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Home , Argilophilus, Driloleirus, Eudrilidae, Eudrilus, Eudrilus eugeniae, Megascolecidae

Turkish Journal of Zoology Turk J Zool (2017) 41: 599-614 http://journals.tubitak.gov.tr/zoology/ © TÜBİTAK Research Article doi:10.3906/zoo-1604-74

A catalogue of the and (: Lumbricina) from South America, with two new records of exotic from Brazil

1, 2 2 José Eriberto DE ASSIS *, José Roberto Botelho SOUZA , Maria Laís Martins VIEIRA , 2 2 1 João Vitor NUNES DE SOUZA , Gilberto Gonçalves RODRIGUES , Martin Lindsey CHRISTOFFERSEN 1 Department of Systematics and Ecology, Exact and Natural Science Center, Federal University of Paraíba, João Pessoa, Paraíba, Brazil 2 Department of Zoology, Biosciences Center, Federal University of Pernambuco, Recife, Brazil

Received: 30.04.2016 Accepted/Published Online: 14.01.2017 Final Version: 17.07.2017

Abstract: A catalogue of the terrestrial Eudrilidae and Megascolecidae is provided, with 10 genera and 18 species reported to date from South America. In addition, we provide the valid names, their synonyms, and the original citation. Eudrilidae and Megascolecidae contain members that have a global distribution, probably as a result of human dispersion. The species of eudrilidEudrilus eugeniae is very common in other regions with a tropical climate, while the megascolecid species gracilis is by far the most common in large cities such as São Paulo and Rio de Janeiro in Brazil. Two samples taken in Brazil revealed two new exotic species from this country: Amynthas robustus (Perrier, 1872) and bicincta (Perrier, 1875), which are described and illustrated. Finally, we provide a taxonomic key to the genera of both families.

Key words: , exotic species, Eudrilidae, Megascolecidae, South America

1. Introduction some islands of the western Pacific Ocean) (Easton, 1982). The Eudrilidae are endemic to the Ethiopian About 40 species are found outside this area (Easton, 1982; region (Reynolds, 1994). They present extreme variations Blakemore, 2002, 2010). Many pheretimoids can jump by in the sexual apparatus, particularly of the female organs means of powerful contractions of the body (Omodeo, (Omodeo, 2000; Blakemore, 2002, 2010). Eudriline 2000). All megascolecids dealt with herein are members of evolution has resulted in economy of sperm production. this group. Modifications in the male apparatus appear to reduce, or Eisen (1900) and Michaelsen (1903, 1935) noticed prevent, gregarine ravages (Sims, 1987). eugeniae that, in the Caribbean, native earthworms were vanishing has achieved a pantropical distribution (Omodeo, 2000). from regions influenced by human activity and were being Megascolecidae is the most diverse family of earthworms replaced by exotic species. By now, it is well known that (Blakemore, 2002); it has over 1000 species (Sims and exotic species dominate in disturbed ecosystems, with Gerard, 1985; Blakemore, 2002). Most of them are native native species predominating in natural, undisturbed to (Blakemore, 2002) and (Sims and environments (Fragoso et al., 1995). Gerard, 1985). One of the family , The rapid pace of destruction in the tropical Diplocardia, is native to North America. Several genera of regions is widely recognized as a threat to biotic diversity. Megascolecidae sensu stricto [i.e. Arctiostrotus, , Peregrine species may eventually cover the tropical Chetcodrilus, Drilochaera, (the only meroic regions. Only a few native species are capable of surviving genus), Kincaidodrilus, Macnabodrilus, Nephrallaxis, and and coexisting with exotic worms in disturbed ecosystems Toutellus], placed under the as of yet poorly defined tribe (Fragoso et al., 1995). Argilophilini Fender & McKey-Fender, 1990 (Gates, 1942; Our aim is to provide a catalogue of the Eudrilidae and James, 1995; Blakemore, 2000, 2010), are native to North Megascolecidae reported to date from the South American America. auct. (= the pheretimoids) is a group continent and describe and report two new exotic species of genera within Megascolecidae that are indigenous to the of pheretimoids for northeastern Brazil. We also provide a so-called Pheretima domain (all of southern Asia, Japan, the taxonomic key to the genera of the two families reported Indo-Australasian archipelago, the Solomon Islands, and herein. * Correspondence: [email protected] 599 DE ASSIS et al. / Turk J Zool

2. Materials and methods James and Brown, 2006): Vinhedo (Righi, 1968); São Data for geographical distribution were collected from Sebastião; Boituva; Campinas; São Paulo (Brown et al., earthworm bibliographies. We provide the following 2006); RJ: Petrópolis (Michaelsen, 1927); Nova Friburgo information: families, genera, species, and other groups, (Brown et al., 2006); MG: Juiz de Fora (Brown et al., 2006); organized alphabetically, with original citation, year of BA: Itajubá and Jequié; SE: Aracajú (Brown et al., 2006); publication, and all related citations from South America. PB: Areia (Guerra and Silva, 1994); PE: Recife (Righi, The synonyms are provided according to Blakemore (2010, 1967a, 1971); Ponta de Pedras; MA: São Luís (Brown et 2014). In the case of Amynthas corticis, those references al., 2006); PA: Vila de Maiauatá, margins of Pará River, pertaining to A. diffringens (Baird, 1869) and its synonyms 20 km southwest of Abaetetuba (Righi, 1972); COL: Valle have been removed from the synonymy of A. corticis, Dep.: Palmira (Feijoo, 2007); San Andrés Island (Reynolds as A. diffringens has been deemed valid by Blakemore and Reynolds, 2001). VEN: Caracas (Michaelsen, 1900a); (2013). We used 3-letter abbreviations for South American Portamer; Isla Margarita (Michaelsen, 1892; Zicsi, 1995a). countries (ARG, Argentina; BOL, Bolivia; BRA, Brazil; GUY and SUR (Beddard, 1887; Michaelsen, 1900a). FRG: CHI, Chile; COL, Colombia; ECU, Ecuador; FRG, French Saint Georges Oyapock (Michaelsen, 1900a; Cernosvitov, Guyana; GUY, Guyana; PER, Peru; SUR, Surinam; URU, 1934, 1935) (Figure 1A). Uruguay; VEN, Venezuela), and 2-letter abbreviations Further distribution: Caribbean region (Zicsi, 1995a). for sampled states in Brazil (AM, Amazonas; BA, Bahia; North America, Africa, Asia, New Caledonia (Gates, DF, Distrito Federal; MG, Minas Gerais; PA, Pará; PB, 1972), and Australia (Blakemore, 2002, 2010, 2014). Paraíba; PE, Pernambuco; PR, Paraná; RJ, Rio de Janeiro; Genus Hyperiodrilus Beddard, 1890 RR, Roraima; RS, Rio Grande do Sul; SC, Santa Catarina; Type: Hyperiodrilus africanus Beddard, 1891 SP, São Paulo); “?” indicates taxonomic uncertainty. New 2. Hyperiodrilus africanus (Beddard, 1891) records herein are based on two samples taken on 28 Distribution: West and Central Africa: from Nigeria, April 2009 and 8 May 2010 in an urban area located in Ghana, and Ivory Coast to Chiloango River in “Portuguese Campina Grande, State of Paraíba, in northeastern Brazil Congo” (Blakemore, 2010); Pontas de Pedras, PE, Brazil (7°13′28″S, 35°52′2″W and 7°12′56″S, 35°53′50″W). They (Righi, 1972; Brown et al., 2006) (Figure 1A). were collected by hand-sorting, fixed in 10% formalin Family MEGASCOLECIDAE Rosa, 1891 solution, and preserved in 70% alcohol. The specimens are Genus Amynthas Kinberg, 1867 deposited in Invertebrate Collection Paulo Young of the Type: Amynthas aeruginosus Kinberg, 1867 Universidade Federal da Paraíba (LIPY). 3. Amynthas aeruginosus (Kinberg, 1867) Amyntas aeruginosus Kinberg, 1867: 101. [Type 3. Results locality: Guam, Marianas Islands. Types in Stockholm Museum: 154]. ?Perichaeta upoluensis Beddard, 1887: Suborder LUMBRICINA (part.) De Blainville, 1828 174, Pl V, figs. 11–12;Perichaeta oeriginosa (?lapsus pro Terricolae (part.) Örsted, 1843; Megadrili Benham, 1890 aeruginosa) Beddard, 1891: 278. Pheretima aeruginosus: Superfamily MEGASCOLECOIDEA Rosa, 1891 Michaelsen, 1900: 253 (syn. oeriginosa [sic]); Cognetti, Family EUDRILIDAE Claus, 1880 1913b: 37. ?Pheretima rennellana Gates, 1959: 11. [Type Genus Eudrilus Perrier, 1871 locality, Hutuna, Lake Te-Nggano, Rennell. Types in British Type: Eudrilus decipiens Perrier, 1871 (= E. eugeniae) Museum, 1957: 11: 5: 25–26]. Amynthas aeruginosus 9829 1031 (part): Sims and Easton, 1972: 211 (?syn. A. upoluensis, A. 1. (Kinberg, 1867) rennellanus); Lee, 1981: 542, figs. 2–6 (?syn. A. upoluensis); Lumbricus eugeniae Kinberg, 1967: 98. [Type locality: Blakemore, 1997: 22, fig. 2 (part = A rennellanus). humid mountains and Valley of St Helena Island (British Distribution: PR (Sautter et al., 2006): Prudentópolis protectorate) in South Atlantic. Types are in Stockholm (Brown et al., 2006; Brawn and James, 2006) (Figure 1B). Museum and/or in British Museum BMNH 1904.10.5.549 Further distribution: Pacific Ocean (Michaelsen, or 550]. 1900a; Blakemore, 2010). Eudrilus decipiens Perrier, 1871: 1176; Eudrilus lacazii + 4. Amynthas corticis (Kinberg, 1867) peregrinus Perrier, 1872: 75, 77/78; Eudrilus boyeri Beddard, Perichaeta corticis Kinberg, 1867; Megascolex (Perichaeta) 1886: 302; Eudrilus sylvicola Beddard, 1887: 372. Eudrilus diffringens Baird, 1869; Perichaeta subquadrangula Grube, jullieni Horst, 1890: 225; Eudrilus roseus Michaelsen, 1892: 1877; Perichaeta sp. Horst, 1878; Megascolex indicus Horst, 224; Eudrilus erudiens Ude, 1893: 71. 1883; Perichaeta peregrina Fletcher, 1886; ?Perichaeta Distribution: BRA (Righi, 1967a): PR (Nunes et al., mirabilis Bourne, 1887; ?Perichaeta sp. Garmann, 1888; 2006; Sautter et al., 2006); Londrina; Ibiaci (Brown et al., Perichaeta heterochaeta Michaelsen, 1891; ?Perichaeta ijimae 2006); SP (Lüderwaldt, 1927; Brown and James, 2006; Rosa, 1891; Perichaeta nipponica Beddard, 1892; ?Perichaeta

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Figure 1. Geographical distribution in South America of exotic Eudrilidae (A) and Megascolecidae (B–D) species: A) Eudrilus euganiae (circle), and Hyperiodrilus africanus (star); B) Amynthas aeruginosus (circle), and Amynthas corticis (square); C) Amynthas gracilis (triangle); D) Amynthas morrisi (star) and Amynthas robustus (circle) (new record from Brazil).

