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A catalogue of aquatic microdrile oligochaetes (Annelida: Clitellata) from


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第 31 卷 增刊 水 生 生 物 学 报 Vol.31, Suppl.
2007 年 12 月 ACTA HYDROBIOLOGICA SINICA Dec., 2007

A catalogue of aquatic microdrile oligochaetes (Annelida: Clitellata)


from South America

Martin Lindsey Christoffersen

Departamento de Sistemática e Ecologia, Universidade Federal da Paraíba, 58059-900, João Pessoa, Paraíba, Brazil
(*Author for correspondence: Tel: +55-083-3216-7773; E-mail: mlchrist@dse.ufpb.br or mlchrist@pq.cnpq.br)

Key words: Alluroididae, Capilloventridae, Haplotaxidae, Naididae (including Tubificidae s.str.), Narapidae, Oligochaeta,
Opistocystidae, Phreodrilidae, and Tiguassuidae.

Abstract
A catalogue of aquatic microdrile oligochaetes exclusive of Enchytraeidae produced 171 nominal taxa (species and
subspecies) reported to date from South America. This number may be compared to the 213 freshwater oligochaetes
known from continental North America north of Mexico. The South American species belong to 55 genera, 41 of which
are Naididae (this family now includes the paraphyletic Tubificidae s.str.). Circa 38.5% of this fauna is endemic to South
America. Full synonyms and detailed South American occurrences are provided. This is the first reassessment of South
American aquatic microdriles in 18 years.

Introduction Reynolds & Wetzel (2008, in prep.). The aquatic


oligochaetes of the world are reviewed by Brinkhurst
There are almost 5000 valid species of Oligochaeta, & Jamieson (1971) and Brinkhurst & Wetzel (1984).
about one-third of which are aquatic (Martin et al., Those interested in the older taxonomic history of
2008). Circa 110 freshwater oligochaetes have been genera and species should refer to Michaelsen (1900)
previously estimated for Central and South America and Sperber (1948). Recently, Pinder (2007) reported
(Rocha, 2006: 23), while 14 marine oligochaete 221 aquatic species for Australia, and Wetzel et al.
species are referred from Brazil (Migotto & Marques, (2008) reported the occurrence of 213 freshwater
2006: 166). However, these estimates were based oligochaetes in continental North America north of
largely on questionnaires sent to South American Mexico. Gavrilov (1979, 1981) and Brinkhurst &
specialists, including the late Dr. Gilberto Righi Marchese (1989) provided an inventory and a guide
(*1937-1999). for the aquatic oligochaetes from South America, the
Most families of microdrile oligochaetes are latter with identification keys. Righi (1984) compiled
relatively small in size, and predominantly aquatic in a guide with identification keys for the aquatic oligo-
nature (Reynolds & Cook, 1976: 2). The microdrile chaetes from Brazil. Berrío-Cárdenas & Velez (2008,
family Enchytraeidae, comprising taxa that are also paper included in this volume) provide a review of the
relatively small in size, is predominantly terrestrial but aquatic Oligochaeta occurring in Colombia. In this
with many species occurring in a variety of freshwater, paper I provide a catalogue of the aquatic microdrile
estuarine, and marine environments. The Enchy- oligochaetes cited to date from the South American
traeidae of South America will be addressed in a continent. Due to space limitations, I have excluded
subsequent publication. the Enchytraeidae as a whole from this present
Major world-catalogues of oligochaetes include catalogue, although many species in this predo-
Reynolds & Cook (1976, 1981, 1989, 1993) and minantly terrestrial family occur in aquatic and
60 ACTA HYDROBIOLOGICA SINICA Vol. 31

semi-aquatic habitats in South America and elsewhere *Brinkhurstia donaldi Omodeo & Coates, 2001.
in the world. Distr: GUY: 3, 4.
This paper has two objectives: to provide a
current starting point for taxonomists, systematists, Kathrynella Omodeo, 1996 (Type: Kathrynella
and ecologists interested in South American aquatic guyanae).
microdrile oligochaetes, and to compile the available *Kathrynella guyanae Omodeo, 1996. Distr:
published information on the biodiversity and dis- GUY: 3.
tribution of this neglected zoological group.
Throughout this catalogue, I use 3-letter abbre- Righiella Omodeo & Coates, 2001 (Type: Righiella
viations for South American countries (ARG, Argentina; jamiesoni).
BOL, Bolivia; BRA, Brazil; CHI, Chile; COL, *Righiella jamiesoni Omodeo & Coates, 2001.
Colombia; ECU, Ecuador; GUY, Guyana; PAR, Distr: GUY: 3.
Paraguay; PER, Peru; SUR, Surinam; URU, Uruguay;
VEN, Venezuela), and 2-letter abbreviations for States CAPILLOVENTRIDAE Harman & Loden, 1984
in Brazil (AL, Alagoas; AM, Amazonas; BA, Bahia; (Type: Capilloventer).
CE, Ceará; DF, Distrito Federal; MG, Minas Gerais; Capilloventer Harman & Loden, 1984 (Type:
MS, Mato Grosso do Sul; MT, Mato Grosso; PA, Pará; Capilloventer atlanticus).
PE, Pernambuco; PI, Piauí; PR, Paraná; RJ, Rio de Capilloventer atlanticus Harman & Loden, 1984.
Janeiro; RN, Rio Grande do Norte; RR, Roraima; RS, Distr: BRA: RJ: 1.
Rio Grande do Sul; SC, Santa Catarina; SP, São
Paulo). Detailed geographical distributions within HAPLOTAXIDAE Michaelsen, 1900 (Type: Ha-
these political units are provided in the appendix. I plotaxis). Syn: Phreoryctidae Claus, 1880.
have only indicated noteworthy ecological associations Haplotaxis Hoffmeister, 1843 (Type: Lumbricus
(e.g., ´in floating Eichhornia´, ´in the mantle cavity of gordioides G. L. Hartmann, 1821). Syn: Phreoryctes
mollusks´, ´50m deep´); more general ecological Hoffmeister, 1845 (Type: Phreoryctes menkeana Hoff-
associations (e.g., ´in fine sediment´, ´850 m high´) have meister, 1843); Georyctes Schlotthauber, 1860 (Type:
not been included in this catalogue. Abbreviations used Georyctes darwinii Keller, 1885); Nemodrilus Cla-
in the accounts below, are as follows: Furth. Distr. = parède, 1862 (Type: Nemodrilus filiformis Claparède,
further distributions (published); Rem. = additional 1862); Lumbricogordius (C. V. Heyden, 1874, in MS)
remarks for the taxon. Noll, 1874 (Type: Lumbricogordius hartmanni Noll,
Species restricted in distribution to South America 1874); Pelodrilus Beddard, 1891 (Type: Pelodrilus
(based upon published records) are preceded by an violaceus Beddard, 1891); Dichaeta Friend, 1896
asterisk (*). More detailed geographical occurrences (Type: Dichaeta curvisetosa Friend, 1896); Hetero-
are listed in the Appendix. chaetella Yamaguchi, 1953 (Type: Heterochaetella
glandularis Yamaguchi, 1953); Adenodrilus Cekano-
Results
skaya, 1959 (Type: Adenodrilus denticulatus Cekanovskaya,
ALLUROIDIDAE Michaelsen, 1900 (Type: Alluroides 1959); Haplolumbriculus (nom. nud.) Omodeo, 1958.
Beddard, 1894). *Haplotaxis aedeochaeta Brinkhurst & Marchese,
Brinkhurstia Jamieson, 1968 (Type: Alluroides 1987. Distr: ARG: 30, 85, 88. BRA: PR: 1-3.
americanus). Haplotaxis gordioides gordioides (G. L.
*Brinkhurstia americana (Brinkhurst, 1964). Syn: Hartmann, 1821). Syn: Lumbricus gordioides G. L.
Alluroides americanus Brinkhurst, 1964. Distr: ARG: Hartmann, 1821; Phreoryctes gordioides (G. L.
25, 28, 83, 85. GUY: 2. BRA: PR: 2, 3; SP: 39; 46, 47. Hartmann, 1821); Phreoryctes (Haplotaxis) gordioides
MS: 1; AM: 2. Rem: This species was named (G. L. Hartmann, 1821); Haplotaxis gordioides (G. L.
Alluroides americanus as a nom. nud. by Cernosvitov Hartmann, 1821); Tubifex uncinarius Ant. Dugès,
(1936), was described by Brinkhurst (1964), and 1837; Clitellio ? (Limnodrilus) uncinarius (Ant. Dugès,
redescribed by Jamieson (1968). 1837); Haplotaxis menkeana Hoffmeister, 1843;
Suppl. Christoffersen: A catalogue of aquatic microdrile oligochaetes (Annelida: Clitellata) from South America 61

Phreoryctes menkeanus (Hoffmeister, 1843); Georyctes tubificids (Tubificidae s.str.) should be regarded as
menkei Schlotthauber, 1854; Georyctes lichtensteinii members of the Naididae whenever the two names are
Schlotthauber, 1860; Nematodrilus filiformis Claparède, regarded as synonyms. The subfamilies of Tubificidae
1862; Phreoryctes filiformis (Claparède, 1862); s.str. (Tubificinae, Rhyacodrilinae, Phallodrilinae,
Phreoryctes heydeni Noll, 1873; Lumbricogordius Limnodrilinae, and Telmatodrilinae) are now included
hartmanni (C. V. Heyden, not publ.) Noll, 1874; within Naididae (Erséus et al., 2008).
Phreoryctes emissarius S. A. Forbes, 1890; Phreoryctes
endeca Giard, 1894; Diachaeta curvisetosa Friend, 1896; Aktedrilus Knöllner, 1935 (Type: Akedrilus mo-
Phreoryctes dichaetus Friend, 1896; Phreoryctes nospermathecus Knöllner, 1935). Syn: Basescuella
endeka var. pachyderma Ferronière, 1899; Haplotaxis Hrabe, 1973 (Type: Basescuella pontica Hrabe, 1973);
endeka var. pachyderma (Ferronière, 1899); ? Haplotaxis Phallodrilus (part), Brinkhurst, 1971.
intermedia Pierantoni, 1904; Haplotaxis forbesi Smith, *Aktedrilus brevis Erséus, 1980. Distr: BRA: BA:
1918. Distr: ARG: 60, 72, 84, 85, 86, 98. Furth. Distr: 1.
Holarctic (Brinkhurst, 1971a: 291). Aktedrilus cuneus Erséus, 1984: 193. Distr: Ecu:
3. Furth. Distr: Europe and Middle East (Erséus, 1989:
Metataxis Righi, 1985 (Type: Metataxis eliae). 16).
Syn: Pelodrilus Beddard, 1891 (Type: Pelodrilus *Aktedrilus dentatus Erséus, 1983. Distr: BRA:
violaceus Beddard, 1891). Rem: Although Brinkhurst RJ: 4.
(1988) placed Metataxis in the order Lumbricina, I Aktedrilus locyi Erséus, 1980. Distr: ECU: 3.
have herein followed Righi (1988: 316) in maintaining Furth. Distr: Bermudas, West coast of United States
this genus in the Haplotaxidae. (Erséus, 1984: 193), and Japan (Takashima & Mawatari,
*Metataxis americanus (Černosvitov, 1939). Syn: 1997: 33).
Phreoryctes americanus Černosvitov, 1939; Pelodrilus Aktedrilus parvithecatus (Erséus, 1978). Syn:
americanus (Černosvitov, 1939); Haplotaxis ame- Bacescuella parvithecata Erséus, 1978; Bacescuella
ricanus (Černosvitov, 1939). Distr: PER: 25. pilicrepus Erséus, 1984. Distr: ECU: 3. Furth. Distr:
*Metataxis bare Righi, 1988. Distr: BRA: RR: 1. Bermuda and France (Erséus, 1984: 196).
*Metataxis eliae Righi, 1985. Distr: VEN: 2.
Allodero Sperber, 1948 (Type: Nais elongata
NAIDIDAE Ehrenberg, 1828 (as Naidina, spe- Stephenson, 1911). Rem: Brinkhurst (1971d: 361) and
lling corrected by Benham, 1890) (Type: Nais). Syn: more recently Milligan (1997), Kathman & Brinkhurst
Tubificidae Vejdovský, 1876 (Type: Tubifex). Rem: (1998), and Wetzel et al. (2008) continue to treat
Based on sequences of 18S rDNA and other molecular Allodero, Aulophorus and Dero as subgenera of Dero.
and morphological data, Erséus & Gustavsson (2002) However, there is also some precedence for conside-
and Erséus et al. (2002) concluded that the family ring these subgenera as full genera. For example,
Naididae is more correctly placed within the subfamily Cekanovskaya (1962) and Liang (1964) regarded
Naidinae of the Tubificidae, with ´Naididae´ thus Dero and Aulophorus as separate genera, while the
becoming a junior synonym of the family Tubificidae. latter author also separated Allodero at the generic
A petition was then submitted by Erséus et al. (2005) level. More recently, Timm (1999, in table) listed
to the International Commission of Zoological Aulophorus as a full genus. The definitions of these
Nomenclature (ICZN, the Commission), requesting three subdivisions overlap considerably, partly owing
the family-group name Tubificidae Vejdovský, 1876, to the inadequate state of knowledge of Allodero
be given precedence over Naididae Ehrenberg, 1828. (Brinkhurst, 1971d: 361). A satisfactory taxonomic
In 2007, however, the Commission voted against this solution is not presently in site. For the purpose of this
proposal, maintaining precedence for Naididae over catalogue, I have decided to list the former three
Tubificidae (ICZN 2007). Erséus et al., 2008) recently subdivisions of Dero as separate genera, hoping that
published a paper summarizing the short history of better taxonomic criteria will be established for their
this issue, with the consequence being that the former separate recognition in the future or, alternatively, that
62
60 ACTA HYDROBIOLOGICA SINICA Vol. 31

if some of these names be demonstrated to be 2, 9, 10, 13, 14, 17; AM: 3, 5. 7. Furth. Distr: Central
untenable, they be synonymized with their closest taxa America (Righi & Hamoui, 2002: 142), North
and their use also be discontinued at the subgeneric America ? (see Kathman & Wetzel, 2003: 548), Africa,
level. and Australasia (Gavrilov, 1979: 104).
Allodero lutzi (Michaelsen, 1926). Syn: Schmardaella Allonais pectinata (Stephenson, 1910). Syn: Nais
lutzi Michaelsen, 1926; Dero lutzi (Michaelsen, 1926); pectinata Stephenson, 1910; Nais denticulata Chen,
Dero (Allodero) lutzi Michaelsen, 1926); Schmar- 1940. Distr: VEN: 22. Furth. Distr: Central America,
daella hylae Goodchild, 1951. Distr: VEN: 6. BRA: North America, Africa, Asia, and Australia (Righi &
SC: 1; SP: 11; RJ: 2, 3; MG: 2. Furth. Distr: Cuba and Hamoui, 2002: 143).
North America (Righi, 1984: 27).
Amerigodrilus Holmquist, 1985 (Type: Limnodrilus
Allonais Sperber, 1948 (Type: Nais pectinata var. chacoensis).
inaequalis Stephenson, 1911). *Amerigodrilus chacoensis (Stephenson, 1931).
Allonais chelata (Marcus, 1944). Syn: ? Schmar- Syn: Limnodrilus chacoensis Stephenson, 1931: 309.
daella filiformis, Moore, 1906; Nais paraguayensis f. Distr: PAR: 3.
chelata Marcus, 1944. Distr: ARG: 1. BRA: SP: 46;
PE: 6; PA: 2, 7, 9, 12, 13, 14, 15; AM: 7, 10. Furth. Amphichaeta Tauber, 1879 (Type: Amphichaeta
Distr: North America (Righi, 1984: 30). leydigi).
Allonais inaequalis (Stephenson, 1911). Syn: Amphichaeta leydigi Tauber, 1879. Distr: ARG: 8,
Nais pectinata var. inaequalis Stephenson, 1911. Dero 58. Furth. Distr: North America (Kathman &
(Allonais) inaequalis (Stephenson, 1911);Nais pec- Brinkhurst, 1998: 46) and Europe (Di Persia, 1977a:
tinata, Stephenson, 1931. Distr: ARG: 33, 35. PAR: 2. 121). Rem: Pavé & Marchese (2005: 188) listed
PER: 44; ECU: 1. SUR: 16. BRA: SP: 38. Furth. Distr: Amphichaeta americana Tauber, 1879 for Entre Ríos
Central America (Righi & Hamoui, 2002: 141), Africa, province, Argentina. This must be a lapsus for
Asia, Australia (Gavrilov, 1979: 104), and USA Amphichaeta leydigi Tauber, 1879. Amphichaeta
(Kathman & Wetzel, 2003: 548). americana Chen, 1944 occurs in the USA (Brinkhurst,
Allonais paraguayensis aequalis (Stephenson, 1971d: 316).
1920). Syn: Nais paraguayensis var. aequalis Ste-
phenson, 1920. Distr: BRA: PE: 3. Furth. Distr: Africa Aulodrilus Bretscher, 1899 (Type: Aulodrilus
and Asia (Righi, 1984: 30). limnobius). Syn: Siolidrilus Du Bois-Reymond Marcus,
Allonais paraguayensis paraguayensis (Michaelsen, 1949 (Type: Siolidrilus adetus).
1905). Syn: Nais paraguayensis Michaelsen, 1905; Nais *Aulodrilus adetus Du Bois-Reymond Marcus,
paraguayensis f. typica (Michaelsen, 1905); Allonais 1949. Syn: Siolidriulus adetus Du Bois-Reymond
paraguayensis (Michaelsen, 1905); Allonais paraguayen- Marcus, 1949. Distr: BRA: MS: 1; PA: 5.
sis paraguayensis (Michaelsen, 1905); Dero (Allonais) Aulodrilus cernosvitovi Du Bois-Reymond Marcus,
paraguayensis paraguayensis (Michaelsen, 1905); ? Nais 1947. Distr: BRA: PA: 4, 7, 9, 12, 13, 14. Furth. Distr:
paraguayensis var. barkudensis Stephenson, 1921; Nicaragua (Righi, 1984: 38).
Allonais paraguayensis ssp aequatorialis (Černosvitov, Aulodrilus limnobius Bretscher, 1899. Syn: Paranais
1938); Dero (Aulophorus) paraguayensis var. aequa- tenuis Černosvitov, 1937; Aulodrilus tenuis (Černosvitov,
torialis Černosvitov, 1938; Dero (Alllonais) para- 1937). Distr: ARG: 28, 38. BRA: PR: 2, 3, 6; SP: 18.
guayensis ssp aequatorialis Černosvitov, 1938; Allonais Furth. Distr: North America (Kathman & Brinkhurst,
lairdi Naidu, 1966; Allonais paraguayensis ghanensis 1998: 186), Europe, and Asia (Righi, 1984: 38).
Hrabe, 1966; Allonais arieli Gluzman de Pascar, 1990. Aulodrilus pigueti Kowalewski, 1914. Distr:
Distr: ARG: 33, 36, 58, 85, 86. URU: 3. PAR: 2. PER: ARG: 8, 32, 42, 58, 63, 83, 85, 86, 106. PER: 3, 45.
40, 43. VEN: 13. SUR: 2, 3, 9. BRA: SP: 12, 14, 34, COL: 1; GUY: 1. BRA: PR: 2, 3; SP: 35; MS: 1. Furth.
39, 40, 46, 47; AL: 1-3; PE: 1, 3, 9, 10, 14; CE: 1; PA: Distr: Cosmopolitan (Brinkhurst, 1971b: 527).
Suppl. Christoffersen: A catalogue of aquatic microdrile oligochaetes (Annelida: Clitellata) from South America 63
61

Aulophorus Schmarda, 1861 (Type: Nais fur- Aulophorus schardei var. huaronensis Piguet, 1928;
cata). Dero (Aulophorus) huaronensis (Piguet, 1928); Dero
*Aulophorus barbatus Righi & Hamoui, 2002. (Aulophorus) shmardei f. huaronensis (Piguet, 1928).
Distr: SUR: 11. Distr: PER: 1. Furth. Distr: Central America (Righi &
*Aulophorus beadlei Stephenson, 1931. Syn: Hamoui, 2002: 140).
Dero (Aulophorus) beadlei Stephenson, 1931. Distr: Aulophorus hymanae (Naidu, 1962). Syn: Dero
PAR: 2. (Aulophorus) hymanae Naidu, 1962. Distr: ARG: 58,
*Aulophorus bimagnasetus Harman, 1974. Syn: 84. Furth. Distr: Asia (Naidu & Srivastava, 1980:
Dero (Aulophorus) bimagnasetus Harman, 1974. Distr: 267).
SUR: 2, 9. Aulophorus indicus (Naidu, 1962). Syn: Dero
Aulophorus borellii (Michaelsen, 1900). Syn: (Aulophorus) indicus Naidu, 1962. Distr: ARG: 84.
Dero sp. Cognetti, 1900; Dero borelli Michaelsen, GUY: 1. Furth. Distr: Bonaire (Righi & Hamoui, 2002:
1900; Aulophorus borellii (Michaelsen, 1900); Dero 136) and India (Stacey & Coates, 1996: 23).
(Aulophorus) borellii (Michaelsen, 1900). Distr: ARG: Aulophorus lodeni (Brinkhurst, 1986). Syn:
1, 85. PAR: 2. GUY: 1. BRA: SP: 47; MS: 1; MT: 1; Aulophorus pectinatus Stephenson, 1931 (non Dero
PA: 1, 2, 6, 7, 9, 10, 13, 14, 16; AM: 6. Furth. Distr: pectinata Aiyer, 1929) (non Nais pectinata Step-
Possibly North America (Gavrilov, 1979: 105; henson, 1910); Nais denticulata Chen, 1940; Dero
Kathman & Brinkhurst, 1998: 74). intermedia Loden & Harman, 1982; Dero (Aulophorus)
*Aulophorus carteri Stephenson, 1931. Syn: lodeni Brinkhurst, 1986. Distr: ARG: 9, 38, 67, 85, 86,
Dero (Aulophorus) carteri Stephenson, 1931. Distr: 107. PAR: 2, 4. COL: 1, 2, 4, 11. GUY: 1. SUR: 9.
ARG: 96. URU: 1. PAR: 2. SUR: 9. BRA: SP: 34; PE: BRA: SP: 39, 40; PA: 3, 15. Furth. Distr: Costa Rica
9, 10, 11, 15; PI: 1; PA: 2, 6, 12-14, 16; MA: 4, 9, 11. (Stacey & Coates, 1996: 23), North America
*Aulophorus costatus Du Bois-Reymond Marcus, (Kathman & Brinkhurst, 1998: 74), Africa (Harman,
1944. Syn: Aulophorus schmardai f. costata Du 1974: 1), and India (Stacey & Coates, 1996: 23).
Bois-Reymond Marcus, 1944; Dero (Aulophorus) Aulophorus schmardai (Michaelsen, 1905). Syn:
costatus (Du Bois-Reymond Marcus, 1944); Dero ? Dero vaga, Stieren, 1892; Dero schmardai Michaelsen,
(Aulophorus) schmardai, Harman, 1974. Distr: ARG: 1905; Aulophorus schmardai var. huaronensis (Piguet,
36, 58, 85, 86. SUR: 9. BRA: SP: 13. 1928). Distr: ARG: 64, 85. PAR: 1. PER: 33. Furth.
Aulophorus furcatus (O. F. Müller, 1774). Syn: Distr: Trinidad (Stieren, 1872: 118).
Nais furcata O. F. Müller, 1774. Dero (Aulophorus) Aulophorus superterrenus Michaelsen, 1912. Syn:
furcatus (O. F. Muller, 1774); ? Aulophorus palustris Dero (Aulophorus) superterrenus (Michaelsen, 1912).
Michaelsen, 1905; Dero sp. Stephenson, 1910; ? Aulophorus Distr: ARG: 1. BRA: PR: 5; SP: 9, 24, 31; RJ: 1; PE:
furcatus var. brevipalpus Golanski, 1911; Aulophorus 18. Furth. Distr: Costa Rica (Righi, 1984: 27), North
stephensoni Michaelsen, 1912; ? Aulophorus afri- America (Frank & Lounibos, 1987: 18), and Asia
canus Michaelsen, 1914; Dero roseola Nicholls, 1921. (Righi, 1984: 27).
Distr: ARG: 8, 14, 26, 35, 37, 38, 42, 43, 50, 54, 58, Aulophorus tonkinensis (Vejdovský, 1894). Syn:
62, 73-75, 82, 85, 86, 88-90, 94, 106. URU: 2; BOL: 4. Aulophorus oxycephalus Schmarda, 1861; Dero
VEN: 1. GUY: 1. SUR: 2-6, 8-10. BRA: PR: 2, 3; SP: (Aulophorus) tonkinensis Vejdovský, 1874; ? Dero
12, 14, 38-40; MS: 1; PE: 7, 8, 16, 17; PA: 2, 6, 10, 11, stuhlmanni Stieren, 1892; ? Aulophorus stuhlmanni
18; AM: 4. Furth. Distr: Central America (Stacey & (Stieren, 1892). Distr: ARG: 35, 39, 85. SUR: 9. Furth.
Coates, 1996: 23), North America, Europe, Africa, and Distr: Africa and Asia (Harman, 1974: 27).
Asia (Cordero, 1931: 350). Aulophorus tridentatus Hrabe, 1966. Distr: VEN:
Aulophorus gravelyi (Stephenson, 1925). Syn: 13. Furth. Distr: Africa (Hrabe, 1966: 380).
Dero (Aulophorus) gravelyi Stephenson, 1925. Distr: Aulophorus vagus Leidy, 1880. Syn: Dero vaga
ARG: 85, 86. Furth. Distr: Asia (Marchese, 1986: (Leidy, 1880); Dero (Aulophorus) vagus (Leidy, 1880);
239). Dero furcatus (part), Bousfield, 1887; non Dero vaga,
Aulophorus huaronensis Piguet, 1928. Syn: Stieren (non Leidy) (= Aulophorus costatus). Distr:
60
64 ACTA HYDROBIOLOGICA SINICA Vol. 31

ARG: 85. PER: 41, 43. COL: 1. BRA: MS: 1. Furth. gaster limnaei). Syn: Derostoma (part), Ant. Dugès,
Distr: North America (Harman et al., 1988: 2237; 1830; Copopteroma Corda, 1837 (Type: Copopteroma
Kathman & Brinkhurst, 1998: 74). nais Corda, 1837); Mutzia Vogt, 1841 (Type: Mutzia
heterodactyla Vogt, 1841); Vetrovermis Imhof, 1888
Bathydrilus Cook, 1970 (Type: Bathydrilus asy- (Type: Vetrovermis hyalinus Imhof, 1888); not p. p.
mmetricus Cook, 1970). Syn: Macroseta Erséus, 1976 Vaillant, 1890 (= Aeolosoma).
(Type: M. rarisetis Erséus, 1976). Chaetogaster crystallinus Vejdovský, 1883. Distr:
*Bathydrilus argentinae Erséus, 1983. Distr: ARG: 15. Furth. Distr: North America (Kathman &
ARG: 31. Brinkhurst, 1998: 42), Europe, Africa, and Asia (Di
Persia, 1975b: 1).
Bothrioneurum Stolc, 1886 (Type: Bothrio- Chaetogaster diaphanus (Gruithuisen, 1828).
neurum vejdovskyanum). Syn: Nais diaphana Gruithuisen, 1828. Distr: ARG:
*Bothrioneurum americanum Beddard, 1894. 21, 38, 58, 84-86, 89-90. URU: 1. PER: 30. BRA: SP:
Distr: ARG: 22, 34, 58, 63, 85, 86, 89, 90. PAR: 1. 38. Furth. Distr: North America, Europe, and Asia
PER: 4, 5, 22. COL: 1. VEN: 3. GUY: 1. BRA: SP: 39, (Gavrilov, 1979: 104).
40; MS: 1. Chaetogaster diastrophus (Gruithuisen, 1828).
*Bothrioneurum brauni Du Bois-Reymond Syn: Nais distropha Gruithuisen, 1828. Distr: ARG:
Marcus, 1949. Distr: BRA: PA: 5. 58, 86, 89-90. URU: 4. BOL: 1. COL: 1. BRA: SP: 12,
Bothrioneurum iris Beddard, 1901. Distr: PER: 7. 15, 16, 31. Furth. Distr: North America, Europe, and
COL: 1. BRA: SP: 11. Furth. Distr: Asia (Righi, 1984: Asia (Righi, 1984: 20).
39). Chaetogaster langi Bretscher, 1896. Syn: Chae-
*Bothrioneurum righii Collado & Schmelz, 2000. togaster spongillae Annandale, 1906; Chaetogaster
Distr: BRA: AM: 12. parvus Pointner, 1914. Distr: URU: 1. SUR: 4, 9.
Bothrioneurum vejdovskyanum Stolc, 1886. Syn: BRA: SP: 7, 12, 19. Furth. Distr: North America,
Bothrioneurum pyrrhum Marcus, 1942. Distr: ARG: Europe, Africa, and Asia (Gavrilov, 1979: 105).
120. BRA: SP: 12, 14, 16, 17. Furth. Distr: North Chaetogaster limnaei limnaei K. Von Baer, 1827.
America, Europe, Africa, and Asia (Brinkhurst, 1971c: Distr: ARG: 87. URU: 1. BRA: SP: 20; MG: 3. Furth.
541). Distr: North America, Europe, and Asia (Righi, 1984:
21).
Branchiura Beddard, 1892 (Type: Branchiura
sowerbyi). Syn: Kawamuria Stephenson, 1917 (Type: Dero Oken, 1815 (Type: Nais digitata). Syn:
Kawamuria japonica Stephenson, 1917). Nais (part) O. F. Müller, 1774; Dero (Dero) Oken,
Branchiura sowerbyi Beddard, 1892. Distr: 1815; Xantho Dutrochet, 1819; Uronais (part), Gervais,
ARG:1, 58, 85, 86, 99-101. URU: 2. COL:1. BRA: SP: 1838; Proto Ørsted, 1843.
16, 35, 39, 40. Furth. Distr: North America, Europe, *Dero botrytis Marcus, 1943. Syn: Dero (Dero)
and Asia (Righi, 1984: 40); cosmopolitan (Brinkhurst, botrytis Marcus, 1943. Distr: ARG: 35, 58, 86, 96, 108.
1971c: 562). URU: 3. COL: 2. BRA: RS: 1; SP: 5, 12, 39, 40, 42;
PE: 4. Furth. Distr: North America (Milligan, 1997:
Bratislavia Kosel, 1976 (Type: Pristina elegans 74).
Finogenova, 1966). Dero cooperi Stephenson, 1932. Syn: Dero (Dero)
Bratislavia dadayi (Michaelsen, 1905). Syn: cooperi Stephenson, 1932. Distr: ARG: 35, 64, 85.
Naidium dadayi Michaelsen, 1905; Bratislavia Furth. Distr: Europe, Africa, and Asia (Naidu &
unidentata (Harman, 1973); Pristina unidentata Srivastava, 1980: 266).
Harman, 1973. Distr: ARG: 38, 42, 43. SUR: 2, 3, 5, 9. Dero digitata (O. F. Müller, 1773). Syn: Nais
BRA: PR: 2, 3; MS: 1. Furth. Distr: North America digitata O. F. Müller, 1773; Dero (Dero) digitata (O. F.
(Harman & Loden, 1978a; Gavrilov, 1981: 177). Müller, 1773); Nais (Proto) digitata (O. F. Müller,
Chaetogaster Von Baer, 1827 (Type: Chaeto- 1773); Uronais digitata (O. F. Müller, 1773); Proto
Suppl. Christoffersen: A catalogue of aquatic microdrile oligochaetes (Annelida: Clitellata) from South America 61
65

digitata (O. F. Müller, 1773); Xantho hexapoda 1910) (= Aulophorus lodeni). Distr: ARG: 1, 37, 85.
Dutrochet, 1819; Dero limosa Leidy, 1852; ? Dero ECU: 1. BRA: PR: 2, 3; SP: 39, 40, 42, 46; MS: 1.
philippinensis Semper, 1877; Dero acuta Bousfield, Furth. Distr: West Indies (Harman, 1974: 22), North
1886; ? Dero mülleri Bousfield, 1886; ? Dero America (Kathman & Brinkhurst, 1998: 70), India,
intermedia Cragin, 1887; Dero michaelseni Svetlov, Australia (Harman, 1974: 22), and Antarctica
1924; D. bonariensis Michaelsen, 1933; D. kawamurai (Gavrilov, 1981: 178).
Kondô, 1936; Dero tanimotoi Kondô, 1936; Dero Dero plumosa Naidu, 1962. Syn: Dero (Dero)
quadribranchiata Cernosvitov, 1937. Distr: ARG: 22, plumosa Naidu, 1962. Distr: ARG: 38, 42, 44, 109.
30, 42, 58, 85, 86, 89, 90. URU: 4. PAR: 1. PER: 40, BRA: SP: 35. Furth. Distr: Grenada (Righi & Hamoui,
42. ECU: 1. COL: 1. VEN: 21. GUY: 1. SUR: 2-4, 6, 2002: 132) and India (Di Persia, 1976: 6).
9, 16. BRA: SP: 38-40, 47; MS: 1; PA: 1, 2, 4, 6, 7, 10, Dero raviensis (Stephenson, 1914). Syn: ? Nais
12, 15, 17, 18; AM: 7. Furth. Distr: Central America tenuidentis Walton, 1906; Nais raviensis Stephenson,
(Stacey & Coates, 1996: 23), North America and 1914; Dero (Dero) raviensis (Stephenson, 1914). Distr:
Philippine Islands (Cordero, 1931: 350). ARG: 95. PER: 45. ECU: 1. COL: 1. SUR: 1, 4, 5, 9.
*Dero evelinae Marcus, 1943. Syn: Dero (Dero) BRA: SP: 38. Furth. Distr: Caribbean (Righi &
evelinae Marcus, 1943. Distr: ARG: 38, 86. URU: 3. Hamoui, 2002: 134), Africa and Asia (Gavrilov, 1981:
VEN: 21. SUR: 15. BRA: RS: 2; PR: 6; SP: 11; AL: 2; 179).
PE: 5, 6; PI: 1; PA: 2, 5, 15. *Dero righii Varela, 1990. Syn: Dero (Dero)
*Dero magna Harman, 1974. Syn: Dero (Dero) righi Varela, 1990. Distr: ARG: 1, 34, 84 85, 89, 90,
magna Harman, 1974. Distr: ARG: 86. SUR: 2, 7. 93. BRA: MS: 1.
Dero multibranchiata Stieren, 1892. Syn: ? Dero Dero sawayai Marcus, 1943. Syn: ? Dero
florifera Oken, 1815. Distr: ARG: 1, 60, 86, 96. BOL: heterobranchiata Michaelsen, 1933; Dero (Dero)
5. SUR: 15, 16. BRA: SP: 40; MS: 1; PE: 14; PA: 1, 4, sawayai Marcus, 1943. Distr: ARG: 58, 85, 86, 89, 90.
9, 10, 13, 14; AM: 4, 7, 12; RR: 2. Furth. Distr: ECU: 1. VEN: 7. SUR: 2, 3, 7, 11, 16. BRA: SP: 15,
Central America (Stieren, 1892: 107; Righi & Hamoui, 33, 38; AL: 2; PE: 3; CE: 2. Furth. Distr: Central
2002: 131). America and India (Stacey & Coates, 1996: 23).
Dero nivea Aiyer, 1930. Syn: Dero (Dero) nivea Dero trifida Loden, 1979. Distr: SUR: 11. Furth.
Aiyer, 1930; Dero palestinica Černosvitov, 1938. Distr: Aruba (Righi & Hamoui, 2002: 130) and North
Distr: ARG: 15, 23, 24, 32, 37, 38, 44, 63, 85, 86, 89, America (Loden, 1979: 584).
90, 92. PER: 28. SUR: 3, 4, 6, 9. BRA: SP: 35, 38-40. *Dero tuna Righi & Hamoui, 2002. Distr: SUR:
Furth. Distr: North America, Europe, Africa, Asia, and 16.
Australia (Harman, 1974: 20). Epirodrilus Hrabe, 1930 (Type: Epirodrilus mi-
Dero obtusa D´Udekem, 1855. Syn: ? Xantho chaelseni Hrabe, 1930).
hexapoda Dutrochet, 1819; Dero perrieri Bousfield, *Epirodrilus antipodum Černosvitov, 1939. Distr:
1886; Dero latissima Bousfield, 1886; Dero communis PER: 7, 8, 12.
Golanski, 1911; Dero polycardia Hayden, 1922. Distr:
ARG: 35, 37, 58, 76-78, 85, 86, 89, 90, 106. BOL: 2, Haemonais Bretscher, 1900 (Type: Haemonais
4. VEN: 21. SUR: 2-4, 7-10, 15, 16. BRA: SP: 15, waldvogeli).
38-40. Furth. Distr: Central America, North America, Haemonais waldvogeli Bretscher, 1900. Syn:
Europe, Middle East, Africa, and Asia (Harman, 1974: Haemonais laurentii Stephenson, 1915; ? Haemonais
21; Righi & Hamoui, 2002: 128). ciliata Hayden, 1922. Distr: ARG: 1, 42. SUR: 2-5, 9.
Dero palmata Aiyer, 1929. Syn: Dero (Dero) BRA: PR: 2, 3; SP: 38-40, 46; AL: 4; PA: 7. Furth.
palmata Aiyer, 1929. Distr: ARG: 85, 86. GUY: 1. Distr: Central America (Righi & Hamoui, 2002: 126),
Furth. Distr: Costa Rica and India (Stacey & Coates, North America, Europe, possibly Africa, and Asia
1996: 23). (Gavrilov, 1979: 105).
Dero pectinata Aiyer, 1930. Syn: Dero (Dero) Ilyodrilus Eisén, 1879 (Type: Ilyodrilus perrieri
pectinata Aiyer, 1930; non Dero pectinata (Sephenson, Eisén, 1879).
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66 ACTA HYDROBIOLOGICA SINICA Vol. 31

Ilyodrilus frantzi Brinkhurst, 1965. Syn: Ilyodrilus *Limnodriloides validus Erséus, 1982. Distr:
frantzi typica Brinkhurst & Cook, 1966; Ilyodrilus URU: 9.
frantzi capillatus Brinkhurst & Cook, 1966. Distr:
ARG: 38. Furth. Distr: Western USA (Brinkhurst, Limnodrilus Claparède, 1862 (Type: Limnodrilus
1971c: 498). hoffmeisteri). Syn: Camptodrilus Eisén, 1879 (Type:
Camptodrilus spiralis Eisén, 1894); Pododrilus (part),
Isochaetides Hrabe, 1966 (Type: Limnodrilus Czerniavsky, 1880.
baicalensis). Limnodrilus claparedeianus Ratzel, 1868. Syn:
Isochaetides baikalensis (Michaelsen, 1901). Syn: Clitellio claparedeianus (Ratzel, 1868); Tubifex clapare-
Limnodrilus baicalensis Michaelsen, 1901; Isochaeta deianus (Ratzel, 1868); Camptodrilus igneus Eisén, 1879;
baicalensis (Michaelsen, 1901). Distr: PER: 7 Furth. Limnodrilus igneus (Eisén, 1879); Clitellio (Limnodrilus)
Distr: Lake Baical (Gavrilov, 1979: 107). igneus (Eisén, 1879); Limnodrilus longus Bretscher,
*Isochaetides lacustris (Černosvitov, 1939). Syn: 1901; Limnodrilus subsalsus (part) Moore, 1905;
Tubifex lacustris Černosvitov, 1939. Distr: PER: 9, 11. ? Limnodrilus gracilis Moore, 1906; Limnodrilus
motomurai Nomura, 1929. Distr: ARG: 1, 14, 42, 45,
Jolydrilus Marcus, 1965 (Type: Jolydrilus jau- 46, 124, 126. URU: 5. COL: 1, 2. Furth. Distr: North
lus). America, Europe, and Asia (Gavrilov & Paz de
*Jolydrilus jaulus Marcus, 1965. Distr: BRA: SP: Tomsic, 1950: 563).
8, 10, 11. Limnodrilus hoffmeisteri Claparède, 1862. Syn:
Limnodrilus hoffmeisteri f. typica Claparède, 1862;
Krenedrilus Dumnicka, 1983 (Type: Krenedrilus Clitellio hoffmeisteri (Claparède, 1862); Clitellio (Lim-
papillatus). nodrilus) hoffmeisteri (Claparède, 1862); Tubifex
*Krenedrilus papillatus Dumnicka, 1983. Distr: hoffmeisteri (Claparède, 1862); Camptodrilus spiralis
VEN: 10. Eisén, 1879; Limnodrilus spiralis (Eisén, 1879);
Camptodrilus californicus Eisén, 1879; Limnodrilus
Limnodriloides Pierantoni, 1903 (Type: Limno- californicus (Eisén, 1879); Clitellio (Limnodrilus) cali-
driloides appendiculatus Pierantoni, 1903). Syn: fornicus (Eisén, 1879); Camptodrilus corallinus Eisén,
Bohadschia Hrabe, 1971 (Type: Bohadschia pieran- 1879; Clitellio (Limnodrilus) corallinus (Eisén, 1879);
toni Hrabe, 1971). Limnodrilus corallinus (Eisen, 1879); ? Limnodrilus
*Limnodriloides basilicus Finogenova, 1986. steigerwaldi Eisén, 1879; Limnodrilus dugesi Rybka,
Distr: PER: 50. 1898; Limnodrilus hoffmeisteri f. dugesi (Rybka, 1898);
*Limnodriloides clavellatus Finogenova, 1986. Limnodrilus gotoi (part) Hatai, 1899; Limnodrilus
Distr: PER: 49. lucasi Benham, 1903; Limnodrilus vejdovskyanus
Limnodriloides claviger Erséus, 1982. Distr: Benham, 1903; Limnodrilus subsalsus (part) Moore,
VEN: 15. Furth. Distr: West Indies (Erséus, 1982: 1905; Limnodrilus aurostriatus Southern, 1909;
221). Limnodrilus aurantiacus Friend, 1911; Limnodrilus
Limnodriloides monothecus Cook, 1974. Syn: hoffmeisteri var. tenellulus Friend, 1912; Limnodrilus
Bohadschia monotheca (Cook, 1974). Distr: VEN: 15. socialis Stephenson, 1912; Limnodrilus hoffmeisteri f.
Furth. Distr: West Indies, Gulf of Mexico to New socialis Stephenson, 1912; Limnodrilus parvus var.
Jersey, Mediterranean Sea, and Pacific coast of USA biannulatus Lastockin, 1927; Limnodrilus pacificus
(Erséus, 1982: 252). Chen, 1940; Limnodrilus hoffmeisteri f. divergens
Limnodriloides rubicundus Erséus, 1982. Distr: Marcus, 1942; Limnodrilus subsalus, Marcus, 1944;
VEN: 16. Furth. Distr: West Indies, East coast of Limnodrilus parvus, Du Bois-Reymond Marcus, 1947,
United States, down to 74 m depth (Erséus, 1982: Gavrilov & Paz de Tomsic, 1950; Limnodrilus
274). hoffmeisteri f. parva, Marcus, 1942. Distr: CHI: 2.
*Limnodriloides sphaerothecus Erséus, 1982. ARG; 8, 14-16, 18, 22, 38, 42-48, 51, 55-58, 63, 72,
Distr: SUR: 17. 76-79, 85, 86, 89, 90, 125. PER: 7, 12, 20, 33-35, 39,
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67

47. COL: 1, 2, 8, 9, 11. GUY: 1. BRA: RS: 3; PR: 1; URU: 6. Furth. Distr: Cosmopolitan (Brinkhurst,
SP: 11, 29, 35, 39, 40, 42, 43, 46, 47. PA: 10. Furth. 1971c: 557).
Distr: Central America, North America, Europe, and
Asia (Gavrilov, 1950: 563). Rem: Cosmopolitan; Limno- Nais O. F. Müller, 1774 (Type: Nais barbata O. F.
drilus hoffmeisteri is perhaps the most ubiquitous species Müller, 1774). Syn: Opsonais Gervais, 1838 (Type:
in the subfamily Tubificinae, worldwide (Mark Wetzel, Nais elinguis O. F. Müller, 1774).
pers. comm.). Nais andina andina Černosvitov, 1939. Distr:
*Limnodrilus neotropicus Černosvitov, 1939. PER: 12, 14, 15, 18, 20. COL: 1. Furth. Distr: India
Syn: Isochaetides neotropicus (Černosvitov, 1939). (Naidu & Srivastava, 1980: 265).
Distr: ARG: 22, 85. PER: 7, 36. COL: 1. BRA: SP: 43, *Nais andina pectinata Martínez & Giani, 1986.
44. Distr: BOL: 3.
Limnodrilus udekemianus Claparède, 1862. Syn: Nais bretscheri Michaelsen, 1899. Distr: ARG:
Clitellio udekemianus (Claparède, 1862); Clitellio 14, 86, 89, 90. VEN: 18. Furth. Distr: North America
(Limnodrilus) udekemianus (Claparède, 1862); Tubifex (Kathman & Brinkhurst, 1998: 86), Europe and Asia
udekemianus (Claparède, 1862); Limnodrilus ornatus (Marchese, 1986: 244).
Eisén, 1879; ? Limnodrilus steigerwaldii Eisén, 1879; *Nais carolina Blanchard, 1849. Distr: CHI; 4.
? Clitellio (Limnodrilus) steigerwaldii (Eisén, 1879); PER: 1.
Clitellio (Limnodrilus) ornatus (Eisén, 1879); Limnodrilus Nais communis Piguet, 1906. Syn: Nais hetero-
wordsworthianus Friend, 1898; Limnodrilus udekemianus chaeta Benham, 1893; ? Nais. parvula Walton, 1906;
var. wordsworthianus (Friend, 1898); Limnodrilus willeyi Nais parviseta Walton, 1906; Nais variabilis var.
Nomura, 1913; Limnodrilus inversus Gavrilov & Paz de punjabensis Stephenson, 1909; Nais communis var.
Tomsic, 1949. Distr: CHI: 2. ARG: 7, 14, 58, 63, 77, punjabensis (Stephenson, 1909); Nais communis var.
78, 81, 83, 85, 86, 121. PER: 26. COL: 1, 2, 10. VEN: caeca Stephenson, 1910; Pterochaeta astronensis
1. BRA: SP: 16, 40, 43. Furth. Distr: Cosmopolitan Pierantoni, 1911; ? Nais communis var. acuta Pointner,
(Brinkhurst, 1971c: 468). 1914; Nais communis f. magenta Marcus, 1943. Distr:
ARG: 2, 7, 37, 39, 41, 42, 55, 63, 85, 86, 89, 90. PER:
Monopylephorus Levinson, 1884 (Type: Mono- 9, 11, 32, 39. ECU: 1. COL: 1, 2, 11. BRA: SP: 3, 15,
pylephorus rubroniveus Levinsen, 1884). Syn: 36, 38, 43, 46, 47; AL: 5; PA: 5. Furth. Distr:
Vermiculus Goodrich, 1892 (Type: Vermiculus pilosus Cosmopolitan (Brinkhurst, 1971d: 330).
Goodrich, 1892); Postiodrilus Bolt, 1926 (Type: Nais elinguis O. F. Müller, 1774. Syn: Opsidonais
Postiodrilus sonderi Bolt, 1926); Littodrilus Chen, elinguis (O. F. Müller, 1774); Nais aralensis
1940 (Type: Rhizodrilus aucklandicus Benham, 1909). Lastockin, 1922. Distr: ARG: 4, 14, 46. URU: 1. PER:
*Monopylephorus moleti Brinkhurst & Marchese, 17, 31. COL: 1. BRA: SP: 38. Furth. Distr:
1987. Distr: ARG: 65, 85. Cosmopolitan (Gavrilov, 1979: 105).
Monopylephorus parvus Ditlevsen, 1904. Syn: Nais pardalis Piguet, 1906. Syn: Nais bretscheri
Rhizodrilus parvus (Ditlevsen, 1904). Distr: BRA: SP: var. pardalis Piguet, 1906; Nais lastockini Sokolskaya,
8. Furth. Distr: Africa and Asia (Erséus, 2002: 198). 1958. Distr: CHI: 3. ARG: 14, 37, 42, 46, 48, 63
Monopylephorus rubroniveus Levinsen, 1884. PER: 18, 32, 40, 43. COL: 1. BRA: SP: 38, 46. Furth.
Syn: Vermiculus pilosus Goodrich, 1892; Rhizodrilus Distr: North America (Kathman & Brinkhurst, 1998:
pilosus (Goodrich, 1892); Vermiculus fluviatilis Ferronière, 86), Europe, and Asia (Gavrilov, 1979: 106).
1899; ? Vermiculus glotini Ferronière, 1899; Monopy- Nais pseudobtusa Piguet, 1906. Syn: Nais obtusa
lephorus glaber Moore, 1905; Rhizodrilus glaber (Moore, var. pseudobtusa Piguet, 1906; ? Pristina variabilis
1905); Rhizophorus kermadecensis Benham, 1915; Friend, 1916; ? Nais incerta Černosvitov, 1942. Distr:
Monopylephorus corderoi Du Bois-Reymond Marcus, COL: 1. Furth. Distr: North America (Kathman &
1949; Rhizodrilus ponticus Hrabe, 1967; Monopy- Brinkhurst, 1998: 84), Europe, Africa, and Afghanis-
lephorus helobius Loden, 1980; Monopylephorus tan (Brinkhurst, 1971d: 340).
rubroniveus ssp. iturupi Finogenova, 1982. Distr: *Nais schubarti Marcus, 1944. Distr: BRA: PR:
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68 ACTA HYDROBIOLOGICA SINICA Vol. 31

2, 3; PE: 3. evelinae Marcus, 1965. BRA: SP: 8.


Nais simplex Piguet, 1906. Syn: Nais variabilis Paranais filiformis (Schmarda, 1861). Syn: Chaetogaster
var. simplex Piguet, 1906; ? Nais tortuosa Walton, filiformis Schmarda, 1861; Schmardaella filiformis
1906; ? Nais sp. Stephenson, 1909. Distr: PER: 21. (Schmarda, 1861). Distr: CHI: 3. ECU: 2. Furth. Distr:
COL: 1, 2. Furth. Distr: North America (Kathman & Japan (Kondô, 1936: 383).
Brinkhurst, 1998: 82), Europe, Africa, and possibly Paranais frici Hrabe, 1941. Syn: Paranais
Tibet (Harman et al., 1988: 2235). mobilis Liang, 1958; Wapsa moliblis (Liang, 1958).
Nais variabilis Piguet, 1906. Syn: ? Nais rivulosa Distr: ARG: 63. Furth. Distr: North America, Europe,
Leidy, 1850; ? Nais japonica Kondô, 1936; ? Nais Africa (Brinkhurst & Coates, 1985: 30) and Asia
bekmani Sokolskaya, 1962; Distr: ARG: 2, 58, 26, 63, (Liang, 1958: 43; Sokolskaya, 1971: 930).
85, 89, 90. BOL: 2, 3. PER: 22, 32. COL: 1, 5. BRA: *Paranais lacteus Černosvitov, 1937. Distr: ARG:
MS: 1. Furth. Distr: Cosmopolitan (Gavrilov, 1981: 26. Rem: Brinkhurst (1971d: 323) transferred this
179). species to Homochaeta lacteus (Černosvitov, 1937).
However, Timm & Grimm (2005) have determined
Olavius Erséus, 1985 (Type: Phallodrilus long- that the genus Homochaeta, and in particular the taxon
issimus Giere, 1979). Homochaeta naidina Bretscher, 1896 − the type
*Olavius bullatus Finogenova, 1986. Distr: PER: species of Homochaeta – should not be included in
48. routine identification keys of Oligochaeta for the
*Olavius crassitunicatus Finogenova, 1986. Distr: following reasons: no type material exists for this
PER: 51, 52. taxon; all subsequent material identified as this taxon,
when available for review, had been misidentified; the
Ophidonais Gervais, 1838 (Type: Nais serpen- original description by Bretscher likely was based
tina). Syn: Serpentina Ørsted, 1843 (Type: Serpentina upon different, immature specimens in the families
quadristriata Ørsted, 1843). Naididae and Tubificidae; the taxon has never been
Ophidonais serpentina (O. F. Müller, 1774). Syn: re-described. Paggi et al. (2006: 1) cited Homochaeta
Nais serpentina O. F. Müller, 1774; Slavinia naidina, while Gómez & Rodrigues (2005: 35) cited
serpentina (O. F. Müller, 1774); Pristina serpentina both Homochaeta naidina and H. lacteus for Argentina.
(O. F. Müller, 1774); ? Nais filiformis Blainville, 1825; Paranais litoralis (O. F. Müller, 1784). Syn: Nais
Serpentina quadristriata Ørsted, 1843; Ophidonais litoralis O. F. Müller, 1784; Blanonais litoralis (O. F.
reckei Floericke, 1892; Nais reckei (Floericke, 1892); Müller, 1784); Uncinais litoralis (O. F. Müller, 1784);
Ophidonais serpentina var. meridionalis Piguet, 1906; Enchytraeus triventralopectinatus Minor, 1863;
Ophidonais serpentina f. typica Michaelsen, 1909; Paranais litoralis var. intermedia Czerniavsky, 1880;
Ophidonais sp. Kleiber, 1911. Distr: ARG: 14, 37. ?Paranais litoralis ssp. orientalis Sokolskaya, 1964;
URU: 7. Furth. Distr: North America (Kathman & Paranais orientalis Sokolskaya, 1964. Distr: ARG: 39,
Brinkhurst, 1998: 50), Europe, and Asia (Gavrilov, 40. Furth. Distr: North America, Europe, Africa, and
1979: 106). Asia (Brinkhurst, 1971d: 318).
*Paranais macrochaeta Černosvitov, 1939. Distr:
Paranadrilus Gavrilov, 1955 (Type: Parana- ARG: 18. PER: 17.
drilus descolei). *Paranais salina Černosvitov, 1939. Distr: PER:
*Paranadrilus descolei Gavrilov, 1955. Distr: 10.
ARG: 1, 38, 59, 67-71, 83, 85, 91, 103. PER: 45. GUY:
1. BRA: PR: 2, 3; MS: 1. Pedonais Collado & Schmelz, 2000 (Type: Pedonais
crassifaucis).
Paranais Czerniavsky, 1880 (Type: Nais lito- *Pedonais crassifaucis Collado & Schmelz, 2000a:
ralis). Syn: Wapsa Marcus, 1965 (Type: Wapsa 225. Distr: BRA: AM: 13.
evelinae). Peloscolex Leidy, 1851 (Type: Peloscolex varie-
*Paranais evelinae (Marcus, 1965). Syn: Wapsa gatus Leidy, 1851).
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61

*P. orghidani Botea, 1987. Distr: VEN: 19. Furth. Distr: Central America (Stacey & Coates, 1996:
*Peloscolex pigueti Michaelsen, 1933. Syn: 24), North America (Kathman & Brinkhurst, 1998:
Trachydrilus plicatus Piguet, 1928. Distr: ARG: 20. 58), Europe, and India (Di Persia, 1973: 281).
PER: 37. *Pristina aequiseta foreli Piguet, 1906. Syn:
Pristina aequiseta f. foreli Piguet, 1906; Naidium
Potamothrix Vejdovský & Mrázek, 1902 (Type: foreli (Piguet, 1906). Distr: BOL: 3.
Potamothrix moldavensi Vejdovský & Mrázek, 1902). Pristina americana Černosvitov, 1937. Syn: Pristina
Syn: Euilyodrilus Brinkhurst, 1963 (Type: Ilyodrilus americana f. typica Černosvitov, 1937; Pristina americana
hammoniensis Michaelsen, 1903). var. loretana Černosvitov, 1937; Pristina peruviana
Potamothrix bavaricus (Öschmann, 1913). Syn: Černosvitov, 1939; Pristina longidentata Harman, 1965;
Tubifex (Ilyodrilus) bavaricus Öschmann, 1913; Eui- Pristinella longidentata (Harman, 1965); Pristina org-
lyodrilus bavaricus (Öschmann, 1913); Ilyodrilus hidani Botea, 1983. Distr: ARG: 1, 15, 26, 63, 85, 86,
bavaricus (Öschmann, 1913). Distr: PER: 2, 16, 18, 89, 90. URU: 2. PER: 2, 16, 18, 22, 38. VEN: 4, 5.
20, 23, 24. Furth. Distr: Cosmopolitan (Brinkhurst & GUY: 1. SUR: 2, 3, 8, 9, 16. BRA: PR: 1-3; SP: 13, 16,
Marchese, 1987: 3157). 39, 40, 43, 46, 47; MS: 1; AL: 4; PA: 1, 2, 6, 7, 9, 10,
Potamothrix heuscheri (Bretscher, 1900). Syn: 14; AM: 1, 4. Furth. Distr: Central America (Stacey &
Tubifex heuscheri Bretscher, 1900; Tubifex (Ilyodrilus) Coates, 1996: 24), North America (Kathman &
heuscheri Bretscher, 1900; Ilyodrilus heuscheri Brinkhurst, 1998: 60), Europe, Africa, and New
(Bretscher, 1900); Euilyodrilus heuscheri (Bretscher, Zealand (Stacey & Coates, 1996: 24).
1900); Ilyodrilus orientalis Černosvitov, 1938; Pristina bilobata (Bretscher, 1903). Syn: ? Naidium
Euilyodrilus orientalis (Černosvitov, 1938). Distr: uniseta Bretscher, 1900; ? Pristina uniseta (Brestscher,
PER: 2. Furth. Distr: Europe, Africa, and Middle East 1900); Pristinella bilobata (Brestscher, 1903); Nai-
(Brinkhurst, 1971c: 490). dium bilobatum Bretscher, 1903. Distr: ARG: 35, 83.
Furth. Distr: Europe, Africa, Middle East, and Asia
Pristina Ehrenberg, 1828 (Type: Pristina longiseta (Brinkhurst, 1971d: 393).
Ehrenberg, 1828). Syn: Naidium O. Schmidt, 1847 Pristina biserrata Chen, 1940. Distr: ARG: 33,
(Type: Naidium luteum O. Schmidt, 1847); Pristinella 37, 86. ECU: 1. GUY: 1. Furth. Distr: East Asia
Brinkhurst, 1985 (Type: Naidium bilobatum). (Stacey & Coates, 1996: 24).
Pristina acuminata Liang, 1958. Syn: Pristinella Pristina breviseta Bourne, 1891. Syn: Naidium
acuminata (Liang, 1958). Distr: ARG: 58, 63. PER: breviseta (Bourne, 1891). Distr: ARG: 8, 14, 46, 49,
29, 38. Furth. Distr: North America and China 58, 60, 83, 85. BRA: SP: 15, 16; PA: 5, 8. Furth. Distr:
(Harman et al., 1988: 2239). North America (Kathman & Brinkhurst, 1998: 60),
Pristina aequiseta aequiseta Bourne, 1891. Syn: and Asia (Gavrilov, 1981: 179).
? Pristina inaequalis (Ehrenberg, 1828); ? Pristinais *Pristina cumanacoensis Botea, 1987. Distr:
inaequalis (Ehrenberg, 1828); Pristina aequiseta VEN: 21.
Bourne, 1891; Pristina aequiseta f. aequiseta Bourne, *Pristina decui Botea, 1987. Distr: VEN: 4.
1891; Naidium aequiseta (Bourne, 1891); Pristina Pristina longiseta Ehrenberg, 1828. Syn: Stylaria
affinis Garbini, 1898; Naidium tentaculatum Piguet, longiseta (Ehrenberg, 1828); Pristina longiseta f.
1906; Pristina tentaculatum (Piguet, 1906); Pristina typica Ehrenberg, 1828; Pristina longiseta longiseta
aequiseta var Michaelsen, 1914; Pristina palustris Ehrenberg, 1828; Pristina leidyi Smith, 1896; Pristina
Schuster, 1915; ? Pristina capiliseta Kondô, 1936; longiseta leidyi (Smith, 1896); Pristina longiseta f.
? Pristina nasalis Kondô, 1936; Pristina evelinae leidyi (Smith, 1896); Pristina leidyi typica Michaelsen,
Marcus, 1943; Pristina sperberae Naidu, 1963. Distr: 1914; Pristina longiseta var. bidentata Černosvitov,
CHI: 6. ARG: 14, 35, 37, 43, 49, 55, 58, 62, 74, 85, 86, 1942; Pristina bidentata Černosvitov, 1942; Pristina
89, 90. BOL: 2. PER: 18, 20, 22, 27, 40-42, 47. ECU: longiseta sinensis Sperber, 1948; Pristina orghidani
1. COL: 1, 2, 11. VEN: 20, 21. GUY: 1. SUR: 2-5, 9, Botea, 1983. Distr: CHI: 0. ARG: 8, 14, 15, 25, 32, 35,
14. BRA: SP: 4, 6, 13, 14, 21, 34, 44; MS: 1; PA: 3. 37, 43, 51, 58, 61, 73-75, 79, 85, 86, 89, 90, 98, 110,
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70 ACTA HYDROBIOLOGICA SINICA Vol. 31

112, 118, 120. URU: 4. PAR: 1. BOL: 2, 3. PER: 21, Pristina jenkinae (Stephenson, 1931).? Pristina rosea,
27, 28, 32, 40, 46. COL: 1. VEN: 4, 5. SUR: 2, 3, 9. Michaelsen & Boldt, 1932, Kondô, 1936, Yamaguchi,
BRA: SP: 3, 6, 35, 38; AL: 6; PE: 3; AM: 4. Furth. 1953 (non Piguet); Naidium roseum, Marcus, 1943
Distr: Central America (Marchese, 1986: 242), North (non Piguet); Pristina idrensis Sperber, 1948; Pristi-
America, Europe, Asia, Australia (Gavrilov, 1979: nella idrensis (Sperber, 1948); Pristina taita Stout,
106), and Hawaii (Harman et al., 1988: 2239). 1956; Pristina nothophagi Stout, 1958. Distr: ARG: 8,
Pristina macrochaeta Stephenson, 1931. Distr: 17, 39, 42-44, 51, 58, 85, 86, 89, 90. PER: 26, 38.
ARG: 1, 33, 61, 63, 85, 86, 92, 109, 111, 112, URU: 1. COL: 1. VEN: 4, 5. GUY: 1. SUR: 12. BRA: SP: 13,
PAR: 2, 4. PER: 27. COL: 1, 2, 4, 6. GUY: 1. SUR: 2, 14, 36, 44-46; PE: 9, 12; AM: 1. Furth. Distr: Central
4, 7. BRA: SP: 13, 32; MS: 1; PE:3, 6, 8; CE: 3. America (Marchese, 1986: 241), North America,
Further Distr: Central America (Righi & Hamoui, Europe, Africa, Asia (Harman et al., 1988: 2239), and
2002: 152) and Afghanistan (Brinkhurst, 1971d: 406). possibly New Zealand (Brinkhurst, 1971d: 394). Rem:
*Pristina marcusi Collado & Schmelz, 2001. Pristinella idrensis was suppressed and synonymyzed
Distr: BRA: AM: 12. with P. jenkinae in Kathman (1985: 1024).
Pristina menoni (Aiyer, 1929). Syn: ? Naidium *Pristina silvicola Collado & Schmelz, 2000b.
mosquensis Udalicov, 1907; ? Naidium heteroseta Udalicov, Distr: BRA; AM: 15.
1907; Naidium menoni Aiyer, 1929; Pristinella menoni Pristina sima (Marcus, 1944). Syn: Naidium
(Aiyer, 1929). Distr: ARG: 1, 35, 85, 106. COL: 1. VEN: simum Marcus, 1944; Pristinella sima (Marcus, 1944).
7-9. SUR: 7. BRA: PR: 2, 3; SP: 36, 46. Futh. Distr: Distr: CHI: 1. ARG: 1, 33, 86. COL: 1. GUY: 1. BRA:
Europe, Africa, and India (Di Persia, 1973: 283). SP: 22, 23, 27, 35. Furth. Distr: Curaçao (Righi &
Pristina notopoda Černosvitov, 1937. Syn: Pristinella Hamoui, 2002: 146), North America and Africa
notopoda (Černosvitov, 1937). Distr: ARG: 28, 53, 85. (Stacey & Coates, 1996: 24).
PER: 40. GUY: 1. Furth. Distr: Europe and Africa Pristina synclites Stephenson, 1925. Distr: ARG:
(Stacey & Coates, 1996: 24). 8, 35, 58, 86, 98, 106, 110. ECU: 1. COL: 2, 8. GUY:
Pristina osborni (Walton, 1906). Syn: Naidium 1. SUR: 8, 15. BRA: SP: 43, 46; MS: 1. Furth. Distr:
osborni Walton, 1906; Pristinella osborni (Walton, 1906); Central America (Righi & Hamoui, 2002: 148), North
Naidium minutum Stephenson, 1914; Pristinella minuta America, Africa, India (Stephenson, 1925: 45;
(Stephenson, 1914); Pristina minuta (Stephenson, 1914). Marchese, 1986: 243) and Indonesia (Ohtaka et al.,
Distr: ARG: 6, 19, 38, 58, 79, 81, 83, 85, 86, 89. 90, 2000: 29).
106, 110, 123. PER: 31, 39. VEN: 13, 21. SUR: 13, 14. *Pristina terrena Collado & Schmelz, 2000b.
BRA: SP: 1, 6; MG: 1, 4; MS: 1; AM: 5, 14. Furth. Distr: BRA: AM: 15.
Distr: Central America (Marchese, 1986: 240), North *Pristina trifida Collado & Schmelz, 2002. Distr:
America, Africa, and Asia (Gavrilov, 1979: 106). BRA: AM: 1.
Pristina proboscidea Beddard, 1896. Syn: Pristina
proboscidea f. typica (Beddard, 1896); Pristina pro- Slavina Vejdovský, 1883 (Type: Nais appendicu-
boscidea var. paraguayensis Michaelsen, 1905; ? Pristina lata).
serpentina Walton, 1906. Distr: CHI: 5. ARG: 27, 33, Slavina appendiculata (D’Udekem, 1855). Syn:
58, 67, 85, 86, 89, 90. PAR: 1. PER: 42. COL: 1, 4-7. ? Nais escherosa Gruithuisen, 1828; ? Nais gracilis
SUR: 2, 3. BRA: SP: 44; MS: 1; CE: 3; PA: 4, 10, 18; Leidy, 1850; ? Slavina gracilis (Leidy, 1850); Stylaria
AM: 4. Furth. Distr: USA (Milligan, 1997: 59), Africa, gracilis (Leidy, 1850); Nais appendiculata D’Udekem,
Asia, and Australia (Harman, 1974: 15). 1855; Nais lurida Timm, 1883; Slavina lurida (Timm,
Pristina rosea (Piguet, 1906). Syn: Naidium 1883); Slavina punjabensis Stephenson, 1909; ? Slavina
roseum Piguet, 1906; ? Naidium luteum, Michaelsen, sp. Stephenson, 1916; ? Slavina montana Stephenson,
1905 (non Schmidt); Naidium jenkinae Stephenson, 1923; Slavina truncata Harman, 1966. Distr: ARG: 37,
1931; Pristinella jenkinae (Stephenson, 1931); 52, 85, 86. URU: 8. COL: 1. GUY: 1. BRA: PR: 6; SP:
Naidium roseum f. jenkinae (Stephenson, 1931); 19, 45. Furth. Distr: Central America, North America,
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61

Europe, Africa, Asia, and New Zealand (Marchese, Stephensoniana trivandrana (Aiyer, 1926). Syn:
1986: 236). Naidium trivandranum Aiyer, 1926; Stephensonia
Slavina evelinae (Marcus, 1942). Syn: Peloscolex trivandrana (Aiyer, 1926); Slavinia trivandrana (Ai-
evelinae Marcus, 1942. Distr: ARG: 1, 11-13, 38, 54, yer, 1926). Distr: ARG: 50, 57, 58, 86. SUR: 9, 16.
58, 85, 86, 89, 90, 106, 114. GUY: 1. SUR: 4, 9. BRA: COL: 1. BRA: PR: 2, 3; MS: 1. Furth. Distr: Central
SP: 13, 19, 25, 26, 39, 40; MS: 1; PE: 4; CE: 2; PA: 3, America (Righi & Hamoui, 2002:143), North America
7-9, 13-16. Furth. Distr: Central America (Stacey & (Kathman & Brinkhurst, 1998: 90), Africa and Asia
Coates, 1996: 24). (Gavrilov, 1981: 179).
*Slavina isochaeta Černosvitov, 1939. Distr:
ARG: 8, 21, 37, 42, 46, 65, 85, 86, 89, 90. PER: 9, 17. Stylaria Lamarck, 1816 (Type: Nereis lacustris).
BRA: SP: 35. Syn: Stylinais Gervais, 1838 (Type: Nais proboscidea
*Slavina sawayai Marcus, 1944. Distr: ARG: 37, O. F. Müller, 1774); Caecaria Floericke, 1892 (Type:
58, 84, 86. BRA: SP: 6. Caecaria rara Floericke, 1892).
Stylaria fossularis Leidy, 1852. Syn: S. kempi
Smithsonidrilus Brinkhurst, 1966 (Type: Smi- Stephenson, 1916. Distr: ARG: 86, 89, 90. Furth. Distr:
thsonidrilus marinus Brinkhurst, 1966). Syn: Mar- North America (Brinkhurst, 1971d: 354), Africa
cusaedrilus Righi & Kanner, 1979 (Type: Marcusae- (Grimm, 1990: 140) and Korea (Yoon et al., 2000:
drilus hummelincki Righi & Kanner, 1979). 250). Rem: Brinkhurst (1971d: 353) considered the
*Smithsonidrilus peruanus (Finogenova, 1986). differences of S. fossularis and S. lacustris to be so
Syn: Marcusaedrilus peruanus Finogenova, 1986. slight, that he included S. fossularis as a species
Distr: PER: 48. inquirenda (Brinkhurst, 1971d: 354).
Stylaria lacustris (Linnaeus, 1767). Syn: Nereis
Spilodrilus Piguet, 1928. (Type: Spilodrilus stell- lacustris Linnaeus, 1767; Nais lacustris (Linnaeus,
atus). 1767); Nais proboscidea O. F. Müller, 1774; Stylaria
*Spilodrilus stellatus Piguet, 1928. Distr: PER: proboscidea (O. F. Müller, 1774); Stylinais probos-
32. cidea (O. F. Müller, 1774); Stylaria paludosa Lamarck,
1816; Stylaria phyladelphiana Czerniavsky, 1880;
Spirosperma Eisén, 1879 (Type: Spirosperma Stylaria scotica Czerniavsky, 1880; Caecaria rara
ferox Eisén, 1879). Syn: Orientodrilus Holmquist, Floericke, 1892; Caecaria silesica Floericke, 1892;
1978 (Type: Peloscolex nikolskyi Lastokin & Caecaria brevirostris Floericke, 1892. Distr: ARG: 36,
Sokolskaya, 1953). 58. Furth. Distr: North America, Europe, Africa and
Spirosperma velutina (Grube, 1879). Syn: Saenuris Asia (Brinkhurst, 1971d: 353).
velutina Grube, 1879; Peloscolex velutinus (Grube,
1879); Peloscolex velutinus velutinus (Grube, 1879); Tectidrilus Erséus, 1982 (Type: Tectidrilus squ-
Embolocephalus velutinus (Grube, 1879); Spirosperma alidus Erséus, 1982).
(Embolocephalus) velutinus Grube, 1879; Psammoryctes Tectidrilus gabriellae Du Bois Reymond Marcus,
velutinus (Grube, 1879); Tubifex velutinus (Grube, 1950. Syn: Peloscolex gabriellae Du Bois-Reymond
1879); Tubifex (Peloscolex) velutinus (Grube, 1879); Marcus, 1950; Tubificoides gabriellae (Du Bois-
Tubifex sarnensis Pierantoni, 1904; Peloscolex Reymond Marcus, 1950). Distr: BRA: SP: 28. Furth.
velutinus sarnensis (Pierantoni, 1904); Peloscolex Distr: West Indies (Erséus, 1982: 257), Atlantic and
cernosvitovi Hrabe, 1958; Peloscolex fontinalis Hrabe, Pacific coasts of North America (Gavrilov, 1979:
1964. Distr: VEN: 1, 11. Furth. Distr: Europe 107).
(Brinkhurst, 1971c: 507). *Tectidrilus intermixtus Finogenova, 1986. Distr:
PER: 49, 50.
Stephensoniana Černosvitov, 1938 (Type: Naidium Tectidrilus verrucosus (Cook, 1974). Syn:
trivandrana). Syn: Stephensonia Aiyer, 1929 (Type: Limnodrilus verrucosus Cook, 1974. Distr: VEN: 14.
Naidium trivandranum). Furth. Distr: Pacific coast of North America (Erséus,
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60 ACTA HYDROBIOLOGICA SINICA Vol. 31

1982: 261). PER: 19. COL: 2. GUY: 1. BRA: SP: 16, 17, 35, 43.
Furth. Distr: Cosmopolitan (Righi, 1984: 36).
Tubifex Lamarck, 1816 (Type: Lumbricus tubifex).
Syn: Lumbricus (part), O. F. Müller, 1774; Nais (part), Tubificoides Lastockin, 1937 (Type: Limnodrilus
Oken, 1815; Blanonais (part) Gervais, 1838; Saenuris het erochaetus).
(part) Hoffmeister, 1842 (Type: Saenuris variegata Tubificoides heterochaetus (Michaelsen, 1926).
Hoffmeister, 1842); Lophochaeta Stolc, 1886 (Type: Syn: Limnodrilus heterochaetus Michaelsen, 1926;
Lumbricus ignota Stolc, 1886); Heterochaeta Claparède, Peloscolex heterochaetus (Michaelsen, 1926); Peloscolex
1863 (non Westwood, 1841) (Type: Heterochaeta gabriellae var. heterochaetus (Michaelsen, 1926).
costata); Edmondsonia Brinkhurst, 1965 (type: Tubifex Distr: PER: 37. Furth. Distr: North America (Milligan,
minor Sokolskaya, 1961); Frearidrilus Dumnicka, 1996: 218) and Europe (Brinkhurst, 1971c: 521).
1987 (Type: Peloscolex pescei Dumnicka, 1981).
Tubifex tubifex blanchardi Brinkhurst & Jamieson, Uncinais Levinsen, 1884 (Type: Nais uncinata).
1971. Syn: Saenuris sp. Gegenbaur, 1852; Tubifex Uncinais uncinata (Ørsted, 1842). Syn: Nais
alpinus Bretscher, 1900; Tubifex ignotus, Brinkhurst, uncinata Ørsted, 1843; Paranais uncinata (Ørsted,
1960 (non Stolc, 1886); Tubifex tubifex chacoensis, 1843); Ophidonais uncinata (Ørsted, 1843). Distr:
Hrabe, 1966; Tubifex tubifex f. blanchardi Brinkhurst GUY: 1. Furth. Distr: North America (Kathmann &
& Jamieson, 1971. Distr: ARG: 63, 85. Furth. Distr: Brinkhurst, 1998: 50) and Europe (Stacey & Coates,
Europe and Africa (Marchese, 1988: 56). Rem: 1996: 42).
Kathman & Brinkhurst (1998: 142) noted the presence
of Tubifex tubifex ´blanchardi´ form in North NARAPIDAE Righi & Varela, 1983 (Type:
America. Narapa).
*Tubifex tubifex kleerekoperi (Marcus, 1944). Narapa Righi & Varela, 1983 (Type: Narapa
Syn: Limnodrilus kleerekoperi Marcus, 1944; Tubifex bonettoi).
kleerekoperi (Marcus, 1944). Distr: BRA: RS: 3. *Narapa bonettoi Righi & Varela, 1983. Distr:
*Tubifex tubifex siolii (Du Bois-Reymond Marcus, ARG: 38, 58, 83, 85, 86. BRA: PR: 2-4; SP: 45; MS:
1947). Syn: Limnodrilus siolii Du Bois-Reymond 1.
Marcus, 1949; Tubifex siolii (Du Bois-Reymond
Marcus, 1947). Distr: BRA: PA: 3, 5-7, 9, 10, 12, OPISTOCYSTIDAE Černosvitov, 1936 (Type:
13-15, 17; AM: 4, 6, 8; RR: 2. Opistocysta).
Tubifex tubifex tubifex (O. F. Müller, 1774). Syn: Opistocysta Černosvitov, 1936 (Type: Pristina
Lumbricus tubifex O. F. Müller, 1774; Nais tubifex (O. flagellum Leidy, 1880).
F. Müller, 1774); Saenuris tubifex (O. F. Müller, 1774); Opistocysta funiculus Cordero, 1948. Syn:
Tubifex rivolorum Lamarck, 1816; Nais filiformis Ant. Pristina flagellum, Černosvitov, 1936 (non Leidy,
Dugès, 1828; Blanonais filiformis (Ant. Dugès, 1828); 1880); Opistocysta flagellum, Marcus, 1944, Du
Tubifex filiformis (Ant. Dugès, 1837); Saenuris Bois-Reymond Marcus, 1947 (non Leidy, 1880). Distr:
variegata (part) Hoffmeister, 1842; Nais sanguinea ARG: 22, 96. GUY: 1. COL: 1; BRA: SP: 11, 35, 39,
(part) Doyère, 1856; Tubifex bonnetti Claparède, 40; PE: 14; PA: 2, 4, 7, 10; AM: 4, 6. Furth. Distr:
1862; ? Tubifex campanulatus Eisén, 1879; Saenuris Possibly Africa (Gavrilov, 1979: 106). Rem:
taurica Czerniavsky, 1880; Saenuris peculiaris Czer- According to Kathman & Brinkhurst (1998: 28), O.
niavsky, 1880; Saenuris diversisetosa f. charcoviensis flagellum is considered a species inquirenda.
Czerniavsky, 1880; Saenuris diversisetosa f. suchumica *Opistocysta serrata Harman, 1970. Syn:
Czerniavsky, 1880; Tubifex (Tubifex) fontaneus Pointner, Opistocysta flagellum, Cordero, 1948 (non Leidy,
1914; Tubifex hattai Nomura, 1926; Limnodrilus 1880); Pristina flagellum, Michaelsen, 1905 (non
kleerekoperi Marcus, 1944; Limnodrilus siolii Du Leidy, 1880). Distr: ARG: 85. PAR: 1. BRA: PR: 2, 3;
Bois-Reymond-Marcus, 1947. Distr: CHI: 2. ARG: 33. MS: 1.
Suppl. Christoffersen: A catalogue of aquatic microdrile oligochaetes (Annelida: Clitellata) from South America 73
61

Phreodrilus kerguelensis Michaelsen, 1903).


Trieminentia Harman & Loden, 1978b (Type: Nesodrilus crozetensis (Michaelsen, 1905). Syn: Phreo-
Opistocysta corderoi). drilus crozetensis Michaelsen, 1905; Hesperodrilus croze-
Trieminentia corderoi (Harman, 1970). Syn: tensis (Michaelsen, 1905); Phreodriloides crozetensis
Opistocysta corderoi Harman, 1970; Opistocysta (Michaelsen, 1905); Phreodrilus (Phreodriloides) croze-
flagellum, Cordero, 1948 (non Leidy, 1880). Distr: tensis Michaelsen, 1905. Distr: ARG: 4. Furth. Distr:
ARG: 8, 35, 60, 85, 86, 89, 90, 106, 115-118. URU: 1, South Georgia and Crozet Islands (Gavrilov, 1979:
10. Furth. Distr: North America (Cordero, 1948: 4; 107).
Milligan, 1997: 21; Kathman & Brinkhurst, 1998: 28).
Phreodrilus Beddard, 1891 (Type: Phreodrilus
PHREODRILIDAE Beddard, 1891 (Type: subterraneus Beddard, 1891). Syn: Hesperodrilus
Phreodrilus). Rem: Brinkhurst (1965) places most Beddard, 1894 (Type: Hesperodrilus branchiatus
genera within Phreodrilus as subgenera, but in a later Beddard, 1894).
cladistic analysis resurrected these genera and Phreodrilus (Phreodrilus) branchiatus (Beddard,
described two new ones (Brinkhurst, 1991). 1891). Syn: Hesperodrilus branchiatus Beddard, 1894;
Antarctodrilus Brinkhurst, 1991 (Type: Hesperodrilus Phreodrilus branchiatus (Beddard, 1894). Distr: CHI:
niger). Syn: Phreodrilus (Gondwanadrilus) Goddard 3. Furth. Distr: Australia (Naidu & Naidu, 1980: 179;
& Malan, 1913 (Type: Phreodrilus africanus Goddard Pinder, 2007).
& Malan, 1913).
Antarctodrilus niger (Beddard, 1894). Syn: Hes- TIGUASSUIDAE Brinkhurst, 1988 (Type: Tigu-
perodrilus niger Beddard, 1894; Phreodrilus niger assu).
(Beddard, 1894); Phreodrilus (Antarctodrilus) niger Tiguassu Righi, Ayres & Bittencourt, 1978 (Type:
(Beddard, 1894); Hesperodrilus albus Beddard, 1894; Tiguassu reginae).
Phreodrilus albus (Beddard, 1894); Hesperodrilus *Tiguassu reginae Righi, Ayres & Bittencourt,
pellucidus Beddard, 1894; Phreodrilus pellucidus 1978. Distr: BRA: AM: 16.
(Beddard, 1894); Phreodrilus albus var. pellucidus
(Beddard, 1894); Phreodrilus africanus Goddard & Discussion
Malan, 1913; Hesperodrilus africanus (Goddard &
Malan, 1913). Distr: ARG: 3, 5. Furth. Distr: South No fewer than 171 nominal species and subspecies of
Africa (Goddard & Malan, 1913: 242; Brinkhurst, aquatic microdriles – exclusive of Enchytraeidae – have
1971b: 631) and Tasmania (Pinder & Brinkhurst, 1997: been referenced in the literature for South America; to
459). Rem: Gondwanaedrilus africanus Goddard & date, 66 (circa 38.5%) of these taxa are known
Malan, 1913 is and incertae sedis taxon, but distinct exclusively from South America. While 23% of these
from Phreodrilus africanus Goddard & Malan, 1913 taxa are shared with North America north of Mexico,
(Pinder, pers. observ.). Both species were described in only 16% are shared with Australia.
different papers of the same volume (Brinkhurst, 1965: I find the record of Limnodrilus variegatus
363). Müller, cited for the Andean Plateau of Argentina
(Miserandino, 2001: 153; 2004: 196) to be somewhat
Insulodrilus Brinkhurst, 1991 (Type: Phreodrilus confusing, and likely a misprint. Perhaps another
lacustris). species of Limnodrilus was intended, because Lum-
Insulodrilus lacustris (Benham, 1903). Syn: Phreodrilus bricus variegatus Müller, 1774 is the type species of
lacustris Benham, 1903; Phreodrilus (Insulodrilus) Lumbriculus Grube, 1844, a taxon that has not
lacustris Benham, 1903; Hesperodrilus lacustris previously been found in the South American
(Benham, 1903). Distr: ARG: 18. Furth. Distr: New continent.
Zealand (Benham, 1903: 204) and Australia (Pinder,
2007). Acknowledgements
Nesodrilus Pinder & Brinkhurst, 1997 (Type: I am particularly indebted to Hong-Zhu Wang, who
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invited me to prepare and present a keynote address Alves, R. G. & G. Strixino, 2000. Distribuiçăo espacial de Oligochaeta

focusing on South American clitellates at the 10th em uma lagoa marginal do Rio Moji-Guaçu, Săo Paulo, Brasil.
Iheringia Série Zoologia 88: 173-180.
International Symposium on Aquatic Oligochaeta
Alves, R. G. & G. Strixino, 2003. The sampling of benthic macro-
(ISAO) in Wuhan, China (October 2006); this paper is
invertebrates using two different methods: Waiting trays and an
based, in part, on that presentation. Prof. Yan-Ling
Ekman collector. Acta Limnológica Brasileira 15(3): 1-6.
Liang, Convener Hong-Zhu Wang, and his numerous Armendariz, L. C., 1999a. Dinámica poblacional de Allonais lairdi
students, in particular Wei-hua Zhao and Hai-jun (Oligochaeta, Naididae) en los Talas, Provincia de Buenos Aires.
Wang, made my stay in China memorable and Ecologia Austral 9: 20-27.
pleasant. The latter presented me with a significant Armendariz, L. C., 1999b. Dero (Aulophorus) costatus (Marcus, 1944)
symbol in the Chinese horoscope, a snake neck emm. Harman, 1974 (Oligochaeta: Naididae) en Los Talas,

pendant, which I used to avoid many diversifying provincia de Buenos Aires. Neotropica La Plata 45: 101-102.
Armendariz, L. C., 2000. Population dynamics of Stylaria lacustris
pressures during the last month of intensive effort that
(Linnaeus, 1767) (Oligochaeta, Naididae) in Los Talas, Argentina.
this manuscript demanded. Besides the deadline, as
Hydrobiologia 438: 217-226.
discussed in conversation with Jörg Römbke, there is
Armendariz, L. C. & I. I. César, 2001. The distribution and ecology of
an intrinsic limit to how much checking of data can be littoral Oligochaeta and Aphanoneura (Annelida) of the natural
accomplished before one becomes a complete social and historical reserve of Isla Martín García, Río De La Plata River,
outcast. I am indebted to Claudia Bérrio-Cárdenas for Argentina. Hydrobiologia 463: 207-216.
providing me with a key reference on Colombian Armendariz, L. C., I. I. César & M. C. Damborenea, 2000. Oligoquetos
oligochaetes. I thank my wife, daughter, students, and en ambientes lénticos de la Reserva Natural e Histórica de la Isla

colleagues for listening to my inordinate demands for Martín García, Río de La Plata Superior, Argentina. Natura
Neotropicalis 31: 73-79.
patience, even though everyone, including myself,
Beddard, F. E., 1894. Preliminary notice of South American Tubificidae
often became very impatient. Despite the inaccuracies
collected by Dr. Michaelsen, including the description of a
that remain, I hope users will find this contribution
branchiate form. Annals and Magazine of Natural History Series 6
useful. I extend my appreciation to Mark J. Wetzel and 13: 205-210.
all participants of the 10th ISAO for welcoming me Beddard, F. E., 1895. A monograph of the order Oligochaeta.
into the ´family´ of aquatic oligochaetologists. Finally, Clarendon Press, Oxford.
I thank Adrian Pinder for providing important updates Beddard, F. E., 1896. Naiden, Tubificiden und Terricolen. I. Limicole
for the phreodrilid data presented herein, and Wetzel Oligochaeten. Ergebnisse der Hamburgische Magelanischen

and an anonymous referee for providing critical and Sammelreise 1892/93 36(6): 1-44.
Berrío-Cárdenas, C. & I. Vélez., 2008. Aquatic Oligochaeta (Annelida:
helpful remarks on the manuscript. Their suggestions,
Clitellata) of the Department of Antioquia, Colombia, South
and particularly the constructive input from Mark
America. Acta Hydrobiologica Sinica (this volume).
during four revisions of this manuscript, have
Bertoldi de Pomar, H., C. Copes, I. Ezcurra de Drago & M. Marchese,
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tributarios en el tramo Goya-Diamante. Los sedimentos de fondo y su
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APPENDIX river (Gluzman de Pascar, 1998a: 141); 41, Salgado lake


(Gluzman de Pascar, 1998a: 141); 42, Baldovino stream (Gluzman
ARGENTINA [ARG]: de Pascar, 1987: 126-128); 43, Zapata stream (Gluzman de Pascar,
1 (Du Bois-Reymond Marcus, 1947: 5; Gavrilov, 1955a: 294; 1955b: 1987: 126); 44, El Carnaval (Gluzman de Pascar, 1987: 126); 45,
302; Gavrilov, 1970: 147; Di Persia, 1980a: 79; Varela et al., 1980: El Gato (Gluzman de Pascar, 1987: 126); 46, El Azul stream
201; Varela, 1990: 224-231; Gluzman de Pascar, 1992: 11; (Gluzman de Pascar, 1987: 126-128); 47, Puerto Nuevo (Cordero,
Marchese & Brinkhurst, 1996: 163; Capello et al., 2004: 136); 2, 1951: 239); 48, El Pescado (Gluzman de Pascar, 1987: 126); 49,
Falkland Islands (Erséus & Grimm, 2002: 77); 3, Port Stanley El Pescado stream (Gluzman de Pascar, 1987: 126-128); 50,
(Michaelsen, 1900: 39; Cognetti, 1905: 7); 4, Tierra del Fuego Rodríguez (Gluzman de Pascar, 1987: 126); 51, Rodriguez
(Michaelsen, 1903: 175; 1921: 1; Brinkhurst & Marchese, 1989:1); stream (Gluzman de Pascar, 1987: 126); 52, Atalaya area
5, Ushuaia (Beddard, 1894: 208); 6, Patagonia (Michaelsen & (Gluzman de Pascar, 1998b: 97); 53, Atalaya channel (Gluzman de
Boldt, 1932: 591); 7, Andean Patagonian Plateau (Miserandino, Pascar, 1987: 126); 54, Punta Lara (Gluzman de Pascar, 1992: 12;
2001: 153; 2004: 196); 1994b: 114); 55, La Balandra channel (Gluzman de Pascar, 1987:
Chaco Prov: 8, Rio Negro (Di Persia, 1979: 45; 1980a: 79; Varela et al., 126); 56, Upper La Plata river: 57, Martín (Gluzman de Pascar,
1980: 201; Marchese, 1986: 236); 9, Madrigon El Puente (Varela, 1987: 126); 58, Martín García island (Armendariz et al., 2000: 75;
1984: 22); 10, Chaco (Di Persia, 1980a: 79); 11, La Cava (Di Armendariz & César, 2001: 212); 59, Palo Blanco (Gluzman de
Persia & Varela, 1980: 1); 12, Barranqueras (Di Persia & Varela, Pascar, 1996: 2);
1980: 1); 13, Tragadero river (Di Persia & Varela, 1980: 1); Santa Fé Prov: 60 (Di Persia, 1976: 15; 1979: 45; 1980b: 79; Di Persia,
San Juan Prov: 14 (Gluzman de Pascar & Moreno de Terzano, 1992: 1980c: 185): 61, Don Felipe river (Di Persia, 1980a: 98); 62, Don
54); 15, San Juan river (Varella et al., 1980: 201; Brinkhurst & Felipe lagune (Di Persia, 1973: 281; 1974a: 37; 1974b: 89), 63,
Marchese, 1987: 3154; Harman et al., 1988: 2233-2238); 16, La Saladillo river basin (Brinkhurst & Marchese, 1987: 3159;
Florida dam (Brinkhurst & Marchese, 1987: 3154); 17, Guanache Harman et al., 1988: 2235-2238; Marchese, 1990: 55; Marchese &
pond (Harman et al., 1988: 2239); Ezcurra de Drago, 1999: 241); 64, Santa Fé (Di Persia, 1980a:
Rio Negro Prov: 18, Mascardi lake (Brinkhurst & Marchese, 1987: 79-98); 65, Mendieta brook (Marchese, 1986: 236; Brinkhurst &
3154-3159; Harman et al., 1988: 2235); Marchese, 1987: 3158-3159); 66, epizoic on pulmonate gastropod
Mendoza Prov: 19, Parque General San Martín lagune (Di Persia, 1981: Ampullaria (Di Persia & Radici De Cura, 1973: 309); 67, San
1); 20, Mendoza (Di Persia, 1980a 79); Javier river (Brinkhurst & Marchese, 1987: 3159; Harman et al.,
21, Maipu (Di Persia, 1980b: 79; 1980d: 117); 1988: 2237); 68, Colastinecito river (Brinkhurst & Marchese,
Misiones Prov: 22 (Černosvitov, 1936b: 75; 1939: 96): 23, Uragua-i 1987: 3159); 69, Colastine river (Brinkhurst & Marchese, 1987:
(Harman et al., 1988: 2237); 24, False Uruagua-i rivers (Harman 3159); 70, Coronda river (Brinkhurst & Marchese, 1987: 3159);
et al., 1988: 2237); 25, Misiones (Černosvitov, 1936a: 19; 1937: 71, Mal Abrigo tributary (Brinkhurst & Marchese, 1987: 3159); 72,
145; 1939: 103-106; 1942: 198); Loreto Dep: 26, Loreto Yacarecito brook (Di Persia et al., 1982: 13); 73, Redonda lagoon
(Černosvitov, 1937: 139); 27, near Loreto (Černosvitov, 1937: (Di Persia, 1974a: 37; 1974b: 89), 74, La Guardia lagoon (Di
136); 28, Pastora brook (Černosvitov, 1937: 140-145; Brinkhurst, Persia, 1973: 281; 1974a: 37; 1974b: 89), 75, El Alemán lagoon
1964: 533); 29, La Plata river (Gomez & Rodrigues, 2007); (Di Persia, 1974a: 37; 1974b: 89);
30, Guayavera brook (Černosvitov, 1937: 145); 31, Argentine Basin, Cordoba Prov: 76, San Roque river (Bonetto et al., 1976: 47; Di Persia,
marine (Erséus, 1983b: 144); 1976: 2); 77, Los Molinos dam (Bonetto et al., 1976: 47; Di Persia,
Santiago del Estero Prov: 32, Termas de Río Hondo pond (Brinkhurst 1976: 2); 78, Tercero river (Di Persia, 1976: 2); 79, La Viña dam
& Marchese, 1987: 3157; Harman et al., 1988: 2237; Gluzman de (Varella et al., 1980: 201; Brinkhurst & Marchese, 1987: 3154;
Pascar & Moreno de Terzano, 1992: 54); Harman et al., 1988: 39); 80, Embalse Río Tercero lake (Bonetto
Buenos Aires Prov: 33 (Pujals, 1986: 149; Gluzman de Pascar, 1993: et al., 1976: 47); 81, San Roque lake (Di Persia, 1973: 284);
47-49): 34, Buenos Aires (Beddard, 1894: 206; Gluzman de Pascar, 82, Concórdia (Varela et al., 1980: 201); 83, Lower Paraná river
1989b: 12): 35, Berisso (Pujals, 1985: 204-209; Pujals, 1988a: 123; (Blattler & Marchese, 2005: 62);
Pujals, 1988b: 203-211; Gluzman de Pascar, 1989a: 8; 1990b: 122; Entre Ríos Prov: 84 (Di Persia, 1975a: 1; Di Persia, 1975c: 13; Di
1994a: 6); 36, Los Talas (Armendariz, 1999a: 24; 1999b: 101; Persia, 1980b: 77; 1980d: 117; Pavé & Marchese, 2005: 188): 85,
2000: 217); 37, Buenos Aires, associated with Eichhornia Paraná river (Rosso de Ferradas & Di Persia, 1975: 61; Marchese
crassipes (Gluzman de Pascar, 1994a: 6; Capello et al., 2004: 135); & Ezcurra de Drago, 1983: 100; Bertoldi de Pomar et al., 1986:
38, La Plata river (Gluzman de Pascar, 1987: 126; Gluzman de 79-92; Marchese, 1987: 209-212; Capelo et al., 2004: 135;
Pascar, 1993: 48; Gomez & Rodrigues, 2007); 39, Samboronbón Marchese & Paggi, 2004: 219; Montalto & Marchese, 2005: 490);
river (Gluzman de Pascar, 1989b: 12; 1998a: 141 ); 40, Salgado 86, Middle Paraná river (Di Persia, 1980b: 77; Marchese &
Suppl. Christoffersen: A catalogue of aquatic microdrile oligochaetes (Annelida: Clitellata) from South America 61
83

