Annelida, Clitellata, Lumbricina, Glossoscolecidae) in South America1∗
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Gaia Scientia 2008, 2(2): p-21 - p-30 Diversity and distribution of Aptodrilus, Glossodrilus, Onoreodrilus and Righiodrilus (Annelida, Clitellata, Lumbricina, Glossoscolecidae) in South America1∗ Martin L. CHRISTOFFERSEN2 Abstract A catalogue of 87 species and subspecies of terrestrial megadrile Glossoscolecidae from South America, belonging to Aptodrilus (11 taxa), Glossodrilus (48 taxa), Onoreodrilus (5 taxa), and Righiodrilus (23 taxa), is presented. Gilberto Righi described 39 of these taxa, predominantly from Brazil, while Andras Zicsi described another 34 taxa, mainly from Ecuador. These genera represent tropical groups of South American endemic species, with the exception of only two species (Glossodrilus nemoralis and G. peregrinus), also found in Central America and the Caribbean. Full synonyms and detailed South American occurrences are provided. KEY WORDS: biodiversity, earthworms, Glossoscolecoidea, Neotropical region. Resumo DIVERSIDADE E DISTRIBUIÇÃO DE APTODRILUS, GLOSSODRILUS, ONOREODRILUS E RIGHIODRILUS (ANNELIDA, CLITELLATA, LUMBRICINA, GLOSSOSCOLECIDAE) NA AMÉRICA DO SUL. Um catálogo de 87 espécies e subespécies de minhocas terrestres da América do Sul, pertencentes aos gêneros Aptodrilus (11 táxons), Glossodrilus (48 táxons), Onoreodrilus (5 táxons) e Righiodrilus (23 táxons), é apresentado. Gilberto Righi descreveu 39 destes táxons, predominantemente do Brasil, enquanto András Zicsi descreveu outros 34 táxons, predominantemente do Equador. Estes gêneros representam grupos tropicais de espécies endêmicas, com apenas duas espécies (Glossodrilus nemoralis e G. peregrinus), encontradas também no Caribe e na América Central. São fornecidas sinonímias completas e ocorrências detalhadas na América do Sul para cada táxon. PALAVRAS-CHAVE: biodiversidade, Glossoscolecoidea, minhocas, região Neotropical. Introduction Progress in our knowledge of glossoscolecids in Glossodrilus + Righiodrilus appear to share a unique pair South America must begin by cataloguing our present of compound calciferous glands on segment 11, Righiodrilus knowledge of this very diverse and highly endemic family differing further by the presence of membranes in these in the Neotropical region. Published data are scattered calciferous glands (Zicsi, 1995) and Glossodrilus by having in numerous small publications in diverse languages, and tubular calciferous glands which have (always?) a reservoir would become more useful and accessible if compiled into (Righi, 1990). a single English publication. On the other hand, Aptodrilus and Onoreodrilus share In this paper I provide a catalogue of the species the presence of 5 pairs of lateral oesophageal calciferous and subspecies of Aptodrilus, Glossodrilus, Onoreodrilus and glands on segments 10-14 (Zicsi, 1990, 1997). Righiodrilus cited to date from the South American continent. Hopefully this effort will serve as a starting point for Fragoso & Brown (2007) recently listed a total of 94 species ecologists and taxonomists to further sample these endemic for these genera in Latin America. South American genera, study species interrelationships, The