601 DE ASSIS et al. / Turk J Zool perkinsi Beddard, 1896; ?Perichaeta molokaiensis Beddard, Perichaeta hawayana Rosa, 1891: 396; ?Perichaeta ijimae 1896; Perichaeta heteropoda Goto & Hatai, 1898; Pheretima Rosa, 1891: 402; Perichaeta bermudensis Beddard, marenzelleri Cognetti, 1906; ?Pheretima silvestrii Cognetti, 1892a: 160; ?Perichaeta pallida Michaelsen, 1892: 227; 1909; Pheretima divergens yunnanensis Stephenson, Perichaeta mandhorensis Michaelsen, 1892: 241, figs. 18-19; 1912; ?Pheretima indica var. cameroni Stephenson, 1932; ?Perichaeta barbadensis (part., spec “b”?) Beddard, 1892a: ?Pheretima sheni Chen, 1935; ?Pheretima directa Chen, 167; ?Perichaeta mauritiana Beddard, 1892a: 170, Pl. X, 1935; Pheretima oyamai Ohfuchi, 1937; Pheretima tajiroensis figs. 5–6; ?Perichaeta kamakurensis Goto & Hatai, 1898: Ohfuchi, 1938; ?Pheretima morii Kobayashi, 1938; Pheretima 68; ?Perichaeta parvula Goto & Hatai, 1898: 68. [Types?]. toriii Ohfuchi, 1941; ?Pheretima clerica Benham 1947; [Non Ohfuchi, 1956 (= parvula)]; ?Perichaeta Pheretima campestris Lee, 1952; ?Pheretima hatomajimensis decimpapillata Goto & Hatai, 1898: 71; Amyntas ijimae: Ohfuchi, 1957; Amynthas nanrenensis James et al., 2005. Beddard (1900: 635); Pheretima hawayana: Michaelsen, Distribution: ARG: Buenos Aires Prov.; Catamarca 1900: 271, 316; Amyntas hawayanus: Beddard, 1900: 420; Prov.; Misiones Prov.; Santa Fé Prov. (Mischis, 2004; ?Pheretima kagoshimensis Takahashi, 1932: 343; ?Pheretima Mischis and Herrera, 2006; Mischis, 2007); Tucumán kamitai Kobayashi, 1934: 5, figs. 4–6; ?Pheretima hawayana Prov.: Potrerillo (Gates, 1972) and Villa Nougues var. barbadensis: Cernosvitov, 1934: 49. (Teisaire, 1991); Córdoba Prov. (Mischis, 1992): Capital, Amynthas gracilis: Sims and Easton, 1972: 235 (in an Calamuchita, Colón, and Santa Maria Deps. (Mischis, A. hawayanus group containing nearly 50 nominal taxa); 1993); San Luis Prov. (Mischis and Brigada, 1985); Salta Easton, 1981: 50 (syn. hawayana, ?kamakurensis, ?parvula, (Teisaire et al., 2003). BOL: Murillo Prov: La Paz Dep.: decimpapillata, ?carnosa, ?kagoshimensis); Easton, 1982: Vale del Rio Zongo, near Cambaya (Righi and Römbke, 728 (syn. hawayana); Sims and Gerard, 1985: 130 (syn. 1987): Cahura St. (Zicsi, 1995b). PER (Michaelsen, 1892). hawayana); Blakemore, 2002, 2008. ECU: Pichincha Prov.: Quito (Zicsi and Csuzdi, 1988). Distribution: CHI: Santiago: Quinta Normal (Zicsi, COL: Palmira, El Cerrito, Caldono, Silva Santander 2004). ARG (Dulout et al., 1969a; Lauría De Cidre, 1972): Rosa, Mocoa Putumayo (Feijoo et al., 2004); Cabuyal Chubut Prov.: Piedra del Aguila; Puerto Madryn (Mischis River (Feijoo et al., 1999); Andean hillside (Feijoo et al., et al., 2006); Neuquén Prov.; San Luís Prov. (Mischis, 2006); Bogotá (Michaelsen, 1900a). VEN: In earth with 2004, 2007; Mischis and Herrera, 2006); Santa Fé Prov. plant arriving by plane in San Juan, Puerto Rico. BRA: (Ljungström et al., 1973); Córdoba Prov.: Sierras Chicas From earth for nematode analysis arriving by plane in (Mischis, 1999); Dep. Capital, Dep. Calamuchita (Mischis, New York (Gates, 1982): RS (Righi, 1965; Knäpper, 1977; 1993); Tucumán Prov.: Tucumán (Teisaire and Roldán, Knäpper and Porto, 1979; Krabbe et al., 1993): Nova 1995; Alderete-De-Majo, 2002a, 2002b, 2002c, 2003a, Petrópolis (Knäpper, 1967); Santo Ângelo; Santa Cruz 2003b); Catamarca Prov.: Las Pirquitas Ville (Mischis and do Sul; Santana do Livramento; Gramado; Caxias do Sul; Righi, 1999); Buenos Aires Prov.: Buenos Aires (Mercadal Estância Velha; Lajeado; Tramandaí (Knäpper, 1972a); Del Barrio and Barrio, 1988; Burela and Cazzaniga, 2001). São Leopoldo (Righi and Knäpper, 1965); Camaquã Mun. URU: Montevideo (Cordero, 1931). ECU (Fragoso and (Lima and Rodriguez, 2007); PR (Voss, 1986; Dionísio et Brown, 2007; Zicsi, 2007). COL: Buenaventura; Palmira; al., 1995; Sautter et al., 1995; Chang, 1997; Tanck et al., Caldono Santander Cauca; Cali; Mocoa Putumayo (Feijoo, 2000; Brown et al., 2004; Sautter et al., 2006): Londrina 2007); Andean hillside (Feijoo et al., 2006); Cabuyal River (Brown et al., 2003); Arapoti (Peixoto and Marochi, 1996); (Feijoo et al., 1999). VEN (Drachenberg, 1992): From SP (Righi, 1965; Brown and James, 2006): northwest of earth with plants arriving by plane at San Juan, Puerto São Paulo State (Caballero, 1976); Campos do Jordão and Rico (Gates, 1982). FRG (Cernosvitov, 1934). BRA São Sebastião (Righi, 1967b); RJ (Zicsi and Csuzdi, 1999): (Moreira, 1903; Righi, 1967b): RS (Michaelsen, 1892; Seropédica (Brown et al., 2006); Nova Friburgo (Gates, 1927; Knäpper, 1972b; Knäpper and Porto, 1979; Krabbe 1954); Parque Nacional do Itatiaia, near Abrigo Rebouças; et al., 1993); Guaíba estuary (Knäpper, 1976); Estrêla MG: Jaboticatuba: Serra do Cipó: Estrada Vespaziano- (Righi and Knäpper, 1965); Santo Ângelo; Santa Cruz do Conceição do Matodentro, km 120 (Righi, 1980); Juiz de Sul; Santana do Livramento; Gramado; Nova Petrópolis; Fora (Brown et al., 2006); AM (James and Brown, 2006) Ivoti; Sapucaio; Canoas; Lajeado; Estrêla; Tramandaí (Figure 1B). (Knäpper, 1972a); Camaquã Mun. (Lima and Rodriguez, Further distribution: Panama to Mexico (Righi, 2007); SC: Blumenau; Schroeder (Brown et al., 2006); PR 1967b), (Beddard, 1893), Asia (Brown et al., 2003), (Voss, 1986; Dionísio et al., 1995; Kobiyama et al., 1995; and Australia (Easton, 1982). Sautter et al., 1995; Peixoto and Marochi, 1996; Chang, 5. Amynthas gracilis (Kinberg, 1867) 1997; Tanck et al., 2000; Brown et al., 2004): Londrina Nitocris gracilis Kinberg, 1867: 102. [Type locality (Brown et al., 2003); Curitiba (Zicsi and Csuzdi, 1999); SP Rio de Janeiro; ?[Perichaeta bournei Rosa, 1890: 110; (Lüderwaldt, 1927; Gates, 1972; Lenko, 1972; Caballero,

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1976); Piracicaba; Itatiaia; Paranapiacaba (Alto da Serra); 1997; Sautter et al., 2006); Curitiba (Brown et al., 2006); São Paulo: Ipiranga; Belém (Lüderwaldt, 1927); São Paulo SP (Brown and James, 2006; James and Brown, 2006); BA: University campus (Jorge and Sawaya, 1967); Barueri, in Salvador (Righi, 1971); AM: Manaus (Michaelsen, 1927) ant colonies (Lenko, 1972); São Roque; Rio Preto; Salto (Figure 1D). de Itú (Vannucci, 1953); RJ (Kinberg, 1867; Vannucci, Further distribution: Barbados to North America, 1953); MG: Paraopeba, Fazenda do Rasgão, Cachoeira Europe, South Atlantic Oceanic islands, Africa, Asia, dos Macacos, and Fazenda Pedra Branca (Righi, 1980); Hawaii (Righi, 1979), and Australia (Easton, 1982). DF: Brasília (Brown et al., 2006); BA: Ituberá (Brown et 7. Amynthas robustus (Perrier, 1872) al., 2006); PB: Areia (Guerra and Silva, 1994); PA: Belém Perichaeta cingulata (part cf. P. darnleiensis): Vaillant, (Brown et al., 2006); AM (James and Brown, 2006): 1867: 234; Perichaeta robusta Perrier, 1872: 112, figs. 67– Manaus (Michaelsen, 1927) (Figure 1C). 68; Perichaeta masatakae Beddard, 1892: 761; ?Pheretima Further distribution: Barbados to Mexico, North campestris Goto & Hatai, 1898: 67 [non campestris Lee, 1952 America, Europe, Africa, Asia (Righi, 1979), Australia (= A. corticis)]; ?Perichaeta parvicystis Goto & Hatai, 1899: (Easton, 1982), Hawaii (Rosa, 1891; Nakamura, 1992), and 18, figs. 8, 8a, 8b; Megascolex robustus: Vaillant, 1889: 76; South Atlantic Ocean (Cernosvitov, 1942). Pheretima lauta Ude, 1905: 405 (429?, 464?; Pheretima lauta 6. Amynthas morrisi (Beddard, 1892) Chen, 1933: 282, fig. 26; ?Pheretima zavattarii Cognetti, Perichaeta morrisi Beddard, 1892a (July): 166, Pl. IX, 1909: 1; Cognetti, 1909: 310; Pheretima ornata Gates, 1927: fig. 1; Perichaeta? barbadensis (?part “a” and/or “c”) Beddard, 20; Pheretima corrugata Chen, 1931: 131; 1933: 278 (non 1892a (July): 167, Pl. IX, fig. 7; Perichaeta? mauritiana P. corrugata kulingiana); ?Pheretima sheni Chen, 1935: 38; Beddard, 1892a: 170; ?Perichaeta pallida Michaelsen, Pheretima robusta: Michaelsen, 1900: 299. 1892 (Sept.): 227; ?Perichaeta amazonica Rosa, 1894: 14; Amynthas robustus: Sims and Easton, 1972: 234; ?Perichaeta sanctijacobi Beddard, 1895: 239; ?Perichaeta Easton, 1981: 56 (syn. mastakae [sic lapsus pro masatakae], insulae Beddard, 1896: 516; ?Perichaeta cupulifera Fedarb, campestris Goto & Hatai, 1898 [non Lee, 1952], lauta, 1898: 445, fig. 1; Amynthas barbadensis: Michaelsen, 1900: corrugata); Blakemore, 2002, 2008. 254 (syn. pallida Michaelsen, 1892: 227, amazonica Rosa, New record: Campina Grande, Paraíba, Brazil: (BRA: 1894:14, sanctijacobi Beddard, 1895: 61, cupulifera Fedarb, PB). 1898: 445); Pheretima hawayana lineata Gates, 1926: 154; Further distribution: Mauritius, Philippines, Myanmar, 1931: 384–386, fig. 24; Pheretima elongata : Ohfuchi, 1956: India, China, Taiwan, Korea, Japan, Okinawa?, Madagascar 148; Pheretima exiloides: Ohfuchi, 1956: 142; Pheretima (Gates, 1972; Blakemore, 2002, 2010); São Paulo, Brazil morrisi: Michaelsen, 1900: 287. (Righi, 1967), cited as P. aspergillum. Amynthas morrisi: Sims and Easton, 1972: 236, figs. Examined material: Specimen: One adult specimen 1A, 1H; Easton, 1981: 55 (syn. exiloides: Ohfuchi, elongata: collected on 8 May 2010 in an urban area located in Ohfuchi); Easton, 1982: 729, fig. 4c; Sims and Gerard, 1985: Campina Grande (7°12′56.60″S, 35°53′50.34″W), State of 132, fig. 47a (syn. barbadensis, mauritiana); Blakemore, Paraíba, Northeast Brazil (UFPB 1460) (Figure 1D). 2002, 2008; Chang et al., 2009: 60, fig. 26. Description: Length: 105 mm. Diameter: 4–5 mm. Distribution: CHI (Zicsi, 1993): Santiago: Quinta Segments: 96. Prostomium: open epilobous. First dorsal Normal (Beddard, 1895). ARG (Mischis, 2004; Mischis pore: 11/12. Setae: perichaetine. Nephropores: absent. and Herrera, 2006): Tucumán Prov. (Teisaire and Roldán, Clitellum: 14–16, annular, darker than the rest of the 1995; Alderete-De-Majo, 2002a, 2002b, 2002c, 2003a, earthworm. Spermathecal pores: 2 pairs in 7/8/9. Female 2003b); Santa Fé Prov. (Ljungström et al., 1973): La pores: single on 14. Male pores: a pair on conical porophores Capital (Righi, 1978); Buenos Aires Prov. (Righi, 1979); in 18. Genital markings: one pair anterior-median to the Córdoba (Mischis, 1993): Sierra Chicas (Mischis, 1999); male pores (Figure 2A). Septa: 5/6–7/8 thickened and Catamarca Prov.: Las Pirquitas Ville (Mischis and Righi, 8/9/10 absent. Spermathecae: 2 pairs in 8 and 9 (Figure 1999). PER (Righi, 1979). PER: Tumbes Prov.: Negritos 2B). Nephridia: meroic. Gizzard: 8–9. Hearts: 10–13. (Gates, 1954, 1972; Fragoso and Brown, 2007; Römbke, Calciferous glands: absent. Ovaries: a pair in 13. Testes: 10 2007); Pichincha Prov.: Cunuyacu (Zicsi and Csuzdi, and 11, seminal vesicles large in 11 and 12, pseudovesicles 1988). BRA (Cernosvitov, 1934): RS (Michaelsen, 1927; in 13–14. Prostate glands: racemose, opening in 18 and Knäpper, 1972b; Knäpper and Porto, 1979; Krabbe et extending into 16–19 (Figure 2C). Intestinal origin: 15, al., 1993); Lageado (Righi and Knäpper, 1965); Novo with a pair of ceca extending into 26–20 (Figure 2D). Hamburgo (Knäpper, 1967); Santo Ângelo; Santa Cruz do Remarks: A. robustus was found in association Sul; Nova Petrópolis; Ivoti; Caxias do Sul; Estância Velha; with E. eugeniae in the first 10-cm layer of with a São Leopoldo. Sapucaio; Tramandaí (Knäpper, 1972a); high concentration of organic matter. A. robustus often Camaquã Mun. (Lima and Rodriguez, 2007); PR (Chang, reproduces by parthenogenesis because of male sterility.

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Figure 2. A) Amynthas robustus, anterior body part in ventral view, genital markings on 18; B) spermatheca with a single diverticulum; C) prostatic gland; D) intestinal caecum.

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A record from America based on a British Museum Chicas (Mischis, 1999); Tucumán Prov. (Teisaire and specimen was erroneously labeled A. gracilis according to Roldán, 1995; Alderete-De-Majo, 2002a, 2002b, 2002c, Gates (1972). Fragoso and Brown (2007) considered this 2003a, 2003b): San Miguel de Tucumán (Dulout et al., species as exotic for Brazil. 1969c; Levin et al., 1969); Santa Fé Prov. (Ljungström et 8. Amynthas rodericensis (Grube, 1879) al., 1973): San Tomé (Righi, 1978); Salta (Teisaire et al., Perichaeta rodericensis Grube, 1879: 554; Perichaeta dyeri 2003). PER (Righi, 1979): From earth around plant in Beddard, 1892: 57, Pl. IX, figs. 2, ,8 Pl. X, fig. 1; Perichaeta baggage by plane arriving in New Orleans (Gates, 1982); sinensis Beddard, 1892a: 158, Pl. IX, figs. 3, 5, Pl. X, figs. 2–4, Huanco Prov: Chogosh (Michaelsen, 1935; Fragoso and 7, 8; ?Perichaeta trinitatis Beddard, 1896: 206; Perichaeta Brown, 2007; Römbke, 2007). VEN (Drachenberg, 1992). monilicystis Michaelsen, 1892: 251, fig. 28; Pheretima BRA: RS (Knäpper and Porto, 1979; Krabbe et al., 1993): rodericensis: Michaelsen, 1900: 299, 318 (syn. dyeri, sinensis, Estrêla (Righi and Knäpper, 1965); São Leopoldo; Novo monilicystis, ?trinitatis). Hamburgo (Knäpper, 1967); Santa Cruz do Sul; Santana Amynthas rodericensis: Sims and Easton, 1972: 235; do Livramento; Gramado; Nova Petrópolis. Ivoti; Caxias Easton, 1982: 729; Sims and Gerard, 1985: 132 (syn. dyeri, do Sul; Estância Velha; Sapucaio; Canoas; Lajeado; sinensis). Tramandaí (Knäpper, 1972a); SC: Lauro Müller (Brown et Distribution: VEN (Drachenberg, 1992): Puerto Rico: al., 2006); PR (Chang, 1997; Sautter et al., 2006): Curitiba Saba (Righi, 1993); Aragua State: Parque Nacional Henri (Brown et al., 2006); SP (Righi, 1967b; Brown and James, Pittier: Rancho Grande (Righi, 1984); Caracas: Monte 2006): Piracicaba; São Paulo: Ipiranga (Lüderwaldt, 1927); Avila, 950 m (Righi, 1989). SUR (Gates, 1972; Fragoso RJ: Rio de Janeiro (Moreira, 1903); MG: Caetanópolis and Brown, 2007). FRG: Morne de Montabe (Cernosvitov, (Righi, 1980); BA: Salvador (Righi, 1971) (Figure 3A). 1934). GUY (Beddard, 1900; Cernosvitov, 1942) (Figure Further distribution: Central and North America, 3A). Atlantic Oceanic islands, Europe, Africa, Middle East, Further distribution: Caribbean region (Zicsi, 1995a), Asia, Australia, Hawaii, and Marquesas Islands (Righi, North America, Europe, Asia, Australia, and Pacific islands 1979). (Easton, 1982). Remarks: Blakemore et al. (2012) added this species Genus Metaphire Sims and Easton, 1972 to the Korean species biodiversity list as an exotic species Type: Rhodopis javanica Kinberg, 1867 (non from Korea. This species is also an exotic species from Woodwardia javanica Michaelsen, 1910 = Lennoscolex Brazil (Fragoso and Brown, 2007). Gates, 1960) 10. Metaphire houlleti (Perrier, 1872) 9. Metaphire californica (Kinberg, 1867) Perichaeta sp. Perrier, 1871: 278; Perichaeta houlleti Pheretima californica (part) Kinberg, 1867: 102 [excl. Perrier, 1872: 99, figs. 31–63; Beddard, 1887: 389; Megascolex Megascolex indicus Horst, 1883 (= A. corticis - see Easton, houlletii: Vaillant, 1889: 75; Perichaeta campanulata Rosa, 1981: 49)]; Perichaeta ringeana Michaelsen, 1890: 10; 1890: 115, t. 1, figs. 9–10; Perichaeta udekemi Michaelsen, 1899: 116; ?Perichaeta sluiteri Horst, 1890: 234, figs. 4–5; 1890: 240, t. 13, fig. 17; Perichaeta guillelmi Michaelsen, ?Perichaeta sumatrana: Beddard, 1892a: 155, Pl. IX, fig. 4. 1895: 32, fig. 7; Pheretima crescentica Fedarb, 1898: 447; ?non Megascolex sumatranus Horst, 1883: 189; Perichaeta Amyntas houlleti: Michaelsen, 1899: 12; Pheretima houlleti: hesperidum Beddard, 1892a: 169; Perichaeta guarini Rosa, Michaelsen, 1900: 273 (syn. campanulata, udekemi, 1894: 13; ?Perichaeta sandvicensis Beddard, 1896: 203; guillelmi); Amyntas kelantanensis Beddard, 1900: 902; Amyntas hesperidum: Beddard, 1900: 417; Pheretima browni Pheretima wimberleyana Stephenson, 1925: 62; Pheretima Stephenson, 1912: 273 (part excluding sexthecal morphs); houlleti var. tortuosa Gates, 1926: 454; Pheretima houlleti Pheretima kiangensis Michaelsen, 1931: 21; Pheretima var. rugosa Gates, 1926a: 157; Pheretima rugosa Gates, 1932: modesta Michaelsen, 1927: 88. [From Myanmar. Types 398; Pheretima campanulata var. penetralis Gates, 1931: 435; in Hamburg 10423]; Pheretima molesta Gates, 1932: 420; Pheretima campanulata var. meridiana Gates, 1932: 457; Pheretima sakaguchii Ohfuchi, 1938: 53, figs. 1–2; ?Pheretima ?Pheretima yapensis Ohfuchi, 1941: 283; Metaphire houlleti sonaiensis Ohfuchi, 1956; Pheretima californica: Michaelsen, houlleti: Sims and Easton, 1972: 238 (houlleti-group); 1900: 258, 275 (syn. ringeana, guarini, ?hesperidum). Metaphire houletti (lapsus pro houlleti): Easton, 1982: 731; Metaphire californica Sims and Easton, 1972: 238; Easton, 1983: 121, tab. 1, 128; Amynthas huangi James et al., 2005: 1981: 57 (syn. ?sakaguchii, ?sonaeiensis); Easton, 1982: 731; 1014, fig. 2E, 2F (on p. 1010). Sims and Gerard, 1985: 132, fig. 47c (syn. hesperidum). Metaphire houlleti (Blakemore, 2002, 2004, 2008, 2010). Distribution: ARG (Dulout et al., 1969b, 1969c; Levin Distribution: FRG: Père Island (Cernosvitov, 1934) et al., 1969; Mischis, 2004): From earth with bougainvillea (Figure 3A). plant in luggage arriving by plane in New York airport Further distribution: Central and North America, (Gates, 1982): Cordoba Prov. (Mischis, 1993): Sierras Europe, Africa, Asia, and Australia (Easton, 1982).