Ezcurra de Drago, 1983: 100; Marchese, 1984: 161; 1986: Iquitos (Harman et al., 1988: 2237).
231-246; 1987: 212; 1994: 303; Bertoldi de Pomar et al., 1986: 92;
Brinkhurst & Marchese, 1987: 3157; Capello et al., 2004: 135-136; CHILE [CHI]:
Montalto & Marchese, 2005: 490; Ezcurra de Drago et al., 2005: 1 (Michaelsen, 1905: 357; Gluzman de Pascar, 1990a: 89); Patagonia: 2,
67; 2007: 255; Galizzi & Marchese, 2007: 535); 87, Middle Esquel-Percy river system (Miserandino & Pizzolón, 2000: 64); 3,
Paraná river, epizoic on pulmonate gastropod Ampullaria (Di Valdivia (Beddard, 1894: 207; 1896: 5; Michaelsen, 1900: 38;
Persia & Radici de Cura, 1973: 309); 88, Entre Ríos (Černosvitov, Piguet, 1928: 81); 4, San Carlos (Cognetti, 1905: 7); 5, Valparaiso:
1937: 145); 89, Saucesito tributary (Pavé & Marchese, 2005: 188); Salto (Beddard, 1895: 293; 1896: 4); 6, near Valparaiso (Cognetti,
90, Las Tunas tributary (Pavé & Marchese, 2005: 188); 91, Las 1905: 6).
Tunas brook, between Paraná and Viale (Gavrilov, 1958: 226); 92,
Ayui Grande brook (Di Persia, 1977b: 13; 1980a: 98); 93, Paraná COLOMBIA [COL]:
Dep. (Di Persia, 1979: 120): 94, Paraná city (Marchese, 1984: 1(Michaelsen, 1914: 202-208; Gaviria, 1993: 238-241);
162); 95, close to Paraná city (Di Persia, 1980d: 119); Antioquia Dep: 2 (Gaviria, 1993: 239): 3, Medellín Mun. (Be-
Corrientes Prov: 96, (Di Persia, 1975a: 5; Di Persia, 1975d: 25; 1976: rrío-Cárdenas & Vélez, 2008); 4, Mutatá Mun. (Berrío-Cárdenas
15; 1979: 44): 97, Paraná river (Righi & Varela, 1983: 7; Bertoldi & Vélez, 2008); 5, Jardín Mun. (Berrío-Cárdenas & Vélez, 2008);
de Pomar et al., 1986: 92; Marchese, 1987: 211; Montanholi- 6, Chigorodó Mun. (Berrío-Cárdenas & Vélez, 2008); 7, Caldas
Martins & Takeda, 1999: 52); 98, Upper Paraná river (Varela et al., Mun. (Berrío-Cárdenas & Vélez, 2008); 8, Santo Domingo Mun.
1983: 103; Capello et al., 2004: 135; Marchese et al., 2005: 485); (Berrío-Cárdenas & Vélez, 2008); 9, Santa Rosa de Osos Mun.
99, Fechada (Takeda, 1999: 35); 100, Pousada das Garças lake (Berrío-Cárdenas & Vélez, 2008); 10, Girardota Mun. (Be-
(Takeda, 1999: 35); 101, Guarana lake (Takeda, 1999: 35); 102, rrío-Cárdenas & Vélez, 2008); (Berrío- Cárdenas & Vélez, 2008);
Baia river (Takeda, 1999: 35); 103, Ivinhema tributary (Takeda, 11, Don Matías Mun. (Berrío-Cárdenas & Vélez, 2008).
1999: 35; Montanholi-Martins & Takeda, 2001: 202); 104, Patos ¨Please see the paper by Berrío-Cárdenas & Vélez (2008), included
lake (Montanholi-Martins & Takeda, 2001: 202); 105, Corrientes elsewhere in this issue of Acta Hydrobiologica Sinica, for an
(Brinkhurst & Marchese, 1987: 3160); 106, La Brava lagune (Di historical review of aquatic oligochaete research in Antioquia Dep.,
Persia, 1973: 283; Bonetto et al., 1978a: 17; 1978b: 57; Di Persia, and elsewhere in Colombia¨.
1980a: 79; Di Persia & Varela, 1980: 1; Varela et al., 1978: 97;
Varela et al., 1980: 201; 1983: 103); 107, Perez lagoon (Varela, ECUADOR [ECU]:
1984: 22); 108, San Cosme Dep. (Di Persia, 1976: 5); 109, Itatí 1 (Coates & Stacey, 1994: 82); 2, Cuenca (Cognetti, 1905: 5); 3,
Dep: Itatí (Di Persia, 1976: 6; 1980a: 98); 110, Totoras lagune Galapagos Islands (Erséus, 1980: 104; 1984: 193-196).
(Bonetto et al., 1978a: 17; 1978b: 57; Di Persia, 1979: 45); 111,
Goya (Di Persia, 1980a: 98); 112, Ituzaingó: Rincón de Santa GUYANA [GUY]:
Marta (Poi de Neiff & Neiff, 1980: 185; Poi de Neiff, 1981: 41); 1 (Coates & Stacey, 1994: 82; Stacey & Coates, 1996: 23-42); 2,
113, near Ituzaingó (Di Persia, 1980a: 107); 114, Mitiñay cannel Kwabanna swamp (Omodeo & Coates, 2001: 39); 3, Essequibo
(Di Persia & Varela, 1980: 1); 115, Galarza (Di Persia, 1979: 45); river: Kurupukari creek (Omodeo, 1996: 12); 4, Amakut (Omodeo
116, Ibera (Di Persia, 1979: 45); 117, Sirena (Di Persia, 1979: 45); & Coates, 2001: 45).
118, González (Di Persia, 1979: 45); 119, González lagune
(Bonetto et al., 1978: 57); PARAGUAY [PAR]:
Tucumán
84 Prov: 120 (Gavrilov, 1948: 228; Gavrilov & Paz de Tomsic, 1 (Michaelsen, 1905: 350-359; Piguet, 1928: 81; Černosvitov, 1939:
1950: 543; Cordero, 1951: 237); 121, del Toro brook (Gavrilov & 97); 2, Makthlawaiya swamp (Michaelsen, 1905: 354; Stephenson,
Paz de Tomsic, 1949: 537); 122, Tucumán (Cordero, 1951: 237); 1931: 299-306); 3, Rio Verde stream (Stephenson, 1931: 309); 4,
123, Sali river (Di Persia, 1973: 284); Chaco, swamps (Carter & Beadle, 1931: 379).
Salta Prov: 124 (Gavrilov, 1970: 147); 125, Tala river (Gavrilov & Paz
de Tomsic, 1950: 543); PERU [PER]:
Jujuy Prov: 126 (Gavrilov, 1970: 147). 1 (Blanchard, 1849: 39; Piguet, 1928: 82); Lake Titicaca: 2
(Černosvitov, 1939: 83-88; Lafont & Juget, 1992: 302); 3, Bahia
BOLIVIA [BOL]: de Plata (Brinkhurst & Marchese, 1987: 3157; Harman et al., 1988:
1 (Martínez-Ansemil & Giani, 1986: 108); 2, Lake Titicaca basin 2237); 4, Uruñi Bay (Černosvitov, 1939: 97); 5, Sucuné
(Martínez-Ansemil & Giani, 1986: 110, 113); 3, Amazon basin (Černosvitov, 1939: 97); 6, Ojerani (Brinkhurst & Marchese, 1987:
(Martínez-Ansemil & Giani, 1986: 113); 4, Amazon river (Ma- 3157; Harman et al., 1988: 2237); 7, Lagunilla Lagunilla
rtínez- Ansemil & Giani, 1986: 113); 5, Amazon river basin, above (Černosvitov, 1939: 94-106); 8, off Faro: Punta Churo (Brinkhurst
84
60 ACTA HYDROBIOLOGICA SINICA Vol. 31

& Marchese, 1987: 3158); 9, Lake Titicaca basin (Černosvitov, 270-359 m deep (Blazejak et al., 2005: 1153).
1939: 101; Harman et al., 1988: 2235): 10, Poopó lake
(Černosvitov, 1939: 84); 11, Lake Umayo (Brinkhurst & Marchese, REPUBLIC OF SURINAME [SUR]:
1987: 3155; Harman et al., 1988: 2235); 12, Saracocha lake 1 Surinam river (Righi & Hamoui, 2002: 134); 2, Lelydorp (Harman,
(Černosvitov, 1939: 89-103): 13, Yanayako river (Coates & Stacey, 1974: 7-31); 3, Onverwacht (Harman, 1974: 7-31), 4, Uitkijk
1994: 82); 14, San Antonio de Esquilache (Černosvitov, 1939: 89); (Harman, 1974: 5-31), 5, Zanderij (Harman, 1974: 9-31), 6,
15, Huenque (Harman et al., 1988: 2235-2238); 16, Huenque river Republiek (Harman, 1974: 20); 7, near Republiek: Coroupina:
(Brinkhurst & Marchese, 1987: 3157); 17, Capachica pond swamp Berseba (Righi & Hamoui, 2002: 127-129); 8, reiver near
(Černosvitov, 1939: 86-95; Marchese, 1986: 236); 18, Murilaya Republiek (Righi & Hamoui, 2002: 127, 128, 148); 9, Paramaribo
river, near Puno Bay (Brinkhurst & Marchese, 1987: 3157; (Harman, 1974: 5-31); 10, Paramaribo: Trench in Cultuurtuim
Harman et al., 1988: 2235-2238); 19, Murilaya stream, near (Righi & Hamoui, 2002: 124, 127); 11, near Paramaribo:
Huenque; on road to Machu Picchu, 81 km from Cuzco: Belwaarde pond (Righi & Hamoui, 2002: 129, 130, 138); 12,
Huairachina: Urumbamba river valley (Brinkhurst & Marchese, about 42 km south of Paramaribo (Righi & Hamoui, 2002: 145);
1987: 3154); 20, Puno Bay (Brinkhurst & Marchese, 1987: 13, north of Paramaribo: swamp at Krepi (Righi & Hamoui, 2002:
3154-3157; Harman et al., 1988: 2235-2237); 21, Puno Bay, near 150); 14, Kabel (Righi & Hamoui, 2002: 144. 150); 15, Ditch at
Uros village (Harman et al., 1988: 2235-2239); 22, Puno Bay at Kabel (Righi & Hamoui, 2002: 124, 127, 131); 16, Zoetwaterput
Ojerani (Harman et al., 1988: 2236-2238); 23, Maranón drainage (Rigfhi & Hamoui, 2002: 126, 127, 129, 131, 134, 143, 148); 17,
(Coates & Stacey, 1994: 82); 24, Uros islands, 20 m deep Continental slope, 1000-1518 m deep (Erséus, 1982: 223).
(Brinkhurst & Marchese, 1987: 3157); 25, Amazon basin: Langui
lake: Layo (Černosvitov, 1939: 109); URUGUAY [URU]:
26, access to Machu Pichu (Brinkhurst & Marchese, 1950: 533; 1987: 1 (Michaelsen, 1903: 175; Cordero, 1931: 345; Cordero, 1951: 231-237;
3157; Harman et al., 1988: 2239); Harman & Loden, 1978: 453);
Madre de Diós Dep: 27, Pampas de Heath, south of Puerto Pardo Montevideo: 2, El Prado (Cordero, 1931: 351; Cordero, 1951: 239); 3,
(Harman et al., 1988: 2237-2239); Prado Botanical Garden (Cordero, 1951: 235); 4, Barra de Santa
Cusco Dep: 28, Huacarpay lagune (Harman et al., 1988: 2237); Lucia (Cordero, 1931: 349-350); 5, Miguelete stream (Cordero,
Ayacucho Dep: 29, Yaracyacu, from bromeliad (Harman et al., 1988: 1931: 350); 6, in brackish water (Du Bois-Reymond Marcus,
2239); 1949b: 1); 7, Maldonado: Piriápolis (Cordero, 1931: 349); 8,
Lima Dep: 30, Cienequella on Lurin river (Harman et al., 1988: 2233); Carrasco stream (Cordero, 1931: 350);
31, Lurin river, near Cieneguilla (Harman et al., 1988: 9, Continental shelf, 256-293 m deep (Erséus, 1982: 218); 10, La
2235-2239); Estacion Parque del Plata: near Solis river (Cordero, 1948: 4).
32, Huaron lake (Piguet, 1928: 80, 81, 82, 91, 92); 33, Murilaya stream
(Brinkhurst & Marchese, 1987: 3154); 34, Huenque (Brinkhurst VENEZUELA [VEN]:
& Marchese, 1987: 3154); 35, Huenque river (Brinkhurst & 1 (Dumnicka, 1983: 255): 2, Amazon basin: Bolivar State: Santa
Marchese, 1987: 3154); 36, Yanayako river, a tributary of the Helena de Uairem: margin of Salto river (Righi, 1985: 24);
Amazon above Iquitos (Coates & Stacey, 1994: 82); 37, Naticocha State of Merida: 3, Los Guaches lagoon (Stacey & Coates, 1996: 24);
lake (Piguet, 1928: 93; Brinkhurst, 1971c: 521); 38, Panguana Aragua Dep: 4, Cuyagua brook, interstitial (Botea, 1987: 69, 70); 5,
(Coates & Stacey, 1994: 82); 39, North Peru (Coates & Stacey, Quebrada de la Toma, hyporeic (Botea, 1987: 70-71);
1994: 82); 6, Maracay, in Hyla spp (Michaelsen, 1926a: 100);
Loreto Dep: 40, Curanja Balta river (Harman et al., 1988: 2237-2239); Falcón Dep: 7, (Dumnicka, 1986: 267); 8, Arroyo El Pitiri (Botea, 1987:
41, Laguna Huacarpay (Harman et al., 1988: 2237); 42, 70), 9, Curimagua (Botea, 1987: 70), 10, Sierra de San Luis
Yarinacoche lake (Harman et al., 1988: 2237); Missiones Prov: 43, (Botea, 1987: 70); 11, Falcón Distr: Paraguana peninsula: Fuente
Tirica stream (Harman et al., 1988: 2235); de Miraca (Dumnicka, 1983: 258); 12, Miraca village (Botea,
44, Amazon (Shain et al., 2008; Martinez-Ansemil & Giani, 1986: 113); 1987: 70);
45, near Iquitos: Yanayako river (Harman et al., 1988: 2236; 13, Marguerita island (Righi & Hamoui, 2002: 138, 142, 144): 14,
Coates & Stacey, 1994: 82); mangroves (Erséus, 1982: 260);
Piura Dep: 46, Piura river (Harman et al., 1988: 2239); Off coast of Punta Piedras: 15, South of Bahia de Amuay, about 15 m
47, Bahia La Plata: beach at Ojerani, 50 m deep; 48, Off coast, shell deep (Erséus, 1987: 272); 16, South of Bahia de Amuay, about 3
sand, 50 m deep (Finogenova, 1986: 45-50); 49, Off coast, 150m; m deep (Erséus, 1987: 273); 17, northeast part of Bahia de Amuay,
50, Off coast, 169 m deep (Finogenova, 1986: 46-48); 51, Off about 4 m deep (Erséus, 1987: 273);
coast, 47-200 m deep (Finogenova, 1986: 46); 52, Off coast, 18, Chama river (Giani, 1978: 270);
Suppl. Christoffersen: A catalogue of aquatic microdrile oligochaetes (Annelida: Clitellata) from South America 61
85

