Suggestion of a Cytotaxonomical Classification

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INTRODUCTION

Fifteen Neotropical freshwater families compose

the Order Siluriformes: Diplomystidae, Cetopsidae,

Aspredinidae, Nematogenydae, Trichomycteridae,

Callichtydae, Scoloplacidae, Astroblepidae, Lori-

cariidae, Pimelodidae, Heptapteridae,

Aucheni-pteridae, Pseudopimelodidae, Ariidae, Doradidae

(R

EIS

et al. 2003).

R

EIS

et al. (2003) proposed that a large part of

the traditional Pimelodidae was included in a

sep-arate family, the Heptapteridae, and another

ad-ditional group constitutes actually the

Pseudop-imelodidae with few genera and species described

at now. According to B

OCKMANN

and G

UAZZELI

(2003) the family Heptapteridae, closely

corre-sponds to the subfamily Rhamdiinae of L

UND

-BERG

et al. (1991) and Heptapterinae of the P

INNA

(1998) and includes many catﬁshes long classiﬁed

together in the Pimelodidae but now placed in a

more restricted version of Pimelodidae (L

UND

-BERG

and L

ITTMANN

2003) and

Pseudopimelodi-dae (S

HIBATTA

2003).

A recent checklist of catﬁshes (Order

Sil-uriformes) presented by F

ERRARIS

(2007) proposed

that the Heptapteridae family is composed by 24

genera and 189 valid species, the Pimelodidae by

29 genera and 93 valid species and

Pseudopimelo-didae by 5 genera and 29 valid species.

The aim of the present paper is update the

cytogenetic data for species of the families

Pime-lodidae, Heptapteridae and Pseudopimelodidae

and suggest a cytotaxonomical classiﬁcation.

MATERIALS AND METHODS

The data here discussed are available from four

distinct hydrographic basins (Upper Paraná,

Paguay, São Francisco and Amazon) found in

ar-ticles, unpublished thesis and congress abstracts.

They take into consideration the diploid, triploid

and haploid numbers, the presence of B and

sex-ual chromosomes and other relevant data (Tables

1, 2, 3). The classiﬁcation of the hydrographic

ba-sin was performed as described by V

ARI

(1992).

Karyological analyses were performed from

ce-phalic kidney cells, with either of the following

protocols: short-term culture cells (F

ENOCCHIO

et

al. 1991) or the conventional air-drying method

(B

ERTOLLO

et al. 1978) and from lymphocyte

cul-ture following the technique of F

ENOCCHIO

and

B

ERTOLLO

(1988). The diploid number was

deter-mined for each specimen studied (Tables 1, 2, 3).

The chromosomes were classiﬁed as metacentrics

(m), submetacentrics (sm), subtelocentrics (st) and

acrocentrics (a) according to their morphology

An update Cytogenetic Review for Species of the Families

Pseu-dopimelodidae, Pimelodidae and Heptapteridae (Pisces,

Sil-uriformes). Suggestion of a Cytotaxonomical Classiﬁcation

Swarça

1

_*

_{Ana Cláudia, Alberto Sérgio Fenocchio}

2

_{and Ana Lúcia Dias}

1

1 _{Universidade Estadual de Londrina, CCB, Caixa Postal 6001, 86051-970 Londrina, PR, Brasil.}

2 _{Universidad Nacional de Misiones, Depto. de Genética, Félix de Azara 1552, 3300, Posadas, Misiones, Argentina.}

Abstract — Fifty three species (48 species valid) belonging to the Pimelodidae, Heptapteridae and

Pseudopimelo-didae families have been studied cytogenetically, and in the present paper the chromosome number, the presence of B chromosomes and other relevant data were examined. The diploid number varies from 2n=42 to 2n=58 and were detected three cases of triploids with 3n=87 in Rhamdia species. The karyotypes show a high fundamental number because there are constituted predominantly by meta/submetacentric chromosomes.

Key words: cytogenetic, Heptapteridae, Pimelodidae, Pseudopimelodidae.

* Corresponding author: Rua Vila Lobos, 793; Jardim Tucanos; CEP 86047-130; Londrina, Paraná, Brazil; e-mail: szwarcy@bol.com.br

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Table 1. Cytogenetic data on the Pimelodidae Family