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Figure 3. Geographical distribution in South America: A) Amynthas rodericensis (circle), Metaphire californica (triangle), Metaphire houlletti (square), Metaphire posthuma (star), and Diplodricodrilus schmardae (square); B) excavatus (circle) and Pheretima darnleiensis (triangle); C) Pheretima bicincta (square) (new record from Brazil) and Polypheretima elongata (square); D) Polypheretima taprobanae (square) and litoralis (circle) from South America.

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11. Metaphire posthuma (Vaillant, 1868) Genus Pheretima Kinberg, 1867 Perichaeta posthuma Vaillant, 1868: 228; Perichaeta affinis Type: Pheretima montana Kinberg, 1867 Perrier, 1872: 106, fig. 66; Megascolex posthumus: Vaillant, 14. Pheretima (Pheretima) darnleiensis (Fletcher, 1899: 72, figs. 12–13;Pheretima posthuma: Michaelsen, 1886) 1900: 295; Pheretima incerta Beddard, 1912: 197; Pheretima Perichaeta darnleiensis Fletcher, 1886; Perichaeta posthuma: Stephenson, 1923: 309 (syn. affinis). cingulata (part, cf. A. robustus); Vaillant, 1868; Perichaeta Metaphire posthuma: Sims and Easton, 1972: 239; Sims darnleiensis (sic) Fletcher, 1886; Perichaeta vaillanti and Gerard, 1985: 132, fig. 47d. Beddard, 1890; Perichaeta martensi Michaelsen, 1892; Distribution: ARG: Buenos Aires Prov.: Buenos Aires Perichaeta bosschae Horst, 1893; Perichaeta martensi javana (Mischis, 2004; 2007) (Figure 3A). Michaelsen, 1894; Perichaeta darnleiensis: Beddard, 1895; Further distribution: Bahamas (Michaelsen, 1900a), Perichaeta padasensis Beddard & Fedarb, 1895; ?Perichaeta Mexico (Fragoso and Brown, 2007), Europe, Asia, Pacific atheca Rosa, 1896; Perichaeta eoa Rosa, 1896; Perichaeta Ocean, and Christmas Island (Michaelsen, 1900a). floweri Benham 1897; Perichaeta madelinae Benham Genus Duplodicodrilus Blakemore, 2008 1897; Perichaeta belli Rosa 1898; Amyntas padasensis Type: Duplodicodrilus schmardae (Horst, 1883) lokonensis Michaelsen, 1899; Pheretima decipiens Beddard, (Blakemore, 2008, 2010) 1912; Pheretima benguetensis Beddard, 1912; Pheretima 12. Duplodicodrilus schmardae (Horst, 1883) kuchingensis Stephenson, 1916; Pheretima borneoensis Ude, Megascolex schmardae Horst, 1883: 194; ?Perichaeta 1925; Pheretima indica birangi Michaelsen, 1928; Pheretima sumatrana: Beddard, 1892a: 155, Pl. IX, fig. 4; Perichaeta pugnatoris Hong & James, 2010; Pheretima tabukensis Hong trityphla Beddard, 1896: 205; ?Perichaeta vesiculata Goto & James, 2010. & Hatai, 1899: 21, figs. 13–15;Amyntas schmardae [sic]: Distribution: BRA: RS (Righi, 1965, 1980; Righi and Michaelsen, 1899: 13; Pheretima schmardae: Michaelsen, Knäpper, 1965): Porto Alegre (Knäpper, 1972a, 1972b); 1900: 302 (syn. trityphla); Pheretima kikuchii Hatai & PR (Chang, 1997; Sautter et al., 2006): Curitiba (Brown et Ohfuchi, 1936: 767; Metaphire schmardae schmardae: al., 2006); SP (Lüderwaldt, 1927; Lenko, 1972; Caballero, Sims & Easton, 1972: 239 (schmardae-group); Metaphire 1976; Brown and James, 2006): Campos do Jordão; São schmardae: Easton, 1981: 58. Sebastião; São Paulo; São José do Rio Preto; Engenheiro Duplodicodrilus schmardae; Blakemore, 2008: 22. Marsilac; Salesópolis (Brown et al., 2006); MG: Conceição Distribution: BRA (Righi, 1967b; Hauser et al., 1975): do Mato Dentro; Tripuí; near Ouro Preto (Righi, 1967b) RS (Knäpper, 1972b; Knäpper and Porto, 1979): Porto (Figure 3B). Alegre: Guaíba estuary (Knäpper, 1976); São Leopoldo Further distribution: Indo-Pacific (Blakemore, 2002, (Brown et al., 2006); Estância Velha; Canoas (Knäpper, 2010). 1972a); SC: Blumenau; Pomerode (Lüderwaldt, 1927); PR Genus Pithemera Sims & Easton, 1972 (Chang, 1997; Sautter et al., 2006): Curitiba (Brown et al., Type: Perichaeta bicincta Perrier, 1875 (= P. bicincta) 2006); SP (Brown and James, 2006): Cotia (Brown et al., 15. Pithemera bicincta (Perrier, 1875) 2006); São Paulo: Colina; RJ: Itatiaia National Park (Righi, Perichaeta bicincta Perrier, 1875; ?Perichaeta violacea 1980); Teresópolis (Michaelsen, 1927) (Figure 3A). Beddard, 1895a: 407; ?Perichaeta violacea Beddard, 1895; Further distribution: Central America (Righi, 1980), ?Pheretima aimerikiensis Ohfuchi, 1941; Pheretima bicincta: Europe, and Asia (Blakemore, 2008, 2010). Michaelsen, 1935b: 107. Genus Perionyx Perrier, 1872 Distribution: ARG (Fragoso and Brown, 2007). Type: Perrier, 1872 New record: Campina Grande, Paraíba, Northeast 13. Perionyx excavatus Perrier, 1872 Brazil. Perionyx gruenewaldi Michaelsen, 1891; Perionyx Further distribution: Asia, Pacific Islands (Blakemore, intermedius Beddard, 1892; Perionyx sp. Friend, 1911; 2002), Central America, Mexico (Fragoso and Brown, ?Perionyx koboensis Stephenson, 1914; Perionyx parvulus 2007), Madeira, Canary Islands (Talavera, 1990), Hungary Stephenson, 1916; Perionyx fulvus Stephenson, 1916; (Csuzdi et al., 2008), London and Dublin (Blakemore, ?Perionyx shillongensis Stephenson, 1920; ?Perionyx turaensis 2010). Stephenson, 1920. Examined material: 8 adult specimens collected Distribution: COL: Alcalá; Valle del Cauca Dep.: on 28 April 2009 in an urban area located in Campina Cartago (Feijoo, 2007) (Figure 3B). Grande (7°13′28.77″S, 35°52′2.74″W), State of Paraíba, Further distribution: Central America and Mexico northeastern Brazil (UFPB 1461) (Figure 3C). (Fragoso and Brown, 2007), Asia, Hawaii, Madagascar, Description: Length: 48–51 mm. Diameter: 2–2.5 Pacific, UK, USA, Australia, and (Blakemore, mm. Segments: 78–102. Prostomium: open epilobous. 1994, 2002). First dorsal pore: 12/13. Setae: perichaetine. Nephropores:

607 DE ASSIS et al. / Turk J Zool absent. Clitellum: 14(1/2)–16, annular, darker than and 4C). Nephridia: meroic. Gizzard: a single one in 8–10. the rest of the earthworm. Spermathecal pores: 5 pairs Hearts: 10–12. Calciferous glands: absent. Ovaries: a pair in 4/5/6/7/8/9. Female pores: closely paired on 14. in 13. Testes: 10 and 11, seminal vesicles paired in 11 and Male pores: a pair on conical porophores in 18. Genital 12. Prostate glands: opening in 18 and extending into 16– markings: absent (Figure 4A). Septa: 5/6–7/8 thickened, 19, racemose. Intestinal origin: 15, with a pair of ceca in 22 and 8/9 absent. Spermathecae: 5 pairs, 5–9 (Figures 4B (Figure 4D).

Figure 4. A) Pheretima bicincta, anterior body part in ventral view, spermathecal pores in 4/5/6/7/8/9, closely paired female pores on 14; B) dorsal view, 5 pairs of spermathecae in 5–9; C) spermatheca with a single diverticulum; D) 1 pair of ceca in 22.

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Remarks: Specimens of P. bicincta were found under Genus Pontodrilus Perrier, 1874 a grape tree sharing the same spot with Pontoscolex Type: Pontodrilus marionis Perrier, 1874 (= Lumbricus corethrurus (Müller, 1857) in the first 10-cm layer of soil. litoralis Grube, 1855) Genus Polypheretima Michaelsen, 1934 18. Pontodrilus litoralis (Grube, 1855) Type: Perichaeta stelleri Michaelsen, 1892 Lumbricus litoralis Grube, 1855: 127; ?[Pontoscolex 16. Polypheretima elongata (Perrier, 1872) arenicola Schmarda, 1861: 11 (?part); Pontodrilus marionis Perichaeta elongata Perrier, 1872: 124, fig. 70;Perichaeta Perrier, 1874: 1582; Pontodrilus bermudensis Beddard, biserialis Perrier, 1875: 1044; Megascolex elongata: Vaillant, 1891: 96; Cryptodrilus insularis Rosa, 1891: 387, fig. 1889: 81; Perichaeta acystis Beddard, 1895a: 423; Perichaeta 11 (of left prostate); Pontodrilus arenae Michaelsen, monocystis Horst, 1899: 202 (lapsus pro acystis Beddard, 1892: 222, fig. 9; Pontodrilus sp. Michaelsen, 1894: 184; 1895); Amynthas elongata: Beddard, 1900: 650; Pheretima Pontodrilus hesperidum Beddard, 1894: 37; Pontodrilus elongata: Michaelsen, 1900: 265; Metapheretima elongata: insularis: Beddard, 1895a: 471; Pontodrilus michaelseni Sims and Easton, 1972: 205 (syn. biserialis) in a species- Eisen, 1895: 73; Pontodrilus ephippiger Rosa, 1898: 277, complex. 281; Pontodrilus matsushimensis Iizuka, 1898: 21, Pl. II; Polypheretima elongata: Easton, 1979: 53; Easton, Pontodrilus ephippiger laysanianus Michaelsen, 1899: 217; 1981: 61 [syn. biserialis; non elongata: Ohfuchi, 1956 (= Pontodrilus matsushimensis chathamianus Michaelsen, Amynthas morrisi)]; Easton, 1984: 733. 1899: 220; Pontodrilus michaelseni hortensis Eisen, 1900: Distribution: PER (Perrier, 1872; Cernosvitov, 1934). 129, 241, figs.; Pontodrilus laccadivensis Beddard, 1903: COL: Valle Dep.: El Cerrito; Palmira; Guacarí (Feijoo, 374; Pontodrilus crosslandi Beddard, 1906: 561; Pontodrilus 2007); San Andrés Island (Reynolds and Reynolds, 2001). albanyensis Michaelsen, 1907a: 185; Pontodrilus VEN: Macuto, near Guayra (Cognetti, 1905); Maracay: bermudensis ephippiger: Stephenson, 1915: 145; Plutellus Experimental Station (Righi, 1989); Bolívar. FRG (Gates, (Pontodrilus) bermudensis: Michaelsen, 1922: 22; 1935: 1972). SUR: Paramaribo (Horst, 1899). GUY: Georgetown 106; Pontodrilus albanyensis var. cygni Jackson, 1931: 94; (Beddard and Feddarb, 1899); BRA: SP (Brown and Plutellus (Pontodrilus) matsushimensis indica Michaelsen, James, 2006; James and Brown, 2006): São Paulo: 1935: 106; Pontodrilus gracilis Gates, 1943: 100. Anhembi (Brown et al., 2006); RJ (Magalhães, 1905); MG: Pontodrilus litoralis: Beddard, 1895a: 469 (syn. Caetanópolis; Curvelo; Banheiro farm; Macacos waterfall; marionis); Michaelsen, 1900: 180 (syn. marionis); Capão Queimado farm (Righi, 1980); BA: Itajubá (Righi, Cognetti, 1901: 404; Michaelsen, 1910c: 89 (syn. marionis, 1971); PE: 50 km from Recife (Righi, 1980); RR (Lavelle crosslandi, albanyensis); Easton, 1984: 114, fig. 1f [syn. and Lapied, 2003) (Figure 3C). arenicola (part.), marionis, bermudensis, insularis, Further distribution: Central America (Gates, 1972), arenae, hesperidum, michaelseni, ephippiger, bermudensis Mexico (Fragoso and Brown, 2007), Africa, Asia, Australia, ephippiger.- Stephenson, 1915, matsushimensis, ephippiger and Pacific islands (Gates, 1972). laysanianus (syn. michaelseni hortensis), matsushimensis 17. Polypheretima taprobanae (Beddard, 1892) chathamianus, michaelseni hortensis, P. sp. Michaelsen, Perichaeta taprobanae Beddard, 1892a: 163; Perichaeta 1894, P. (P. ) sp. Michaelsen, 1930, laccadivensis, crosslandi, pauli Michaelsen, 1897: 234; Pheretima taprobanae typica: albanyensis, P. sp. Michaelsen, 1913, albanyensis cygni, Michaelsen, 1900: 308; Amynthas taprobanae: Beddard, matsushimensis indica, P. sp. Gates, 1936, gracilis]. 1900: 648 (syn. pauli); Pheretima taprobanae: Stephenson, Distribution: CHI: Galapagos Islands (Blakemore, 1923: 312; Metapheretima taprobanae: Sims and Easton, 2002). COL: Cartagena (Michaelsen, 1910). VEN: 1972: 233; Polypheretima taprobanae: Easton, 1979: 45 (syn. Morrocoy National Park: Playa Mero; Cayo Cico; Cayo pauli); Easton, 1982: 734. Sombrero (Easton, 1984; Righi, 1989; Zicsi, 1995a). BRA: Polypheretima taprobaena (laps.): Easton, 1979: 45. RS (Michaelsen, 1900b, 1910); SC: Florianópolis; São Distribution: ARG: Tucumán Prov.: Tucumán (Teisaire Francisco Island; SP (Lüderwaldt, 1927; Righi, 1967b; and Roldán, 1996; Mischis, 2004; Mischis and Herrera, Brown and James, 2006): Santos (Michaelsen, 1910); 2006). BRA (Moreira, 1903; Righi, 1965): RS (Righi and Itanhaém (Lüderwaldt, 1927); Cananéia, in mangrove Knäpper, 1965): Santa Cruz do Sul; São Leopoldo (Brown (Righi, 1968); RJ (Moreira, 1903); Angra dos Reis, in et al., 2006); Porto Alegre: Guaíba estuary (Knäpper, mangrove (Righi, 1968); PE: Itamaracá Island (Brown et 1976); SP: Piracicaba (Lüderwaldt, 1927; Righi, 1967b; al., 2006) (Figure 3D). Brown and James, 2006); RJ: Rio de Janeiro (Michaelsen, Further distribution: Atlantic, Indian, and Pacific 1927) (Figure 3D). oceans, fully marine (Easton 1984). Further distribution: Mexico (Fragoso and Brown, Remarks: Blakemore et al. (2012) added this species to 2007), Africa, Asia, Australia, and Pacific islands (Gates, Korean species biodiversity list as an exotic species from 1972), London (Blakemore, 2010, 2014). Korea.

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4. Discussion 2b. Esophageal glands unpaired in 9–11; paired None of the 18 species catalogued herein are endemic to calciferous glands in 13; several intestinal South America. Most of these species were transported gizzards...... Hyperiodrilus unintentionally, through soil or other materials that could 3a. Gizzard present...... 5 contain earthworms, probably soon after the establishment 3b. Gizzard absent or rudimentary...... 4 of trade routes between the Old and the New Worlds 4a. Prostates tubular; chaetae 8 per segment (i.e., (Brown et al., 2006; Blakemore, 2009). lumbricine); nephridia not holoic (absent from The habitat of Eudrilidae is probably the forests of anterior segments...... Pontodrilus central Africa. Eudrilus eugeniae has been introduced 4b. Prostate nontubular; holoic (i.e., 1 pair of nephridial in many other localities of the world. This species can tubules per segment); digestive system without ceca... only survive in temperatures above 10 °C and requires ...... Perionyx temperatures of 21–26 °C for (Sims and 5a. Intestinal ceca present...... 6 Gerard, 1985). Hyperiodrilus africanus is also considered 5b. Intestinal ceca absent; crescent markings an introduced species by Michaelsen (1910). It is common absent...... Polypheretima in western Africa, and its type species is from Kew Gardens. 6a. Intestinal ceca originating in or near 27...... 7 Megascolecid earthworms are widely distributed in 6b. Intestinal ceca originating in or near 22...... Pithemera South America and their presence on this continent most 7a. Male pores nonsuperficial within lateral apertures of of the time indicates disturbed by human activities. copulatory pouches...... 8 Amynthas gracilis is by far the most common species in 7b. Male pores superficial discharging directly onto the many areas. It has been known for over a century on this surface...... Amynthas continent, and it was first described for Brazil in 1867 by 8a. Spermathecal duct free of nephridia...... 9 Kinberg (Brown et al., 2006). The species listed herein represent threats to our 8b. Nephridia on spermathecal ducts...... Pheretima native species, leading to a loss in the diversity of local 9a. Large, eversible, and doubled intromittent organs earthworms. present...... Diplodricodrilus 9b. Intromittent organs absent, male pores not Taxonomic key to the genera of Eudrilidae and developed...... Metaphire Megascolecidae from South America. Adapted from Blakemore (2010). Acknowledgments 1a. Male pore on 17; spermathecae absent from The authors heartily thank Dr Robert J Blakemore pretesticular segments; as coelomic cavities (Biodiversity Laboratory, College of Natural Science, combining with ovaries (“euprostate” present, holoic, Hanyang University, Seoul, South Korea) and Alexander setae lumbricine)...... Eudrilidae...... 2 Feijoo M (Universidad Tecnológica de Pereira, Apartado 1b. Male pore on 18; spermathecae present in pretesticular Aéreo 97, Pereira, Colombia) for valuable contributions segments (prostates tubular to racemose), to the manuscript. Many thanks to Jürgen Römbke for nephridia holoic or meroic; setae lumbricine to providing us an important reference on earthworms. perichaetine...... Megascolecidae...... 3 Finally, we acknowledge CNPq and FACEPE for the 2a. Esophageal glands unpaired in 10 and 11, paired postdoctoral fellowship to JE De Assis and a productivity calciferous glands in 12; single esophageal grant to ML Christoffersen (CNPq Proc. N. 302388/2014- gizzard...... Eudrilus 1).

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