 (Type: G. unica Jamieson, Citation1962)
Haplodrilus (Gatesia) Jamieson, Citation1962
39*. H. (G.) unica (Jamieson, Citation1962)
Syn: Gatesia unica Jamieson, Citation1962
Distr: ARG: Misiones Prov: Loreto (Jamieson Citation1962, p. 619).
Haplodrilus (Haplodrilus) Eisén, Citation1900
40*. H. (H.) amazonicus Righi, Citation1983
Distr: BRA: PA: 6.
41*. H. (H.) borellii (Rosa, Citation1895)
Syn: Ocnerodrilus borellii Rosa, Citation1895
Distr: PAR: Asunción (Rosa Citation1895, p. 3; Rosa Citation1896, p. 89).
42*. H. (H.) iheringi Michaelsen, Citation1926
Distr: BRA: SP: 1, 8.
43*. H. (H.) michaelseni (Cognetti, Citation1900)
Syn: Ocnerodrilus michaelseni Cognetti, Citation1900
Distr: PAR: Asunción (Cognetti Citation1900, p. 1). BRA: PR: 1, 2; MT: 13.
44*. H. (H.) tagua Righi, Ayres & Bittencourt, Citation1978
Distr: BRA: AM: 1.
Ilyogenia Beddard, 1893
Type: I. africana Beddard, 1893
Syn: Ocnerodrilus (Ilyogenia) Beddard, 1893
45*. I. comondui (Eisén, Citation1900)
Syn: Ocnerodrilus comondui Eisén, Citation1900; Enicomodrilus comondui (Eisén, Citation1900).
Distr: ARG (Fragoso & Brown Citation2007).
46*. I. paraguayense (Rosa, Citation1895)
Syn: Ocnerodrilus (Ilyogenia) paraguayensis Rosa, Citation1895.
Distr: ARG: Jujuy Prov: San Lorenzo (Cognetti Citation1902, p. 1). PAR: Central Prov: Asunción (Rosa Citation1895, p. 1; Rosa Citation1896, p. 89). BOL: Gran Chaco Prov: Santa Cruz Dep: Aguayrenda; Tarija Prov. (Cognetti Citation1902, p. 1).
47. I. tuberculatus Eisén, Citation1900
Distr: BOL: Vaca Diez Prov: Beni Dep: Guayaramerín: Nueva Cuba (Zicsi Citation1995a, p. 604).
Furth. Distr: Guatemala (Römbke & Hanagarth Citation1994, p. 14).
Kerriona Michaelsen, Citation1924
Type: K. luederwaldti Michaelsen, Citation1924
48*. K. garbei Michaelsen, Citation1924
Distr: BRA: ES: 1.
49*. K. limae Righi, 1980
Distr: BRA: SP: 1, 9.
50*. K. luederwaldti Michaelsen, Citation1924
Distr: BRA: RJ: 1.
Liodrilus Eisén, Citation1900 (Pro: Leidrilus Eisén, Citation1900)
Type: Ocnerodrilus eiseni Beddard, Citation1891
Syn: Ocnerodrilus (Leidrilus) Eisén, Citation1900; Ocnerodrilus (Guamaia) Righi, Citation1975
51*. L. eiseni (Beddard, Citation1891)
Syn: Ocnerodrilus eiseni Beddard, Citation1891
Distr: GUY (Beddard Citation1891, p. 563).
52*. L. ipu (Righi, Citation1975)
Syn: Ocnerodrilus ipu Righi, Citation1975; Ocnerodrilus (Guamaia) ipu Righi, Citation1975.
Distr: BRA: PA: 7.
53*. L. mendesi Righi, Citation1994
Distr: BRA: PB: 1.
Lourdesia Righi, Citation1994
Type: L. paraibaensis Righi, Citation1994
54*. L. paraibaensis Righi, Citation1994
Distr: BRA: PB: 1.
Nematogenia Eisén, Citation1900
Type: Pygmaeodrilus lacuum Beddard, 1893
Syn: Ocnerodrilus (Nematogenia) Eisén, Citation1900
55. N. lacuum (Beddard, 1893)
Syn: Pygmaeodrilus lacuum Beddard, 1893; Ocnerodrilus lacuum (Beddard, 1893); Ocnerodrilus (Nematogenia) lacuum (Beddard, 1893); N. josephinae Cognetti, 1904.
Distr: BRA: MT: 2, 6, 11, 15, 16, 17; RO: 9–11.
Furth. Distr: Central America and Africa (Righi Citation1990, p. 72).
Rem: Blakemore (Citation2002, p. 86) suggests that this is possibly a senior synonym of N. panamaensis. In this case, N. lacuum would also occur in Asia (Gates Citation1972, p. 272).