four genera share 8 pairs of chaetae per segment, and promote efforts to protect the endangered habitats in disposed in longitudinal series, have one pair of intraclitellar which endemic species may still be found. Compiling the male pores, and are metandric and metagynic (Righi, 1990; available information on the biodiversity and distribution Zicsi, 1990, 1995, 1997). It is difficult to decide at our of a neglected zoological group in a megadiverse and present state of knowledge if any of these characters may only cursorily sampled continent remains a priority for be apomorphic. those interested in biodiversity studies and conservation biology. 1I dedicate this paper to the memory of Gilberto Righi. 2 Departamento de Sistemática e Ecologia, Universidade Federal da Paraíba, 58059-900, João Pessoa, Paraíba, Brazil; [email protected] 22 M. L. Christoffersen Material and methods Distr: ECU: Azuay Prov: Cuenca and Gualaceo; Cañar Prov: Paredones (Cognetti, 1904: 15); Chimborazo I have used 3-letter abbreviations for South American Prov; Cotopaxi Prov; Pichincha Prov: near Quito and countries (ARG, Argentina; BOL, Bolivia; BRA, Brazil; Santo Domingo (Zicsi & Csuzdi, 1988: 216); Carchi Prov; COL, Colombia; ECU, Ecuador; GUY, Guyana; PAR, Cotopaxi Prov; Tungurahua Prov; Boliva Prov; Chimborazo Paraguay; PER, Peru; VEN, Venezuela) and 2-letter Prov. (Zicsi, 1995: 101). abbreviations for sampled States in Brazil (AM, Amazonas; AP, Amapá; MA, Maranhão; MT, Mato Grosso; PA, Pará; 5*. A. fuhrmanni Michaelsen, 1913 RJ, Rio de Janeiro; RO, Rondônia; RR, Roraima; RS, Rio Previous name: Thamnodrilus fuhrmanni (Michaelsen, Grande do Sul; SP, São Paulo). 1913). Syn: Rhinodrilus (Aptodrilus) uncinatus var. Michaelsen, Results 1913. Distr: ECU: Pichincha Prov: near Quito and Santo Species marked with an asterisk (*) are previously only Domingo; Cotopaxi Prov: San Francisco de las Pampas, reported from South America. Distr. = Distribution; Syn. Ebenda, and Maranchito; Chimborazo Prov: Pohaluin, San = Synonymy. Juan Triglona, and close to Juan de Valesco, Wiese, and LUMBRICINA De Blainville, 1830 Paramo; Bolivas Prov: Las 4-Esquinas; Bolivar Prov: Cashca Syn: Terricolae Örsted, 1850; Megadrili Benham, 1890; Totoraz (Zicsi & Csuzdi, 1988: 212); Napo Prov. (Zicsi, Diplotesticulata Yamaguchi, 1953; Crassiclitellata Jamieson, 1997: 167). COL: Finca Camelia, near Titribi (Michaelsen, 1988 1914: 204); Cauca valley (Zicsi, 1997: 167); Valle Dep: GLOSSOSCOLECOIDEA Michaelsen, 1900 Palmira (Feijoo, 1993; 2007); Antioquia Dep: Angelopolis, GLOSSOSCOLECIDAE Michaelsen, 1900 to the south of Medellin (Michaelsen, 1918: 159). Glossodrilus genus-group Aptodrilus Cognetti, 1904 6*. A. griseus Zicsi, 1997 Type: A. excelsus Cognetti, 1904 Distr: ECU: Bolívar Prov: Cascha-Totóras (Zicsi, 1997: Syn: Rhinodrilus (Aptodrilus) Cognetti, 1904 162). 1*. A. baloghi (Zicsi & Csuzdi, 1988) 7*. A. lacteus Zicsi, 1997 Syn: Thamnodrilus baloghi Zicsi & Czusdi, 1988; Distr: ECU: Cañar Prov; Cotopaxi Prov. (Zicsi, 1997: Glossodrilus baloghi (Zicsi & Csuzdi, 1988: 212). 