Monagas Dep: 19, Caripe river, hyporeic (Botea, 1987: 71); 20, beach Belo Horizonte: 2 (Michaelsen, 1927: 369);
near Santa Cruz (Botea, 1987: 68); 3, Ibirité reservoir (Callisto et al., 2005: 1);
Sucre Dep: 21, Jaguaracual stream, near Cumanacoa, hyporeic (Botea, 4, Serra do Cipó: Capivara river (Righi, 1973: 294).
1987: 68-69);
Central Dep: Caracas: Jardin Botánico: 22, Laguna del Leste (Righi & MATO GROSSO DO SUL [MS]:
Hamoui, 2002: 126, 127, 131, 151); 23, Laguna del Oeste (Righi Pantanal: 1, Rio Negro (Takeda et al., 2000: 53).
& Hamoui, 2002: 143).
MATO GROSSO [ MT]:
BRAZIL [BRA]: 1 (Michaelsen, 1900: 522);
2, Carandazinho, near Corumbá (Cognetti, 1900: 1).
ALAGOAS [AL]:
Maceió: 1(Marcus, 1944: 51); PARÁ [PA]:
Satuba: 2 (Marcus, 1943: 35; 1944: 65); 3, Catolé river: 3 (Marcus, 1, Tapajós river (Du Bois-Reymond Marcus, 1947: 6); 2, Fordlândia
1944: 51); 4, Catolé river, near Satuba (Marcus, 1944: 50-65; Du (Du Bois-Reymond Marcus, 1947: 4- 9; 1949a 2; 1949b: 2-3); 3,
Bois-Reymond Marcus, 1947: 3; 1949: 1); near Fordlândia (Marcus, 1942: 157; Du Bois-Reymond Marcus,
5, Paulo Afonso falls (Marcus, 1943: 21); 1947: 1949a: 2); 4, Belterra (Du Bois-Reymond Marcus, 1947: 9;
Garça Torta: 6 (Du Bois-Reymond Marcus, 1947: 9). 1949b: 2); 5, near Belterra (Marcus, 1942: 157; Du Bois-Reymond
Marcus, 1947: 4-11; 1949a: 1-2; 1949b: 3-10); 6, Cururú river, a
AMAZONAS [AM]: tributary of Tapajós river (Du Bois- Reymond Marcus, 1947:
1 (Righi & Hamoui, 1989: 409; Collado & Schmelz 2001: 4; 2002: 2); 1-11);
Laguna del este (Righi & Hamoui, 2002: 2, Transamazonian highway Santarém: 7 (Marcus, 1942: 157; Du Bois-Reymond Marcus, 1947:
(Righi et al., 1978: 11); 4-11; 1949a: 2-9; 1949b: 2);
3, Madeira river (Marcus, 1944: 51); Óbidos: 8 (Marcus, 1942: 157; Du Bois-Reymond Marcus, 1949a: 2);
4, near Humaitá (Marcus, 1943: 112; Du Bois- Reymond Marcus, 1947: 9, Trombetas river (Marcus, 1942: 157; Du Bois- Reymond Marcus,
1-11; 1949a: 2; 1949b: 2); 1947: 4-11); 10, Acará river (Du Bois-Reymond Marcus, 1947:
5, Solimões river (Marcus, 1944: 51; Righi, 1978: 485); 9-11; 1949b: 2); 11, south of Belém (Du Bois-Reymond Marcus,
Maués: 6 (Du Bois-Reymond Marcus, 1947: 7-11; 1949b: 2); 1947: 1-10); 12, São Manuel river (Du Bois-Reymond Marcus,
7, Lago Grande Curuay (Marcus, 1944: 51; Du Bois- Reymond Marcus, 1947: 4-11); 13, Cuminá river (Marcus, 1942: 157; Du
1944: 65; 1947: 6); Bois-Reymond Marcus, 1947: 4-11); 14, Salgado lake (Marcus,
8, near Vila Franca (Du Bois-Reymond Marcus, 1947: 11); 9, at Vila 1942: 157; Du Bois-Reymond Marcus, 1947: 4-11); 15, Cupari
Franca (Du Bois-Reymond Marcus, 1947: 6); 10, Lago Grande river (Marcus, 1942: 157; Du Bois-Reymond Marcus, 1947: 4-11;
Curuay, near Óbidos (Du Bois-Reymond Marcus, 1947: 4); 11, 1949a: 2; 1949b: 3); 16, Alemquer river (Marcus, 1942: 157; Du
Amazon river, opposite Óbidos (Du Bois- Reymond Marcus, 1947: Bois-Reymond Marcus, 1947: 7-9); 17, Juruena river (Marcus,
6); 12, Central Amazon (Irmler, 1989: 31; Collado & Schmelz, 1944: 51; Du Bois- Reymond Marcus, 1947: 6-11);
2000a: 228; 2001: 2); 13, Central Amazônia, in soil (Collado & Itaituba: 18 (Du Bois-Reymond Marcus, 1947: 1-9).
Schmelz, 2000a: 225); 14, Curari island (Righi, 1978: 485);
Manaus: 15 (Collado & Schmelz, 2000b: 510, 513); 16, Manaus–Rio PERNAMBUCO [PE]:
Branco highway, km 45 (Righi et al., 1978: 6). São Bartolomeu: 1 (Marcus, 1944: 50): 2, São Francisco river (Du
Bois-Reymond Marcus, 1947: 9); 3, São Francisco river, near
BAHIA [BA]: Belém (Marcus, 943: 35; 1944: 50-135); 4, near São Bartolomeu
Salvador: 1, Farol da Barra beach (Erséus, 1980 100). (Marcus, 1942: 157; 1943: 38);
São Caetano Mun: 5, Ipojuca river, near Antonio Olinto (Marcus, 1944:
CEARÁ [CE]: 65);
Araripe: 1, Paraíso reservoir; Cabo Mun: 6, Gurjaú reservoir (Marcus, 1944: 51-65); 7, Cabo (Du
Fortaleza: 2 (Marcus, 1942: 157; 1943: 35); 3, in colony of sponge Stoella Bois-Reymond Marcus, 1947: 1); 8, Bom Jardim (Marcus, 1944:
agilis f. iheringi (Marcus, 1943: 112; 1944: 150; Du Bois - Reymond 150; Du Bois- Reymond Marcus, 1947: 1);
Marcus, 1947: 9). Recife: 9 (Marcus, 1944: 50): 10, Capoeira (Marcus, 1944: 50), 11, São
Bento (Marcus, 1944: 50); 12, Tegipió (Marcus, 1943: 130); 13,
MINAS GERAIS [MG]: Jaboatão (Marcus, 1944: 69); 14, Jaboatão river (Marcus, 1944:
1 (Righi, 1973: 299); 50; Du Bois-Reymond Marcus, 1947: 60);
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86 ACTA HYDROBIOLOGICA SINICA Vol. 31

Goiana: 15 (Marcus, 1944: 50); (Marcus, 1943: 38); 6, Araquá river (Marcus, 1943: 108-129; 1944:
Mercês: 16 (Du Bois-Reymond Marcus, 1947: 1); 59); 7, Araquá river, near São Pedro (Marcus, 1943: 13);
Salgadinho: 17 (Du Bois-Reymond Marcus, 1947: 1); Cananéia: 8, in mangrove (Marcus, 1965: 63-77);
Garanhuns: 18 (Marcus, 1944: 50). Itanhaém: 9 (Lopez et al., 2005: 15);
PIAUÍ [PI]: 10 Cubatão (Marcus, 1965: 77);
1, Caicós reservoir (Marcus, 1944: 65; Du Bois- Reymond Marcus, São Paulo: 11 (Marcus, 1942: 157-208; 1943: 40-108; 1944: 69); 12,
1947: 6). Pinheiros (Marcus, 1943: 12-38); 13, Pinheiros river (Marcus,
1942: 157, 202; 1943: 23-112; Du Bois-Reymond Marcus, 1943:
PARANÁ [PR]: 12; 1944: 4); 14, Jardim Europa (Marcus, 1943: 23); 15, Jardim
1, Paraná river (Montanholi-Martins & Takeda, 1999: 52): 2, Patos lake Europa brook (Marcus, 1942: 202; 1943: 12-129); 16, Tietê river
(Montanholi-Martins & Takeda, 2001: 202); 3, Ivinhema tributary (Marcus, 1942: 175-202; 1943: 12-129; Du Bois-Reymond
(Montanholi- Martins & Takeda, 2001: 202); 4, Upper Paraná Marcus, 1949a: 2); 17, Lapa (Marcus, 1942: 200-202); 18, Vila de
river (Takeda et al., 2001: 241); 5, Caiobá cliff (Marcus, 1943: Caxingui (Du Bois- Reymond Marcus, 1947: 13); 19, Caxingui
93); brook (Marcus, 1942: 157; 1943: 13; 1944: 54); 20, Ibirapuera
Curitiba: 6 (Marcus, 1943: 40; 1944: 78). park, epizoic on pulmonate gastropod Australorbis (Du Bois-
Reymond Marcus, 1947: 2); 21, Nove de Julho avenue (Marcus,
RIO DE JANEIRO [RJ]: 1943: 112); 22, Cidade Jardim (Marcus, 1944: 68); 23, Itaim
1, Barra de Maricá (Lopez et al., 2005: 22); (Marcus, 1944: 68); 24, Botanical garden (Lopez et al., 2005: 15);
Rio de Janeiro: 2, Guanabara Bay, 22 m deep (Harman & Loden, 1984: 25, Canindé (Marcus, 1942: 157); 26, Santana (Marcus, 1942:
51); 3, Rio de Janeiro, in Hyla spp (Michaelsen, 1926a: 100; 1926b: 157); 27, Cantareira (Marcus, 1944: 68);
231; 1926c: 3); São Sebastião: 28, coast (Du Bois-Reymond Marcus, 1950: 1);
Nova Iguaçu: 4, in Bufo crucifer (Rodrigues & Maldonado Jr., 1982: 13; 29, Pirituba, near São Paulo (Mendes et al., 1951: 290);
Rodrigues, 1986: 27); Campinas: 30, near Campinas (Marcus, 1943: 12): 31, Atibaia farm
Búzios: 5, João Faema beach (Erséus, 1983a: 1052). (Marcus, 1943: 12); 32, Atibaia river (Marcus, 1943: 109);
Rio Claro: 33, near Mogi das Cruzes (Marcus, 1943: 35);
RORAIMA [RR]: Pirassununga: 34, Emas reservoir, near Pirassununga (Marcus, 1943:
1, Boa Vista – Bonfim highway, km 120 (Righi, 1988: 316); 2, Boa 60-112);
Vista (Righi, 1984: 26-36). Mogi-Guaçu: 35, Mogi-Guaçu river: Diogo lake (Alves & Strixino,
2000: 177; 2003: 4);
RIO GRANDE DO SUL [RS]: Brotas Mun: 36, Jacaré Pepira river (Gorni & Alves, 2007: 519); 37,
Porto Alegre: 1 (Marcus, 1943: 38); 2, vicinity of Porto Alegre; 3, Brotas, in bryophytes (Gorni & Alves, 2007: 518);
Quadros lagoon (Marcus, 1944: 71-73). Araraquara Mun: 38, associated with gastropod Pomacea bridgesii (Reeve,
1856) (Gorni & Alves, 2006: 1060);
SANTA CATARINA [SC]: 39, Tietê river: Bariri (Pamplin et al., 2005: 67); 40, Ponte Nova
Lauro Muller Mun: 1, Novo Horizonte, in Hyla (Righi, 1972: 150). reservoirs (Pamplin et al., 2005: 67); 41, São Paulo, in Hyla spp
(Michaelsen, 1926a: 100; 1926b: 231); 42, Limoeiro reservoir
(Marcus, 1943: 38); 43, Mongolinho river (Alves et al., 2006: 434);
SÃO PAULO [SP]: 44, Água Branca stream (Alves et al., 2006: 434); 45, Gouveia
1 (Marcus, 1943: 129); stream (Alves et al., 2006: 434); 46, Pinheirinho stream (Alves &
Ribeira de Iguape: 2, Ribeira de Iguape river (Marcus, 1943: 13); 3, in De Lucca, 2000: 115; Alves et al., 2006: 434); 47, Santa Clara
Ephydatia sponge colony (Marcus, 1943: 21-108); 4, near São stream (Alves & De Lucca, 2000: 115).
Pedro (Marcus, 1943: 112); São Pedro: 5, São Pedro station

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