56. N. panamaensis (Eisén, Citation1900).
Syn: Ocnerodrilus (Nematogenia) lacuum var. panamaensis Eisén, Citation1900; Pygmaeodrilus lacuum var. panamaensis (Eisén, Citation1900).
Distr: VEN: In earth with anthurium roots arriving by plane at San Juan, Puerto Rico (Gates Citation1982, p. 67); San Juan (Gates Citation1979, p. 163). BRA: SP: 1, 3.
Futh. Distr: Central America, Africa (Gates Citation1982, p. 67), and Asia (Gates Citation1972, p. 272).
Ocnerodrilus Eisén, 1878
Type: O. occidentalis Eisén, 1898
57*. O. andinus Righi, Citation1981
Distr: ECU: Pichincha Dep: Los Tayos: cave near Quito (Righi Citation1981, p. 242).
58*. O. hendriei paulistus Righi, 1968
Syn: O. (Ilyogenia) hendriei ssp. paulistus Righi, 1968
Distr: BRA: SP: 4.
59*. O. ibemi Righi, 1968
Syn: O. (Ilyogenia) ibemi Righi, 1968; Ilyogenia ibemi (Righi, 1968).
Distr: BRA: SP: 10.
60. O. occidentalis Eisén, 1878
Syn: O. hendriei Eisén, 1893; O. (O.) occidentalis var. sinensis Eisén, Citation1900 ; O. occidentalis var. arizonae Eisén, Citation1900; O. tenellus Gates, Citation1945.
Distr: ARG: Cordoba Prov: Primeiro river; Santa Fé Prov: La Capital (Righi Citation1990, p. 70). BOL: Vaca Diez Prov: Beni Dep: Guayaramerín (Zicsi Citation1995a, p. 604). COL: Guajira Dep: Hacha river (Zicsi Citation1995b, p. 53); Medellín (Michaelsen Citation1914, p. 202). BRA: SC: 2; PR: 1–3; SP: 1, 4; MS: 4; MT: 3; PA: 5; AM: 3.
Furth. Distr: Central and North America (Righi Citation1990: 69), Europe (Rota Citation2004), Africa (Blakemore Citation2002, p. 89), Asia, including Japan (Eisén Citation1900, p. 115; Gates Citation1945, p. 223; Blakemore et al. Citation2006, p. 173), and Australia (Blakemore Citation1999, p. 182).
Rem: Blakemore (Citation2002, p. 89) suggests that Ilyogenia africana Beddard, 1893 might be a synonym of Ocnerodrilus occidentalis. Beyond extending the known distribution of O. occidentalis to Africa, this necessitates considering Ilyogenia a junior synonym of Ocnerodrilus, because I. africana is the type species of the former genus. Lee (Citation1985) remarks that this species also reaches temperate regions. As anticipated by Lee (Citation1985), this species is spreading to temperate regions.
61*. O. potyura Righi, Citation1994
Distr: BRA: PB: 2.
Paulistus Michaelsen, Citation1926
Type: P. taunayi Michaelsen, Citation1926
62*. P. taunayi Michaelsen, Citation1926
Distr: BRA: BA: 2.
Rem: This species may be considered doubtful, as the genus Paulistus is of uncertain position within the family (Omodeo Citation2000, p. 179).
Phoenicodrilus Eisén, Citation1895
Type: P. taste Eisén, Citation1895
63. P. calwoodi (Michaelsen, 1899)
Syn: Ocnerodrilus (Ilyogenia) calwoodi Michaelsen, 1899.
Distr: COL: Antioquia Dep: near Medellin (Michaelsen Citation1914, p. 204).
Furth. Distr: Central America (Righi Citation1994, p. 93) and Canary islands (Cognetti Citation1906, p. 147).
64*. P. taste Eisén, Citation1895
Distr: CHI: Juan island: Wide channel (Eisén Citation1895, p. 63; Zicsi Citation1993, p. 172).
Pygmaeodrilus Michaelsen, 1890
Type: P. quilimanensis Michaelsen, 1890
65*. P. amapaensis Righi, Citation1988
Distr: BRA: AP: 2.
Quechuona Gates, 1941
Type: Rhinodrilus cameliae Michaelsen, 1934
66*. Q. kixo Righi, Citation1981
Distr: ECU: Los Tauos (Righi Citation1981, p. 238).
67*. Q. michaelseni Jamieson, Citation1962
Distr: ARG: Misiones (Jamieson Citation1962, p. 622).
68*. Q. roseni (Michaelsen, Citation1923)
Syn: Quechua roseni Michaelsen, Citation1923.
Distr: PER: Junine Prov: Junin lagune (Michaelsen Citation1923, p. 1).
Xibaro Righi, Citation1981
Type: X. ashmolei Righi, Citation1981
69*. X. ashmolei Righi, Citation1981
Distr: ECU: Los Tayos (Righi Citation1981, p. 241).