159). Distr: ECU: Cotopaxi Prov: San Francisco de las 8*. A. locksai Zicsi, 1997 Pampas; near Quito and Santo Domingo; Pichincha Prov: Syn: Glossodrilus loksai (Zicsi, 1989). Saloya; 32 km from Selva Alegre (Zicsi & Csuzdi, 1988: Distr: ECU: Pichincha Prov: Loa; Imbabura Prov. 214); Napo Prov: Puerto Misahuali; Pataza Prov: Puyo; (Zicsi, 1997: 163): Tungurahua Prov: Pilaguin; Chimborazo Cañar Prov; Azuay Prov; Imbabura Prov. (Zicsi, 1988a: Prov: Juan de Valesco; Chimborazo; Cotopaxi Prov: Pujili; 318); Sucumbios Prov. (Zicsi, 1997: 158). Cotoxapi (Zicsi, 1989: 172). 2*. A. botari (Zicsi & Csuzdi, 1988) 9*. A. ruvidus Cognetti, 1904 Syn: Thamnodrilus botari Zicsi & Csuzdi, 1988. Syn: Thamnodrilus ruvidus (Cognetti, 1904). Distr: ECU: Chimborazo Prov. (Zicsi & Csuzdi, 1988: Distr: ECU: Carchi Prov: Tulcan (Cognetti, 1904: 16). 214); Cotopaxi Prov: Carchi: Chiles volcano (Zicsi, 1997: 166). 10*. A. tandajapaensis Zicsi, 1997 Distr: ECU: Pichincha Prov: Tandajapa (Zicsi, 1997: 3*. A. chilensensis Zicsi, 1997 160). Distr: ECU: Carchi Prov: Chiles volcano (Zicsi, 1997: 166). 11*. A. uncinatus (Michaelsen, 1910) Syn: Rhinodrilus (Aptodrilus) uncinatus Michaelsen, 1910; 4*. A. excelsus (Cognetti, 1904) Thamnodrilus uncinatus (Michaelsen, 1910); ?Rhinodrilus Syn: Glossodrilus excelsus Cognetti, 1904; Rhinodrilus (Aptodrilus) uncinatus var. Michaelsen, 1913. (Aptodrilus) excelsus (Cognetti, 1904); Thamnodrilus excelsus Distr: ECU (Michaelsen, 1910: 135); Cotopaxi Prov; (Cognetti, 1904); Andioscolex excelsus (Cognetti, 1904). Pichincha Prov; Chimborazo Prov. (Zicsi & Csuzdi, 1988: Diversity and distribution of some Glossodrilus in South America 23 218); Carchi Prov; Imbabura Prov; Sucumbios Prov. (Zicsi, Distr: PER: Loreto Prov: Panguana ecological station. 1997: 166). COL: Valle Dep: Palmira (Feijoo, 2008). ECU: Pichincha Prov; Cotopaxi Prov. (Righi & Römbke, 1987: 532); Tungurahua Prov. (Zicsi, 1995: 102). Glossodrilus Cognetti, 1905 Type: G. parvus Cognetti, 1905 21*. G. dudichi Zicsi, 1995 Syn: Andioscolex Michaelsen, 1927 (Type: Tykonus Distr: ECU: Pichincha Prov. (Zicsi, 1995: 98). peregrinus Michaelsen, 1897); G. (Tonperoge) Righi & Ayres, 1975 (Type: G. parecis Righi & Ayres, 1975) 22*. G. excelsus (Cognetti, 1904) Syn: Glossoscolex excelsus Cognetti, 1904. 12*. G. antisanae Zicsi, 1989 Distr: ECU: Azuay Prov: Cuenca; Cañar Prov: Gualaceo; Distr: ECU: Pichincha Prov: Antisana; Nono; Pifo; Paredones (Cognetti, 1904: 16); Bolivar Prov; Chimborazo Papallacta; Pasochoa National Park (Zicsi, 1989: 181); Prov. (Zicsi, 1995: 100). Cotopaxi Prov. (Zicsi, 1995: 94). 23*. G. fragilis Zicsi, 1989 13*. G. antunesi (Righi, 1971) Syn: G. crucifer subsp. fragilis Zicsi, 1989. Syn: Andioscolex antunesi Righi, 1971; G. baiuca Hamoui Distr: ECU: Pichincha Prov: Pichincha; Pasochoa & Donatelli, 1983. National Park; Chimborazo Prov: Chimborazo (Zicsi, Distr: BRA: MA: Caxias Mun: margin of Brocos lake 1989: 178); Imbabura Prov; Tungurahua Prov; Loja Prov. (Righi, 1972: 161); AP: Serra do Navio; Capivara waterfall; (Zicsi, 1995: 102). middle Amaparí river (Righi, 1971: 58); RR (Hamoui & Donatelli, 1983: 143): Maracá island; Bonfim (Righi, 1991: 24*. G. griseus Zicsi & Feijoo, 1994 392). Distr: COL: Valle Dep: Palmira (Feijoo, 1993; 2007); El Cerrito: Páramo de Los Dominguez; Florida: Páramo 14*. G. baloghi Zicsi, 1998 de Tinajes (Zicsi & Feijoo, 1994: 55). Distr: