Neotropical Biology and Conservation 3(1):39-48, january - april 2008 © by Unisinos

A Catalogue of the Piscicolidae, Ozobranchidae, and Arhynchobdellida (Annelida, Clitellata, Hirudinea) from South America

Catálogo das famílias Piscicolidae, Ozobranchidae e Arhynchobdellida (Annelida, Clitellata, Hirudinea) da América do Sul

Martin Lindsey Christoffersen1 [email protected]

Abstract

A catalogue of the families Americobdellidae, Cyclobdellidae, Cylicobdellidae, Erpobdellidae, Euhirudinea, Hirudinidae, Macrobdellidae, Ozobranchidae, Piscicolidae, Salifidae, Semiscolecidae, and Xerobdellidae produced 66 nominal species reported to date from South America. 86% of this fauna is endemic to the South American continent. Synonyms and detailed South American occurrences are provided. This is the first reassessment of South American leeches other than Glossiphoniidae in 26 years. An inventory of this little- studied group may be important from an applied point of view. Leeches have very high endemicity rates in South America, perhaps unparalleled in freshwater environments. Arguably these animals could serve as important model organisms for ecological studies, particularly as environmental indicators.

Key words: biodiversity, environmental indicators, leeches, Neotropical region.

Resumo

Um catálogo das famílias de Americobdellidae, Cyclobdellidae, Cylicobdellidae, Erpobdellidae, Euhirudinea, Hirudinidae, Macrobdellidae, Ozobranchidae, Piscicolidae, Salifidae, Semiscolecidae e Xerobdellidae produziu 66 espécies nominais assinalados até o momento para a América do Sul. Do total, 86% desta fauna é endêmica para o continente sul-americano. Sinonímias e ocorrências detalhadas para a América do Sul são fornecidas. Esta é a primeira reavaliação da fauna de hirudíneos não-glossifonídeos da América do Sul em 26 anos. Um inventário deste grupo pouco estudado poderá ser importante de um ponto de vista aplicado. Sanguessugas têm um índice muito alto de endemismo na Améri- ca do Sul, talvez sem paralelo em ambientes aquáticos. Estes organismos poderiam servir como um importante modelo para estudos ecológicos de ambientes aquáticos, particular- 1 Departamento de Sistemática e Ecolo- mente como indicadores de qualidade ambiental. gia, Universidade Federal da Paraíba, Campus I, 58059-900, João Pessoa, Pa- raíba, Brazil. Palavras-chave: biodiversidade, indicadores ambientais, sanguessugas, região Neotropical.

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Introduction difficult, because even very inconspi- leeches that prey mainly on inverte- cuous external characters may someti- brates, swallowing and smashing them The diverisity and endemism of neo- mes be reliable for species separation, with a long muscular pharynx (Tron- tropical hirudineans, which are a result while some anatomical characters telj et al., 1999, p. 141), having aban- of limited dispersal range of the majo- appear to be highly variable within the doned the blood feeding habits of their rity of species, is well known (Moore, species and sometimes even among ancestors (Siddall, 2002, p. 1). San- 1911; Siddall and Borda, 2004, p. 14). individuals (Sket, 1968). guivorous hirudinids feed primarily on South America is especially important Arhynchobdellidan evolution involved blood, assisted by their toothed jaws because of the large number of ende- a move to land. They contain large, (Trontelj et al., 1999, p. 141). They mic species. aquatic hirudinids with serrate muscu- make tripartite incisions in the skin of Monotypic South American Americo- lar jaws, or smaller terrestrial blood- their hosts, from which they draw on bella lies at the base of the erpobdellid feeding groups. Many arhynchobdelli- the upwelling blood. Hirudo medici- leeches, being an example of the im- dans possess multiple eyespots for 3- nalis sucks blood directly through ex- portance of South America in leech dimensional vision. The eggs of arrhyn- ternal mammalian skin, usually until evolution and diversification (Siddall chobdellidans (Erpobdella and Hirudo) gorged, and then drops off from its et al., 2001, p. 346). The widespread are small (circa 100 μm), and their de- host. The blood ingested by starved Asian Barbronia weberi, on the other velopment and gastrulation undergoes leeches amounts from six to nine ti- hand, is an example of an accidental modifications parallel to those of me- mes their weight in one meal. Occa- introduction of leeches. This species gadrile eggs in relation to microdriles sionally, H. medicinalis invades the feeds predominantly on chironomid (Omodeo, 1998, p. 63), a clear evolu- nostrils of its mammalian host. This larvae (Siddall and Budinoff, 2005, p. tionary response to a terrestrial exis- species has been used in medicine 470). The mostly New World ´medical tence. Arhynchobdellidans deposit co- since ancient times (Orevi et al., leeches´, Macrobdellidae and Semisco- coons with a hard surface, inherited 2000, p. 122). No fewer than 7 medi- lescidae, originated about 85 Mya ago, from their oligochaete ancestors. Ne- cally useful substances have already with the opening of the South Atlantic vertheless, in Erpobdella punctata, a been identified from the medical le- (Phillips and Siddall, 2005, p. 559). large proportion of the unprotected co- ech: hirudin, preventing coagulation Ringuelet (1981a, p. 192) lists Ichthyo- coons were destroyed by predators, of ingested blood, an anaesthesic, a bdlla (?) cichlae Diesing, 1850 for Rio primarily by snails (Sawyer, 1971, p. spreading factor, an anti-plasma, an de Janeiro, Brazil. However, Ichthyo- 202). The habit of cementing the pro- anti-trypsin, an anti-inflammatory, bdella De Blainville, 1827 is a sy- tective cocoon to a substrate of some and an anti-biotic (Sawyer et al., nonym of Piscicola De Blainville, 1818 sort appears to be the original state in 1982, p. 412). (Pinto, 1923, p. 902). This species is leech evolution, exibited in acanthob- In this paper I provide a catalogue of not listed in later revisions of Sawyer dellidans and branchiobdellidans, and the piscicolids, ozobranchids, and (1986) and Epshtein et al., 1994). Sin- retained in piscicolids and erpobdelli- arhynchobdellidans cited to date from ce the efforts of Ringuelet (1981a), no ds alike (Siddall and Burreson, 1996, the South American continent. The re- species inventories of the leeches and p. 282). The aquatic habits of the erpo- maining leeches of South America, their allies from South America have bdellids appear to represent a secon- belonging to the Glossiphoniidae, have been attempted. dary return to water. Those groups that been treated in another paper. Piscicolidae and Ozobranchidae form return to water retain some aspects of the sister group to the Arhynchobdelli- terrestrialism, particularly those invol- Material and methods da (Apakupakul et al., 1999, p. 356). ved in cocoon deposition and early The ozobranchids are distinguished by ontogeny. The terrestrial deposition of I have used 3-letter abbreviations for the presence of lateral digitiform bran- cocoons in Hirudinidae is interpreted South American countries (ARG, Ar- chiae (MacCallum and MacCallum, as a means to reduce the destruction of gentina; BOL, Bolivia; BRA, Brazil; 1918), being the only non-piscicolid the offspring by aquatic predators such CHI, Chile; COL, Colombia; ECU, leeches to live in the sea, but they are as snails, insect larvae, and fish (Kuts- Ecuador; FRG, French Guyana; PAR, unique in being permanent ectoparasi- chera and Wirtz, 2001, p. 132). Paraguay; PER, Peru; URU, Uruguay; tes of sea turtles (Sawyer, 1986, p. 700). A dominant feature of the South Ame- VEN, Venezuela) and 2-letter abbrevi- Among the arrhynchobdellidans, the rican leach fauna is the presence of ter- ations for sampled States in Brazil Erpobdellidae have long been used as restrial predacious forms, virtually la- (AM, Amazonas; ES, Espírito Santo; model organisms for ecological studi- cking in Africa. Semiscolex represents GO (Goiás); MG, Minas Gerais; PA, es and species interactions, and as in- a totally predaceous species (Sawyer, Pará; PB, Paraíba; RJ, Rio de Janeiro; dicators for freshwater toxicology (Si- 1986, p. 701). Erpobdellidae is a group RS, Rio Grande do Sul; SC, Santa Ca- ddall, 2002, p. 1). Their taxonomy is of non-sanguivorous macrophagous tarina; SP, São Paulo.

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Results Distribution. BRA (Pinto, 1923, p. Distribution. S. America (Cordero, 1011). 1937a, p. 16). ARG (Ringuelet, 1968, Species marked with an asterisk (*) are Further distribution. Arctic seas (Sa- p. 378). previously only reported from South wyer, 1986, p. 676). America. A systematic index is provi- Pontobdella Leach, 1915 ded in Appendix 1. Myzobdella Leidy, 1851 Type. Pontobdella muricata Linnaeus, Type: M. lugubris Leidy, 1851 1758 (= P. verrucosa Fleming, 1811) Order RHYNCHOBDELLIDA R. Syn: Illinobdella Meyer, 1940 Synonym. Albione Savigný, 1820 Blanchard, 1894 *M. platense (Cordero, 1933) *P. variegata Baird, 1859 PISCICOLIDAE Johnston, 1865 Original name. Piscicola platensis Distribution. ARG: Patagonia (Weber, Syn: Ichthyobdellidae Leuckart, 1863 Cordero, 1933. 1915, p. 21). Distribution. URU: Rio de la Plata, on Bathybdella Burreson, 1981 Hoplias malabaricus (Bloch) (Pisces) *P. zonata Apáthy, 1905 Type: B. sawyeri Burreson, 1981) (Cordero, 1933, p. 450); on fish (Rin- Distribution. CHI: Calbuco: Ancud guelet, 1968, p. 378; Volonterio et al., gulf (Weber, 1915, p. 20). *B. sawyeri Burreson, 1981 2004, p. 21). Distribution. ECU: Galapagos rift, Stibarobdella Leigh-Sharpe, 1925 2447 m deep (Burreson, 1981, p. 486; *M. uruguayensis Mañé-Garzón and Type. Pontobdella macrothela Sch- Burreson and Segonzac, 2006, p. 15). Monteiro, 1977 marda, 1861 Distribution. URU, on fish (Volonte- Synonym. Pontobdellina Harding and Branchellion Savigný, 1822 rio et al., 2004, p. 21). Moore, 1927 (part); Pentabdella Type: B. torpedini Savigný, 1822 Remarks. This species parasites the Llewellyn, 1966 gills of the yellow catfish Rhamdia B. lobata Moore, 1952 sapo (Vallenciennes, 1840) (Mané- *S. dispar (Cordero, 1937) Distribution. CHI (Sawyer, 1986, p. Garzón and Monteiro, 1977, p. 59). Original name. Pontobdella dispar 675). Cordero, 1937. Further distribution. Eastern Pacific Platybdella Malm, 1863 Distribution. BRA (Cordero, 1937a, p. 13). up to California (Sawyer, 1986, p. 675). Type. P. anarrhichae (Diesing, 1859) Synonym. Crangonobdella Selensky, *S. loricata (Harding, 1924) *B. gnesios Ringuelet, 1985 1914; Sanguinothus De Silva and Bur- Original name. Pontobdella loricata Distribution. CHI, host Myliobatis don-Jones, 1961 Harding, 1924. chilensis Philippi (Pisces) (Ringuelet, Distribution. BRA: South, parasitic on 1985, p. 168). *P. chilensis Moore, 1910 Angel shark (Soto, 2003, p. 691). Previous name. Cryobdella chilensis B. parkeri Richardson, 1949 (Moore, 1910). *S. macrothela (Schmarda, 1861) Synonym. B. callorhynchus Szidat, Distribution. CHI (Sawyer, 1986, p. Original name. Pontobdella macro- 1972. 663). thela Schmarda, 1861. Distribution. ARG: Buenos Aires Remarks. The species was collected Distribution. S. Amer. (Cordero, 1937a, Prov: Punta Redonda, piscean host in Valparaíso, on the catfish Aphos po- p. 5). BRA: SC: parasitic on the whaler (Tanzola and Botté, 1998, p. 65). rosus (Vallenciennes, 1837) (Moore, shark (Pisces) (Soto, 2000, p. 713). Further distribution. Australia and 1910, p. 28). New Zealand (Sawyer, 1986, p. 675). *S. planodiscus Baird, 1859 P. patagonica Ringuelet, 1945 Original name. Pontobdella planodis- *B. ravenelii (Giard, 1851) Previous name. Cryobdella patagoni- cus Baird, 1859. Distribution. CHI: Concepcion bay, pis- ca (Ringuelet, 1945). Distribution. ARG: Patagonia (Weber, cean host (Morillas et al., 1987, p. 342). Distribution. ARG (Ringuelet 1945, p. 1915, p. 21). 109). Johanssonia Selensky, 1914 Further distribution. Sandwich Islan- Trachelobdella Diesing, 1850 Type: J. kolaensis Selensky, 1914 ds (Sawyer, 1986, p. 663). Type. T. mulleri Diesing, 1850

J. arctica (Johansson, 1898) *P. tentaculata (Cordero, 1937) *T. australis R. Blanchard, 1900 Synonym. Ichthyobdella pantopodum Original name. Ichthyobdella tentacu- Distribution. ARG: Ushuaia, Tierra Selensky, 1914. lata Cordero, 1937. del Fuego (Weber, 1915, p. 23).

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*T. mulleri Diesing, 1850 Americobdella Caballero, 1956 Previous names. Helobdella adaio- Synonym. Tracheobdella kollari Die- Type. Macrobdella valdivbiana De phthalma (Ringuelet, 1980); Bibula sing, 1850. Philippi, 1872 adaiophthalma (Ringuelet, 1980). Distribution. BRA (Pinto, 1923, p. 1023). Synonym. Cardea R. Blanchard, 1917; Distribution. COL (Ringuelet, 1980, Macrobdella De Philippi, 1872 (non p. 6). OZOBRANCHIDAE Pinto, 1921 Verrill,1872); Philippia Apáthy, 1905 (non Gray, 1840, mollusk; non Signo- Blanchardiella biolleyi Dequal, 1916 Bogabdella Richardson, 1969 ret, 1869, hemipteran). Original name. Helobdella biolleyi Type. Bogabdella diversa Richardson, (Dequal, 1916). 1969 *A. valdiviana (De Philippi, 1872) Distribution. ECU: Pun (Ringuelet, Synonym. Colombobdella Mañé-Gar- Previous names. Macrobdella valdi- 1981a, p. 193). zón, 1973 viana De Philippi, 1872; Philippia val- Further distribution. Costa Rica (Rin- diviana (De Philippi, 1872); Cardea guelet, 1981a, p. 193). *B. ringueleti (Mané Garzón, 1973) valdiviana (De Philippi, 1872). Original name. Colombobdella rin- Distribution. CHI (Moore, 1931, p. *Blanchardiella cameliae M. Weber, gueleti Mané-Garzón, 1973. 1220): Corral (Weber, 1915, p. 112); 1913 Distribution. COL: Meta Dep: Villa- Valdivia (Pinto, 1923, p. 1078); Parque Previous names and synonym. Helo- vicenso Mun: Peralonso: Pavitos dam, Oncol, near Santiago, 495 m (Siddall bdella cameliae (M. Weber, 1913); on the tortoise Podocnemis vogli Wa- and Borda, 2004, p. 3). Blanchardiella bogotaensis M. Weber, gler (Mané-Garzón, 1973, p. 129). 1913; Bibula cameliae (M. Weber, CYCLOBDELLIDAE Ringuelet, 1972 1913). Ozobranchus De Quatrefages, 1852 Distribution. COL: Cundinamarca: Type. O. branchiatus (Menzies, 1791) Orchibdella Ringuelet, 1945 Bogotá; Tolima: La Camelia coffee Synonym. Lophobdella Poirer and Ro- Type. O. pampeana Ringuelet, 1945 plantation; Popayán: Tambo (Weber, chebrune, 1884 1915, p. 120). *O. diaguita Ringuelet, 1978 O. margoi (Apáthy, 1890) Distribution. ARG: Subandean regi- *Blanchardiella columbiensis M. We- Original name. Pseudobranchellion on (Ringuelet, 1978, p. 264). ber, 1913 margoi Apáthy, 1890. Previous names. Dacnobdella colum- Distribution. ARG: La Plata, on *O. pampeana Ringuelet, 1945 biensis (M. Weber, 1913); Macrobdella dolphin Pontoporia blainvillei (Ger- Distribution. ARG: Subtropical and colombiensis M. Weber, 1913; Hypso- vais and D´Orbigný) (Soto, 2001, p. pampean domains (Ringuelet, 1945, p. bdella columbiensis (M. Weber, 1913); 173). 122; 1949, p. 155; 1976a, p. 101): Rio Bibula columbiensis (M. Weber, 1913). Further distribution. Italy, parasiti- de la Plata (Paggi et al., 2006, p. 5). Distribution. COL (Weber, 1913, p. zing turtle Thalassochelis corticata 731). (Pinto, 1923, p. 1093). *O. peruviensis Ringuelet, 1976 Distribution. PER: Andean región *Blanchardiella ecuadoriensis Dequal, Unoculubranchiobdella Solano Lobo (Ringuelet, 1976a, p. 101); San Geró- 1916 Peralta, Rodrigues Matos and Maues nimo stream (Ringuelet, 1981a, p. 193). Previous name. H. ecuadoriensis (De- Serra-Freire, 1998 qual, 1916). CYLICOBDELLIDAE Ringuelet, Distribution. ECU: León: Vallevicio- *U. expansa Solano Lobo Peralta, Ro- 1972 so (Ringuelet, 1981a, p. 193). drigues Matos and Maués Serra-Frei- re, 1998 Blanchardiella M. Weber, 1913 *Blanchardiella festai Dequal, 1916 Distribution. BRA: PA: Belém: Zoo- Type. B. fuhrmanni M. Weber, 1913) Previous name. Oxyptychus festai botanical Park, parasite of Podocnemis Synonym. Hypsobdella M. Weber, 1913 (Dequal, 1916). expansa (Chelonia) (Solano Lobo Pe- (Type: H. columbiensis M. Weber, 1913); Distribution. ECU (Dequal, 1916, p. 1). ralta et al., 1998, p. 161). Bibula R. Blanchard, 1917 (Type: Blan- chardiella fuhrmanni M. Weber, 1913) *Blanchardiella fuhrmanni M. Weber, Order ARRHYNCHOBDELLA R. (non Blanchardella Moniez, fish cesto- 1913 Blanchard, 1894 de); Clepsine (part) Savigný, 1822 Previous name. Bibula fuhrmanni (M. Weber, 1913). AMERICOBDELLIDAE Caballero, *Blanchardiella adaiophthalma Rin- Distribution. COL: Eastern cordille- 1956 guelet, 1980 ra; Boca del Monte (Weber, 1913, p.

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731); Páramo Cruz Verde; Del Ruiz ge: Piara (Siddall, 2001a, p. 11). PER: Erpobdella De Blainville, 1818 cordillera (Weber, 1915, p. 117). VEN: near Huaron (Ringuelet, 1960, p. 251). Synonym. Trochaeta Dutrochet, 1817 Merida (Ringuelet, 1981b, p. 226). ECU: Cajamarca: between Huarón and (Type: T. subviridis Dutrochet, 1817); Cahunayos. COL: Aquacatal; Suiza; Nephelis Savigný, 1822; Herpobdella *Blanchardiella octoculata M. Weber, Cafetal La Camelia (Weber, 1915, p. Agassiz, 1846; Nephelopsis Verrill, 1913 106); Cundinamarca (Ringuelet, 1872 (Type: N. obscura Verrill, 1872); Previous names. Helobdella octocu- 1981a, p. 193). VEN: Mérida (Ringue- Archaeobdella Grimm, 1876 (Type: A. lata (M. Weber, 1913); Bibula octocu- let, 1981b, p. 226); Mucuchíes desert: esmonti Grimm, 1876); Dina R. Blan- lata (M. Weber, 1913); Erpobdella oc- Lake Mucubaji (Ringuelet, 1981a, p. chard, 1892 (Type: D. lineata (O. F. toculata (M. Weber, 1913). 191). BRA (Ringuelet, 1971a, p. 99). Müller, 1774)); Mooreobdella Pawlo- Distribution. COL: Cundinamarca: Further distribution. Trinidad (We- wski, 1955 (Type: Hirudo octoculata near Santa Isabel (Ringuelet, 1980, p. ber, 1915, p. 107). Linnaeus, 1758); Fadejewobdella 6): Eastern cordillera: Bogotá (Weber, Lukin, 1962 (Type: F. quinqueannula- 1915, p. 125). *C. intermedium (Nonato, 1946) ta Lukin, 1929) Original name. Liostomum intermedia *Blanchardiella paramoensis M. We- Nonato, 1946. *E. dubia (Ringuelet, 1958) ber, 1913 Distribution. ARG. PAR (Ringuelet, Original name. Bathracobdella dubia Previous names. H. paramoensis (M. 1981a, p. 193). BRA: Central and sou- Ringuelet, 1958. Weber, 1913); Bibula paramoensis (M. thern (Nonato, 1946, p. 287). Distribution. ARG: Santa Cruz Prov: Weber, 1913). Argentine lake (Ringuelet, 1958, p. Distribution. COL: Eastern Cordille- *C. joseense (Grube and Örsted, 1859) 121). CHI: Antofagasta (Ringuelet, ra: Ruiz and Paramo Cruz Verde (We- Original and previous names. Cen- 1972b, p. 345). ber, 1915, p. 122). tropygus joseensis Grube and Örsted, 1859; Liostomum joseensis Grube and *E. wuttkei Kutschera, 2004 *Blanchardiella peruana Ringuelet, Örsted, 1859; Centropygus jocensis Distribution. South America (Kuts- 1960 (Grube and Örsted, 1959); Centropy- chera, 2004, p. 153). Previous names. H. peruana (Ringue- gus joscencis (Grube and Örsted, let, 1960a); Bibula peruana (Ringue- 1859); Nephelis tergestina R. Blan- Lumbricobdella Kennell, 1886 let, 1960). chard, 1892. Type. L. schaefferi Kennell, 1886 Distribution. PER (Ringuelet, 1960a, p. Distribution. ARG (Weber, 1915, p. 253): Huanuco: between Tingo Maria 105; Ringuelet, 1944, p. 55). PAR: San *L. chamensis Dequal, 1917 and Carpfish (Ringuelet, 1981a, p. 193). Bernardino; Asuncion; San José. VEN: Distribution. VEN: Chama: between Puerto Cabello; Caracas. COL: Bogo- Mérida and Zulia (Ringuelet, 1981a, p. *Blanchardiella tamboensis M. We- tá (Weber, 1915, p. 104). BRA (Ba- 193). ber, 1913 dham, 1923, p. 243): RS; SC: Blume- Previous names. Helobdella tambo- nau; Florianópolis (Desterro) (Weber, HIRUDINIDAE Whitman, 1886 ensis (M. Weber, 1913); Bibula tam- 1915, p. 104); SP: Barueri, in ant colo- boensis (M. Weber, 1913). nies (Lenko, 1972, p.10); RJ; ES; GO; Hirudo Linnaeus, 1758 Distribution. COL: Popayán: Eastern AM (Pinto, 1923, p. 1074). Type. H. medicinalis Linnaeus, 1758 Cordillera: Tambo: (Weber, 1915, p. 124). *C. lumbricoides Grube, 1871 H. medicinalis Linnaeus, 1758 Cylicobdella Grube, 1871 Previous name. Liostomum lumbricoi- Distribution. PER: San Martin Dep; Type. Centropygus joseensis Grube des (Grube, 1871). Ayacucho Dep (Beltran et al., 1998, p. and Örsted, 1859 Distribution. ECU: Sigsig. COL: Cun- 1123). BRA: RJ (Pinto, 1923, p. 1033). Synonym. Liostomum R. Blanchard, dinamarca. VEN. FRG. BRA: Northe- Further distribution. Throughout 1896 (non Wagler, 1931, nom. rej. Rin- ast (Ringuelet, 1981a, p. 193). Europe (Kutschera and Utevsky, 2006, guelet, 1948); Centropygus Grube and Remarks. Liostomum lumbricoides p. 1). Örsted, 1859; Cylicobdella Grube, 1871 was synonymyzed with L. joseensis by Remarks. The South American recor- Pinto (1923, p. 1072), but revalidated ds of this species are doubdtful, as there C. coccineum (Wagler, 1831) as Cylicobdella lumbricoides by Rin- is now uncertainty about the identity of Original and previous name. Liosto- guelet (1981a, p. 193). material referred to the two sister speci- mum coccineum Wagler, 1831; Centro- es in Europe, H. medicinalis and H. ver- pygus coccineus (Wagler, 1831). ERPOBDELLIDAE R. Blanchard, bana Carena, 1820 (Trontelj and Ute- Distribution. BOL: Apolobamba ran- 1894 vsky, 2005). There is now molecular

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evidence that commercially available ro, 1937b, p. 44). ARG. URU: Monte- Semiscolex Kinberg, 1866 European medical leeches are not Hi- video (Pinto, 1923, p. 1047). VEN: Type. S. juvenilis Kinberg, 1866 rudo medicinalis (Siddall et al., 2007). Caracas. BRA (Ringuelet, 1972a: 99); Synonym. Cyclobdella Weyenbergh, South (Ringuelet, 1981a, p. 192). 1866 MACROBDELLIDAE Richardson, 1969 SALIFIDAE Johansson, 1910 *S. coecus Ringuelet, 1936. Limnobdella R. Blanchard, 1893 Synonym. Trematobdellidae Johans- Distribution. ARG (Ringuelet, 1936, Type. L. mexicana R. Blanchard, 1893 son, 1913 p. 379). Synonym. Potamobdella Caballero, 1932 Barbronia Johansson, 1918 *S. glaber (Weyenbergh, 1877) L. mexicana (R. Blanchard, 1893) Type. B. rouxi Johansson, 1918 Original name. Cyclobdella glabra Distribution. VEN: Puerto Cabello. Synonym. Vivabdella Richardson, 1970 Weyenbergh, 1877. BRA: MT: Cuiabá (Weber, 1915, p. 82). (part) Distribution. CHI: Fritillar. ARG: Further distribution. Mexico (Blan- Córdoba. PAR: Asunción; San Bernar- chard, 1893, p. 1). B. weberi (R. Blanchard, 1897) dino (Weber, 1915, p. 97). BOL/PER: Previous name. Alboglossiphonia we- Formosa (Ringuelet, 1953a, p. 224). Oxyptychus Grube, 1851 beri (R. Blanchard, 1897). Type. O. striatus Grube, 1851 Distribution. BRA (Pamplin and Ro- *S. intermedius Ringuelet, 1942 Synonym. Argyrobdella Cordero, 1937 cha, 2000, p.723). Distribution. ARG (Ringuelet, 1945, (Type: Nephelis ornata Weyenbergh, Further distribution. United States, p. 122). 1883); Limnobdella R. Blanchard, Afghanistan, India, Southeast Asia, and 1893; Diplobdella Pinto, 1920 (Type: Australia (Govedich et al., 2002, p. *S. juvenilis Kinberg, 1866 Limnobdella brasiliensis Pinto, 1920). 225). Distribution. CHI. ARG: Córdoba. URU: Montevideo. PAR: Apa River; *O. brasiliensis (Pinto, 1920) SEMISCOLECIDAE Scriban and Au- Asunción; Alto Chaco; San Bernardi- Original and previous name. Limno- trum, 1934 no (Weber, 1915, p. 95). BRA: RS (De bdella brasiliensis Pinto, 1920; Diplo- Avila Goulart, 1963, p. 5); BA (Weber, bdella brasiliensis (Pinto, 1920). Patagoniobdella Ringuelet, 1972 1915, p. 95). Distribution. ARG. BRA (Cordero, Type. Semioscolex variabilis R. Blan- 1937b, p. 40): SP (Ringuelet, 1981a, chard, 1900 *S. notatus Cordero, 1937 p. 192); RJ: Campo Belo; MG: Lassan- Distribution. BRA: PB (Cordero, ce; GO: Araguary and Ipé Arcado (Pin- *P. ademonia Ringuelet, 1976 1937a, p. 24). to, 1920, p. 1); Pirenópolis (Brandão Distribution. ARG: Patagonia: Nahu- and Garda, 2000, p. 171). el Huapi andean lake (Ringuelet, *S. similis (Weyenbergh, 1877) 1976b, p. 61). Original name. Nephelis similis *O. inexpectatus Ringuelet, 1945 Weyenbergh, 1877. Distribution. URU. ARG (Ringuelet, *P. fraterna Ringuelet, 1976 Distribution. South America (Corde- 1945, p.116; 1949, p. 154; De Avila Distribution. CHI: (Ringuelet, 1981b, ro, 1937a, p. 60). ARG (Ringuelet, Goulart, 1963, p. 2). p. 226; 1985, p. 163); Neltume lake 1945, p. 120; 1949, p. 154). URU (Rin- (Siddall and Borda, 2004, p. 6). ARG: guelet, 1981a, p. 192). BOL: Cocha- *O. ornatus (Weyenbergh, 1883) Patagonia: Lacar lake (Ringuelet, bamba (Ringuelet, 1953a, p. 215); San- Original and previous name. Nephe- 1976b, p. 61). ta Cruz Dep: Volcán lagune (Siddall, lis ornata Weyenbergh, 1883; Argyob- 2001b, p. 2). PAR (Ringuelet, 1948, p. della ornata (Weyenbergh, 1883). *P. variabilis (R. Blanchard, 1900) 239). PER: Amazon (Shain et al., Distribution. ARG (Cordero, 1937b, Original name. Semiscolex variabilis 2007). BRA: RJ (Ringuelet, 1981a, p. p. 50). BRA: RS (De Avila Goulart, R. Blanchard, 1900 192). 1963, p. 3). Distribution. South America (Cordero, 1937b, p. 62). CHI: Fritillar; Concepci- *S. zonatus Oka, 1931 *O. strenuus Ringuelet, 1948 on (Weber, 1915, p. 98); Villarica and Distribution. South America (Oka, Distribution. PAR (Ringuelet, 1948, p. Huerquehue lakes (Siddall and Borda, 1931, p. 323). BRA (Oka, 1932, p. 236). 2004, p. 6). ARG (Weber, 1915, p. 100; 316). Ringuelet, 1945, p. 121). PAR: San Ber- *O. striatus Grube, 1851 nardino (Weber, 1915, 98). BRA: RS XEROBDELLIDAE Moore, 1946 Distribution. South America (Corde- (De Avila Goulart, 1963, p. 5). Synonym. Diestecotomatidae Ringue-

44 Volume 3 number 1 january - april 2008

039a048_Christoffersen.pmd 44 22/4/2008, 22:30 A Catalogue of the Piscicolidae, Ozobranchidae, and Arhynchobdellida (Annelida, Clitellata, Hirudinea) from South America

let, 1953; Mesobdellidae Ringuelet, Discussion References 1972; Nesophilaemonidae Ringuelet, 1982 No less than 66 nominal species of APAKUPAKUL, K.; SIDALL, M.E.; BURRE- Hirudinea belonging to the Piscicolidae, SON, E.M. 1999. Higher level relationships of Diestecostoma Vaillant, 1890 Ozobranchidae and Arhynchobdellida leeches (Annelida: Clitellata: Euhirudinea) ba- sed on morphology and gene sequences. Mole- Type. D. mexicana (Baird, 1869) have been refered in the literature for South cular Phylogenetics and Evolution, 12:350-359. Synonym. Heterobdella Baird, 1869 America, 56 of which are endemic to the BADHAM, C. 1923. On Centropygus joseensis, (pre-occupied); Hygrobdella Caballero, South American continent. Of the eight a leech from Brazil. Quarterly Journal of Mi- 1940 species that are not restricted to South croscopical Science London, 67:243-256. America, Cylicobdella coccineum extends BELTRAN, F. de E., MELGAR A., R.; TELLO C., R. 1998. Nasal infection by Hirudo medici- *D. trujillensis Ringuelet, 1976 its range into the Caribbean, and the other nalis leech. In: I. TADA; S. KOJIMA; M. TSU- Distribution. PER: Coastal zone (Rin- seven are more widely distributed in the JI (eds.), ICOPA IX: 9th International Congress guelet, 1976c, p. 67); La Libertad: Tru- world. More than 1/4 of the known species in Parasitology, Makuhari Messe, Chiba, Japan, jillo (Ringuelet, 1981a, p. 193). were described by the Argentinian Raul August 24-28, 1998. Bologna, Monduzzi Edito- Adolfo Ringuelet (†1914-1982), who has re, p. 1-1314. BLANCHARD, R. 1893. Révision des hirudi- Mesobdella R. Blanchard, 1893 thus contributed most to our knowledge nées du Musée de Turin. Bolletino dei Musei di Type. Hirudo gemmata C. E. Blan- of South American hirudineans belonging Zoologia ed Anatomia Comparata della Real chard, 1849 to these groups. Università di Torino, 8(145):1-32. Leeches represent very promising BRANDÃO, R.A.; GARDA, A.A. 2000. Scinax *M. gemmata (C. E. Blanchard, 1849) model organism for ecological studies, fuscomarginatus (NCN). Predation. Herpetolo- gical Review, 31:171. Original name, synonyms, and pre- especially in South America, where their BURRESON, E.M. 1981. A new deep-leech, Ba- vious names. Hirudo gemmata C. E. high degree of endemicity and their thybdella sawyeri, n.gen., n.sp., from thermal vent Blanchard, 1849; Hirudo cylindrica C. habitat diversification is very great. By areas on the Galapagos Rift. Proceedings of the E. Blanchard, 1849; Hirudo brevis Gru- inventariating the leech fauna of South Biological Society of Washington, 94:483-491. be, 1871; Mesobdella brevis (Grube, America, I hope to provide a first step BURRESON, E.M.; SEGONZAC, M. 2006. Morphological variability of Bathybdella sawyeri 1871). towards documenting the diversity of (Hirudinea: Piscicolidae) from hydrothermal Distribution. CHI (Ringuelet, 1942, these animals in our megadiverse and vents on the Galapagos Rift and the South East p. 364): Puerto Montt; Frutillar; El faunistically almost unknown continent, Pacific Rise. Zootaxa, 1286:15-21. Huito (Weber, 1915, p. 92); Valdivia particularly with regards to non- CORDERO, E.H. 1933. Notes sur les hirudinées. and Chiloé (Pinto, 1923, p. 1055); near vertebrate, non-arthropod groups. II. Piscicola platense n. sp., dun poisson sud- américain, Hoplias malabaricus (Bloch). Anna- Santiago: Chan-Chan Alto (Siddall Hopefully leeches will prove usefull as les de Parasitolologie Paris, 11:450-462. and Borda, 2004, p. 3); Central Chile, indicators of environmental conditions, CORDERO, E.H. 1937a. Hirudineos neotropi- between 36 oS and 38 oS: lakes Chica especially in freshwater habitats, where cales y subantárcticos. Nuevos, críticos o ya co- de San Pedro, Grande de San Pedro, their high rate of endemicity may be nocidos del Museo Argentino de Ciencias Natu- Quiñero, and Lanalhue (Munoz et al., shown to be almost unparalleled. rales. Anales del Museo Argentino de Ciências Naturales, 39:1-78. 2001, p. 177). CORDERO, E.H. 1937b. Los hirudineos del Nor- Acknowledgements deste del Brasil, I. Especies recogidas por Clemen- *M. notohilica Ringuelet, 1953 te Pereira en 1933 y por el autor en 1935. Anais Distribution. ARG (Ringuelet, 1953b, I am particularly indebted to Hong-Zhu da Academia Brasileira de Ciências, 9:13-26. p. 187). Wang, who invited me for a keynote DE AVILA GOULART, A.D. 1963. Hirudofau- na do município de Porto Alegre (Estado do Rio presentation on South American Grande do Sul, Brasil) (Annelida, Hirudinea). Nesophilaemon Nybelin, 1943 clitellates at the Tenth International Iheringia Série Zoologia, 29:1-8. Synonym. Philaemon Lambert, 1898 Symposium on Aquatic Oligochaeta in DEQUAL, L. 1916. Viaggio del dott. E. Festa (Type: P. pungens Lambert, 1898); Xenob- Wuhan, China. This paper is based in nel Darien, nell´ Ecuador e regioni vicine. XXV. della Richardson, 1975; Micobdella Richar- part on that presentation. I am very Irudinei. Bollettino del Museo di Zoologia et Ana- tomia Comparata di Torino, 31:1-20. dson, 1975; Castrabdella Richardson, 1975. grateful to Claudia Berrío and Imelda EPSHTEIN, V.M.; UTEVSKY, A.Y.; UTE- Vélez for sending me an important VSKY, S.Y. 1994. The system of fish leeches *N. skottsbergi (Johansson, 1924 ) paper by Ringuelet. Boris Sket read the (Hirudinea: Piscicolidae). Genus, 5:401-409. Original name. Philaemon skottsber- manuscript and offered several GOVEDICH, F.R.; BAIN, B.A.; DAVIES, R.W. gi Johansson, 1924. suggestions for improving this review, 2002. First record of the Asian freshwater leech Distribution. CHI: Juán Fernandez most of which have been incorporated Barbronia weberi (Blanchard, 1897) (Euhirudi- nea: Erpobdellidae) in Australia. The Victorian Archipelago (Johansson, 1924, p. 442; into this text. A productivity Naturalist, 119:225-226. Ringuelet, 1955, p. 137; Siddall and scholarship from CNPq provided JOHANSSON, L. 1924. Ein neuer Landblutegel Borda 2004, p. 16). financial support for this research. aus den Juan Fernandez Inseln. Natural History

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of Juan Fernandez Zoology, 3:439-460, 1 pl. PAGGI, A.C.; OCON, C.; TANGORRA, M.; entos sobre la hirudofauna neotrópica y transici- KUTSCHERA, U. 2004. The freshwater leech CAPITULO, A.R. 2006. Response of the zoo- onal mexicana. Physis, 27:367-380. Helobdella europaea (Hirudinea: Glossiphonii- benthos community along the dispersion plume RINGUELET, R.A. 1972a. Alguns hirudíneos dae): An invasive species from South America? of a highly polluted stream in the receiving wa- del Musem d´Histoire Naturelle de Paris. Phy- Lauterbornia, 52:153-162. ters of a large river (Rio de La Plata, Argentina). sis, 31:99-103. KUTSCHERA, U. 2006. The infamous blood Hydrobiologia, 568:1-14. RINGUELET, R. A. 1972b. Hirudíneos neotró- suckers from Lacus Verbanus. Lauterbornia, PAMPLIN, P.A.Z; ROCHA, O. 2000. First re- picos de Colombia, Cuba y Chile con la descrip- 56:1-4. port of Barbronia weberi (Hirudinea: Salifidae) ción de una nueva especie de Oligobdella (Glos- KUTSCHERA, U.; WIRTZ, P. 2001. The evolu- from South America. Revista de Biologia Tropi- siphoniidae). Physis Seccion B, 31(83):345-352. tion of parental care in freshwater leeches. The- cal, 48:723. RINGUELET, R.A. 1976a. Un curioso hirudi- ory in Biosciences, 120:115-137. PHILLIPS, A.J.; SIDDALL, M.E. 2005. Phylo- neo alto andino del Perú (Orchibdella peruvien- LENKO, K. 1972. Minhocas e sanguessugas geny of the New World medicinal leech family sis nov. sp., Hirudiniformes Cyclobdellidae). Li- (Annelida: Oligochaeta: Hirudinea) em ninhos Macrobdellidae (Oligochaeta: Hirudinida: Arhyn- mnobios, 1:101-104. de Camponotus rufipes (Insecta, Hymenoptera: chobdellida). Zoologica Scripta, 34:559-564. RINGUELET, R.A. 1976b. Dos hirudinoes nue- Formicidae). Revista Brasileira de Entomologia, PINTO, C. 1920. Contribuição ao estudo dos vos del género Patagoniobdella (Hirudinoidea 16:7-12. hirudíneos do Brasil (Limnobdella brasiliensis, Semiscolecidae) de los lagos andino patagóni- MACCALLUM, W.G.; MACCALLUM, G.A. nov. sp.). Brasil-Médico, 34:1-3. cos de la República Argentina. Limnobios, 1918. On the anatomy of Ozobranchus branchi- PINTO, C. 1923. Ensaio monographico dos hiru- 1:3:61-66. atus (Menzies). Bulletin of the American Museum díneos. Revista do Museu Paulista, 13:853-1118. RINGUELET, R.A. 1976c. Una nueva Dieste- of Natural History, 38:395-408. RINGUELET, R.A. 1936. Sobre una nueva es- costoma de la zona costera del Perú (D. rujillen- MANÉ-GARZÓN, F. 1973. Un nuevo tipo de pecie del género Semiscolex Kinberg (Hirudi- sis n. sp Hirudinea Diestecostomatidae) y diag- Hirudinea Colombobdella ringueleti n.gen. n.sp. nea). Notas del Museo de la Plata, Sección Zoo- nosis de esta familia. Neotropica, 22(68):67-76. parásito de una tortuga de Colombia. CON- logía, 1:379-388. RINGUELET, R.A. 1978. Hirudineos nuevos o GRESSO LATINOAMERICANO DE ZOOLO- RINGUELET, R.A. 1942. Sobre dos hirudineos ya descritos de la Argentina y del Uruguay. Lim- GIA, 5, Montevideo, Actas, 1:129-137. del sur de Chile: Mesobdella gemmata (E. Bl.) y nobios, 1:258-268. MANÉ-GARZÓN, F.; MONTEIRO, R. 1917. Helobdella similis Ring. Physis Buenos Aires, RINGUELET, R.A. 1980. Hirudineos terrestres Myzobdella uruguayensis n. sp. (Hirudinea Pis- 19:364-378. nor andinos y alto andinos de América del Sur. cicolidae) parásito de las branquias del bagre RINGUELET, R.A. 1944. Los hirudineos argen- Neotropica, 26:3-11. amarillo Rhamdia sapo (Val.). Revista de Biolo- tinos de los géneros Helobdella R. Bl., Batraco- RINGUELET, R.A. 1981a. Hirudinea. In: S. H. gia del Uruguay, 5:59-65. bdella Vig., Cylicobdella Gr. y Semioscolex HURLBERT; G. RODRIGUEZ, G.; N. D.DOS MOORE, J. P. 1910. Platybdella chilensis sp. nov. Kinb. Revista del Museo de la Plata, Sección SANTOS (eds.), Aquatic biota of tropical South Revista Chilena de Historia Natural, 14:28-30. Zoología, 4:5-94. America. Being a compilation of taxonomic bi- MOORE, J.P. 1911. Hirudinea of southern Pata- RINGUELET, R.A. 1945. Hirudineos del mu- bliographies for the fauna and flora of inland gonia. Reports of the Princeton University Ex- seo de La Plata. Revista del Museo de la Plata, waters of the tropical portion of South Ameri- pedition to Patagonia 1896-1899, 3:669-689. Sección Zoología, 4:95-137. ca). Part 2. Anarthropoda. San Diego, San Die- MOORE, J.P. 1931. A remarkable South Ameri- RINGUELET, R.A. 1948. Notas sobre hirudi- go University, p.191-196. can leech. Archivos de la Sociedad de Biologia neos neotropicales, V. Especies de la Republica RINGUELET, R.A. 1981b. Some advances in de Montevideo, 5(Suppl.):1220-1225. del Paraguay. Notas del Museo de la Plata, Sec- the knowledge of Neotropical leeches. Limnobi- MORILLAS, J.; GEORGE-NASCIMENTO, M.; ción Zoología, 13:213-244. os, 2:226. VALERIA, H.; KHAN, R.A. 1987. Trypanoso- RINGUELET, R.A. 1949. Notas sobre hirudi- RINGUELET, R.A. 1985. Sinopsis de los hiru- ma humboldti n.sp. from the Chilean catshark, neos neotropicales. VI. Presencia del genero dineos de Chile (Annelida). Boletín de la Socie- Schroedichthys chilensis (Guichenot, 1848). Glossiphonia en la Argentina y otras adiciones dad Biológica de Concepción, 56:163-179. Journal of Protozoology, 34:342–344. al conocimiento de la hirudofauna de los países SAWYER, R.T. 1971. Phylogenetic development MUNOZ, E.; MENDOZA, G.;VALDOVINOS, del Plata. Notas del Museo de la Plata, Sección of brooding behaviour in Hirudinea. Hydrobio- C. 2001. Evaluación de la biodiversidad en cin- Zoología, 14:141-159. logia, 37:197-204. co sistemas lénticos de Chile central: macroin- RINGUELET, R.A. 1953a. Notas sobre hirudí- SAWYER, R.T. 1986. Leech biology and beha- vertebrados bentónicos. Gayana, 65:173-180. neos neotropicalis. VIII. Algunas especies de vior. Oxford, Clarendon Press, 374 p. NONATO, E.F. 1946. Sobre sanguessugas do Bolivia y Peru. IX. Rehabilitación del género SAWYER, R.T.; DAMAS, D.; TOMIC, M.T. 1982. gênero Liostoma Wagler. Boletim da Faculdade ¨Cyclobdella¨ Weyenbergh. Notas del Museo Eva Anatomy and histochemistry of the salivary com- de Filosofia Ciências e Letras da Universidade Perón, 16:215-224. plex of the giant leech Haementeria ghilianii (Hi- de São Paulo, Zoologia, 11:287-331, pls 1–7. RINGUELET, R.A. 1953b. Notas sobre hirudi- rudinea, Rhynchobdellida). Archives de Zoologie OKA, A. 1931. Étude morphologique d´une nou- neos Neotropicales, VII. Un nuevo haemadipsi- Expérimentale et Genérale, 122:411-425. velle espèce de Semioscolex (Hirudinea) . Pro- do del género Mesobdella Blanch. Notas del Mu- SHAIN, D.H.; GUAVARA, M.I.A.; CORAL, ceedings of the Imperial Acadademy of Tokyo, seo de la Plata, Sección Zoología, 16:187-193. A.L.; CRAMPTON, G.R.; ALBERT, J.S. 2008. 7:323-326. RINGUELET, R.A. 1955. Sobre la sanguijuela A survey of freshwater annelids in the Peruvian OKA, A. 1932. Hirudinées extra-européenes du terrestre de Juan Fernandez (Philaemon skotts- Amazon. Acta Hydrobiologica Sinica Supple- Musée Zoologique Polonais. Annales del Mu- bergi Joh., Hirudinea). Investigación Zoológica ment (in press). seum de Zoologie Polonici, 9:313-328. en Chile, 2:137-142. SIDDALL, M.E. 2001a. Leeches of Laguna Vol- OMODEO, P. 1998. History of Clitellata. Itali- RINGUELET, R.A. 1960. Hirudineos terrestres can, Bolivia, including a new species of Helob- an Journal of Zoology, 65:51-73. del Perú. CONGRESSO SUDAMERICANO della (Clitellata: Hirudinea). American Museum OREVI, M.; ELDOR, A.; GIGUZIN, I.; RIGBI, DE ZOOLOGÍA, 1, La Plata, Actas y Trabajos, Novitates, 3313:1-11. M. 2000. Jaw anatomy of the blood-sucking lee- p. 251-256. SIDDALL, M.E. 2001b. Hirudinea from the Apo- ches, Hirudinea Limnatica nilotica and Hirudo RINGUELET, R.A. 1968. Llave o clave para el lobamba in the Bolivian Andes, including three medicinalis, and its relationship to their feeding reconocimiento de las sanguijuelas conocidas de new species of Helobdella (Clitellara: Hirudinea). habits. Journal of Zoology, 250:121-127. la República Argentina (Hirudinea) y apuntami- American Museum Novitates, 3341:1-14.

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SIDDALL, M.E. 2002. Phylogeny of the leech matics and Evolutionary Research, 37:141-147. family Erpobdellidae (Hirudinea: Oligochaeta). TRONTELJ,P.; UTEVSKY, S.Y. 2005. Celebrity Invertebrate Systematics, 16:1-6. with a neglected taxonomy: Molecular systema- SIDDALL, M.E.; APAKUPAKUL, K.; BURRE- tics of the medicinal leech (genus Hirudo). Mole- SON, E.M.; COATES, K.A.; ERSÉUS, C.; GEL- cular Phylogenetics and Evolution, 34:616-624. DER, S.R.; KALLERJO, M.; TRAPIDO-RO- VOLONTERIO, O.; LEON, E.L.; DE LEON, SENTHAL, H. 2001. Validating Livanow: Mo- R.P. 2004. Infestation dynamics and histopatho- lecular data agree that leeches, branchiobdelli- logy of two species of freshwater leeches (Hiru- dans and Acanthobdella peledina form a mono- dinea : Piscicolidae) on teleost fish from Uru- phyletic group of oligochaetes. Molecular Phylo- guay. Comparative Parasitology, 71:21-28. genetics and Evolution, 21:346-351. WEBER, M. 1913. Hirudinées columbiennes. SIDDALL, M.E.; BORDA, E. 2004. Leech col- Voyage d´Exploration scientifique en Columbie. lections from Chile including two new species Memoires de la Societé Neuchateloise des Sci- of Helobdella (Annelida : Hirudinida). Ameri- ences Naturelles, 5:731-747. can Museum Novitates, 3457:1-18. WEBER, M. 1915. Monographie des hirudinées SIDDALL, M.E.; BUDINOFF, R.B. 2005. Dna- sudamericaines. Imprimaterie Attinger Frères, barcoding evidence for widespread introductions Neuchatel, 134 p. of a leech from the South American Helobdella triserialis complex. Conservation Genetics, Submitted on August 17, 2007 6:467-472. Accepted on March 17, 2008 SIDDALL, M.E.; BURRESON, E.M. 1996.Le- eches (Oligochaeta?: Euhirudinea), their phylo- geny and the evolution of life-history strategies. Hydrobiologia, 334:277-285. SIDDALL, M.E.; TRONTELJ, P.; NKAMANY, M.; MACDONALD, K.S. 2007. Diverse mole- cular data demonstrate that commercially avai- lable medicinal leeches are not Hirudo medici- nalis. Proceedings of the Royal Society B Biolo- gical Sciences 274(1617):1481-1487. SKET, B. 1968. Zur Kenntnis der Egel-Fauna Ju- goslawiens. Rasprave SAZU Class IV 11:127-197. SOLANO LOBO PERALTA, A.; RODRIGUES MATOS, E.; MAUES SERRA-FREIRE, N. 1998. Unoculubranchiobdella expansa gen.nov., sp.n. (Hirudinea: Ozobranchidae) parasito de Podocnemis expansa (Chelonia: Pelomedusidae). Entomologia y Vectores, 5:161-177. SOTO, J.M.R. 2000. Marine leech, Stibarobdella macrothela (Schmarda, 1861) (Hirudinea, Piscico- lidae), parasitic on the whaler shark, Carcharhinus brachyurus (Gunther, 1870) (Chondrichthyes, Car- charhinidae), in southern Brazilian waters. Revista Brasileira de Biologia, 60:713-714. SOTO, J.M.R. 2001. First record of a turtle lee- ch, Ozobranchus margoi (Apáthy, 1890) (Anne- lida, Clitellata), on La Plata dolphin, Pontoporia blainvillei (Gervais & D´Orbigny, 1844) (Ceta- cea, Platanistoidea). Mare-Magnum, 1:173. SOTO, J.M.R. 2003. The marine leech Stibaro- bdella loricata (Harding, 1924) (Hirudinea, Pis- cicolidae), parasitic on the angel shark Squatina spp. and sandtiger shark Carcharias taurus Ra- finesque, 1810 (Chondrichthyes: Squatinidae, Carchariidae) in southern Brazilian waters. Bra- zilian Journal of Biology, 63:691-694. TANZOLA, R.D.; BOTTÉ, S.E. 1998. Presencia de Branchellion parkeri Richardson, 1949 (Hiru- dinea: Piscicolidae) sobre Sympterygia bonapar- tei Muller & Henle, 1841 (Pisces: Rajidae) en el Atlantico Sur. Natura Neotropicalis, 29:65-68. TRONTELJ, P.; SKET, B.; STEINBRÜCK, G. 1999. Molecular phylogeny of leeches: Congruence of nu- clear and mitochondrial rDNA data sets and the ori- gin of bloodsucking. Journal of Zoological Syste-

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039a048_Christoffersen.pmd 47 22/4/2008, 22:30 Martin Lindsey Christoffersen

Appendix 1. Systematic Index

Order Rhynchobdellida Piscicolidae Bathybdella Branchellion Johanssonia Myzobdella Platybdella Pontobdella Stibarobdella Trachelobdella Ozobranchidae (turtle leeches) Bogabdella Ozobranchus Unoculubranchiobdella Order Arhynchobdellida Americobdellidae Americobdella Cyclobdellidae Orchibdella Cylicobdellidae Blanchardiella Cylicobdella Lumbricobdella Erpobdellidae Erpobdella Hirudinidae Hirudo Macrobdellidae Limnobdella Oxyptychus Salifidae Barbronia Semiscolescidae Patagoniobdella Semiscolex Xerobdellidae Diestecostoma Mesobdella Nesophilaemon

48 Volume 3 number 1 january - april 2008

039a048_Christoffersen.pmd 48 22/4/2008, 22:30