70*. X. medioporus Zicsi & Csuzdi, Citation2002
Distr: ECU (Zicsi & Csuzdi Citation2002, p. 125).
Conclusion
Seventy species of Ocnerodrilidae have been registered from tropical and subtropical South America. Sixty of these are known only from South America, while 10 are more widely distributed.
The South American species of Ocnerodrilidae may be divided into three groups: Subtropical, Subtropical‐Tropical, and Tropical. There are 17 subtropical species, 16 of which are endemic in South America and only Eukerria macdonaldi also occurs in the west coast of North America (Table ). There are 15 species spanning both tropical and subtropical South America (Table ). Five of these occur in other continents, Eukerria saltensis and Ocnerodrilus occidentalis having become not only pantropical but also broadly distributed in temperate latitudes of the Holarctic region. The remaining group of 38 species are mainly restricted to tropical South America, with only 4 species reaching into Central America, Africa or Asia (Table ).
Ocnerodrilidae are considered originally endemic to the Neotropical region (Reynolds Citation1994, p. 12). Independent of their status as nested within the Megascolecoidea or elevated to an independent lineage, it should be stressed that Sims’ (Citation1980) classification of the Lumbricina makes sense from a phylogenetic point of view, because Megascolecoidea plus Glossocolecoidea can be justified as representing a sister clade to Lumbricoidea, both morphologically as well as biogeographically. While Lumbricoidea are Laurasian and have small ovaries with single egg strings, both Glossoscolecoidea and Megascolecoidea are Gondwanan and have large, multilobed ovaries arranged into multiple egg strings, as well as presenting specialized oesophageal or intestinal gizzards along the alimentary canal. Ocnerodrilids also share with all megascolecoids and most glossoscolecoids the development of extramural calciferous glands.
The above views on phylogeny are not followed by all authors, however. For example, Omodeo (Citation2000) considers the Glossoscolecidae as being a part of the Lumbricoidea.
Acknowledgements
Many thanks to Jörg Römbke for providing me with an important reference on oligochaetes. This research was conducted with a scholarship from CNPq, the Brazilian National Science Foundation.
References
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Appendix I. Localities in Brazil
AM: 1: Sucundari: Transamazonian highway, stretch Humaitá–Jacaréacanga, km 523 (Righi et al. Citation1978, p. 28, 30); 2: Manaus–Itacoatiara highway, km 26: Ducke reserve (Righi et al. Citation1978, p. 27); 3: Codajás (Righi Citation1988a, p. 338); 4: Sucundari: Saibroso creek (Righi et al. Citation1978, p. 33).
AP: 1: Marajó island (Righi Citation1968a, p. 180); 2: Serra do Navio (Righi Citation1988, p. 345).
BA: 1: Itajubá (Righi 1971, p. 3); 2: Itabuna (Michaelsen Citation1926, p. 317).
BRA: 1 (Alves & Dal Pai Macha Citation1984, p. 1); 2 (Moreira Citation1903, p. 125).
ES: 1: Porto Cachoeira (Michaelsen Citation1924, p. 80).
GO: 1: Britânia (Brown et al. Citation2006, p. 354).
MG: 1: Serra do Cipó: Vespasiano–Conceição do Mato Dentro highway, km 128 (Righi Citation1971b, p. 388); 2 (Michaelsen Citation1927, p. 370).
MS: 1 (Righi Citation1990, p. 71); 2: Maracajú and Terenos (Righi Citation1984a, p. 19); 3: Ladário, near Corumbá (Righi Citation1990, p. 65, 74); 4: Bela Vista (Righi Citation1994, p. 93).
MT: 1 (Righi Citation1984b, p. 195); 2: Pontes e Lacerda (Righi Citation1984b, p. 195, 198; Righi & Guerra Citation1985, p. 147); 3: Poconé (Righi Citation1984a, p. 19; Righi Citation1994, p. 93); 4: Cuiabá (Righi Citation1984c, p. 13); 5: Cuiabá–Rondonópolis highway, km 25 (Righi Citation1972, p. 151); 6: Cáceres (Righi Citation1984b, p. 195, 198; Righi & Guerra Citation1985, p. 146); 7: Pontes–Lacerda–Vilhena highway, km 30: Três Corações farm, near Branco river (Righi & Guerra Citation1985, p. 147); 8: Pontes–Lacerda–Vilhena highway, km 45, Três Irmãos farm (Righi & Guerra Citation1985, p. 147); 9: Pontes–Lacerda–Vilhena highway, km 60: Sapé farm (Righi & Guerra Citation1985, p. 146, 147); 10: Pontes e Lacerda–Vilhena highway, km 88, Cerejeiras farm, near Seixas river (Righi & Guerra Citation1985, p. 147); 11: Pontes e Lacerda–Vilhena highway, km 123, Paraíso farm (Righi & Guerra Citation1985, p. 147); 12: Nova Alvorada (Righi & Guerra Citation1985, p. 147; Righi Citation1990, p. 74); 13: Urucúm, near Corumbá (Cognetti Citation1900, p. 3); 14: Ladário, near Corumbá (Cognetti Citation1900, p. 6); 15: Tabuleta (Righi & Guerra Citation1985, p. 146); 16: Vila Bela da Santíssima Trindade (Righi Citation1984b, p. 198; Righi & Guerra Citation1985, p. 146); 17: near Guaporé river (Righi & Guerra Citation1985, p. 146).
PA: 1: Belém: Guamá river (Righi Citation1971a, p. 2); 2: Amazonia national park: Tapajós: Uruá and Mergulhão streams (Righi Citation1983, p. 927); 3: Amazônia national park, Tapajós: Saita: Tapajós river (Righi Citation1983, p. 927); 4: Marajó island (Righi Citation1968b, p. 545): 5: Marajó island: Arari waterfall (Righi Citation1968a, p. 183; Righi Citation1988a, p. 338); 6: Amazonia national park: Tapajós: Uruá creek (Righi Citation1983, p. 932); 7: Belém (Righi Citation1975, p. 78).
PB: 1: João Pessoa (Righi Citation1994, p. 99, 102); 2: Mari (Righi Citation1994, p. 96).
PE: 1: Cabo (Righi Citation1971a, p. 1).
PR: 1 (James & Brown Citation2006, p. 56); 2 (Sautter et al. Citation2006, p. 298); 3: Jaguapitã (Nunes et al. Citation2006, p. 331).
RJ: 1: Itatiaia (Michaelsen Citation1924, p. 79).
RO: 1: BR 364 highway, stretch Presidente Médici–Cacoal, km 12 (Righi Citation1988b, p. 120); 2: BR‐364 highway, stretch Pimenta Bueno–Vilhena, km 5: near Riozinho river (Righi Citation1988b, p. 120); 3: BR 364 highway, stretch Pimenta Bueno–Vilhena, km 42: São Vitorino farm (Righi Citation1988b, p. 120); 4: Mirante da Serra highway, km 32 (Righi Citation1988b, p. 121); 5: BR‐364 highway, stretch Pimenta Bueno–Vilhena, km 57 (Righi Citation1988b: 121); Loc. 6: Mirante da Serra highway, km 13 and km 62 (Righi Citation1988b, p. 120); 7: RO‐421 highway, km 7: Tabapuá river (Righi Citation1988b, p. 120); 8: BR‐364 highway, stretch Arquimenes–Jaru, km 24 (Righi Citation1988b, p. 120); 9: BR‐364 highway, stretch Jaru–Ouro Preto do Oeste, km 30 (Righi Citation1988b, p. 120); 10: Presidente Médici–Cacoal stretch, km 61 (Righi Citation1988b, p. 120); 11: Pimenta Bueno–Espigão do Oeste highway, km 18 (Righi Citation1988b, p. 120).
RR: 1: Maracá reserve (Righi & Guerra Citation1985, p. 147); 2: Boa Vista–Bonfim highway, km 100, Santana farm (Righi & Guerra Citation1985, p. 147); 3: Bonfim (Righi & Guerra Citation1985, p. 147).
RS: 1: Camaquã (Lima & Rodríguez Citation2007, p. 15); 2: Estrela (Righi & Ayres Citation1975, p. 309); 3: Jaguapitã (Nunes et al. Citation2006, p. 331); 4: Porto Alegre (Michaelsen Citation1927, p. 370).
SC: 1: Blumenau (Righi Citation1968c, p. 370); 2: Lauro Müller (Brown et al. Citation2006, p. 354).
SE: 1: Umbauba: Nova Esperança farm (Righi Citation1980a, p. 240).
SP: 1 (Brown & James Citation2006, p. 147); 2: Iporanga Mun: Paiva cave (Righi Citation1995, p. 362); 3: Botucatu (Brown et al. Citation2006, p. 353); 4: São Paulo: Anhembi (Righi Citation1968c, p. 370, 373; Righi Citation1968b, p. 545; Righi Citation1971b, p. 3); 5: Ilha Bela (Brown et al. Citation2006, p. 354); 6: Birigui (Brown et al. Citation2006, p. 354); 7: Rio Claro (Brown et al. Citation2006, p. 354); 8: Piracicaba (Michaelsen Citation1926, p. 321); 9: Salesópolis: Boracéia biological station (Righi Citation1980b, p. 2); 10: São Sebastião: Pitangueira creek (Righi Citation1968c, p. 372; Brown & James Citation2006, p. 147).