Genera, Species Locality HB n 2n Karyotype FN Bs Size B HS References Bergiaria

B. westermanni R. São Francisco, MG SF 56 42m,sm + 14st 98 0-5 small Dias and Foresti (1993)

Calophysus

C. macropterus R. Negro; R. Solimões, _AM AM 50 22m+18sm+10a (90) Ramirez-Gil et al. (1998)

Iheringichthys

I. labrosus R. Mogi-Guaçu, SP UP 56 26m+14sm+12st+4a 96* 0-2 micro Dias and Foresti (1990)

I. labrosus R. Paraná, PR UP 56 42m,sm+42st,a 98 Garcia et al. (1990)

I. labrosus R. Tibagi, PR UP 56 32m+8sm+6st+10a (96) 0-3 micro Carvalho et al. (2004)

I. labrosus D. Jurumirim, SP UP 56 22m+18sm+10st+6a 96 0-2 acrocentric Vissotto et al. (1999b)

Parapimelodus

P. valenciennes R. Guaiba, RS PA 56 Costa and Reggi (1986)

Pimelodus

P. blochii R. Solimões, AM AM 58 Della-Rosa et al. (1980)

P. blochii R. Araguai, Barra do _{Garças, MT} PA 56 36m/sm+20st/a 92* Faria et al. (2000)

P. blochii R. Araguai, Barra do _{Garças, MT} PA 56/58 36m/sm+20st/a_14m+8sm+36a 92*_80* Silva et al. (2004)

P. clarias Argentina PA 56 Fenocchio et al. (1994)

P. clarias Uruguai PA 26 Gonzales (1994)

P. fur 56 30m+14sm+12a 100* Toledo and Ferrari (1976b)

P. fur R. São Francisco, MG SF 54 32m+8sm+6st+8a (94) Garcia and Moreira Filho₍₂₀₀₅₎

P. maculatus 56 30m+14sm+12a 100* Toledo and Ferrari (1976b)

P. maculatus R. São Francisco, MG; _{R. Mogi-Guacu, SP} SF;_UP 56 40m,sm+16st,a 96 Dias and Foresti (1993)

P. maculatus R. Guaíba, RS PA 56 Costa and Reggi (1986)

P. maculatus R. Tibagi, Pr UP 56 20m+20sm+10st+6a (96) Swarça et al. (2001b)

P. maculatus R. Sapucaí; D. Furnas, _MG UP 56 40m/sm+16st/a 96 Marques et al. (1998)

P. maculatus R. Paranapanema; _{D. Jurumirim, SP} UP 56 20m+20sm+10st+6a 96 Vissotto et al. (1999a)

P. maculatus Delta Paranaense, Arg. UP 56 24m+14sm+12st+6a 94* Heras and Mendoza (2002)

P. maculatus R. Paraguai, MS PA 56 22m+16sm+10st+8a (94) Souza et al. (2003)

P. maculatus R. São Francisco, MG SF 56 32m+12sm+12st (100) Garcia and Moreira Filho (2005)

P. maculatus R. Paraná, Porto Rico/PR UP 56 20m+20sm+10st+6a (96) Borin and Martins-Santos ₍₂₀₀₂₎

P. cf. maculatus R. Jarí Almerim, PA 58 30m/sm+28st/a 88* Souza et al. (2000)

P. maculatus R. Tejuco; R. Araguari, _MG UP 56 Moreira et al. (2004)

P. ornatus R. Paraná, PR UP 56 18m+22sm+6st+10a 96* _{Martins-Santos (1996)}Abucarma and

P argenteus R. Paraguai, MS PA 56 24m+16sm+12st+4a (96) Souza et al. (2003a)

P. mysteriosus R. Paraguai, MS PA 56 26m+20sm+2st+8a (102) Souza et al. (2003a)

P. heraldoi R. Tibagi, PR UP 56 22m+22sm+6st+6a (100) Souza et al. (2004)

P. ortmanni Caxias Reservoir, R, _{Iguaçu, PR} UP 56 24m+18sm+8st+6a 98* 1-4 small _{Martins-Santos (2004)}Borin and

P. absconditus R. Paraná, Porto Rico/PR 56 24m+18sm+8st+6a (98) _{Martins-Santos (2002)}Borin and

P. ornatus R. Paraná, Porto Rico/PR 56 20m+18sm+8st+10a (94) _{Martins-Santos (2002)}Borin and

P. sp 56 30m+14sm+12a 100* Toledo and Ferrari (1976b)

P. sp 26 Scheel (1973)

P. sp R. São Francisco,MG SF 56 40m/sm+16st/a 96* Dias and Foresti (1993)

P. sp R. São Francisco, MG SF 56 32m+12sm+6st+6a (100) Garcia and Moreira Filho ₍₂₀₀₅₎

P. sp R. Iguaçu, PR UP 56 24m+26sm+4st+2a (106) Souza et al. (2004)

P. sp Caxias Reservoir, _{R, Iguaçu, PR} UP 56 30m+14sm+8st+4a 100* 0-4 small Borin and Martins-Santos ₍₂₀₀₄₎

Pinirampus

P. pirinampu R. Paraná, PR UP 50 22m+12sm+4st+12a (84) _{Martins-Santos (2000)}Vasconcelos and

P. pirinampu R. Tibagi, PR UP 50 26m+12sm+2st+10a (88) Swarça et al. (1999)

P. pirinampu R. Paraná, Corrientes, _ARG UP 50 18m+14sm+4st+14a (81) Sanchez and Fenocchio (2005)

P. pirinampu R. Araguari, MG SF 50 Molina and Morelli (2004)

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Genera, Species Locality HB n 2n Karyotype FN Bs Size B HS References Luciopimelodus

L. pati R. Paraná, Corrientes, _ARG UP 50 16m+14sm+8st+12a (80) Sanchez and Fenocchio (2005)

Megalonema

M. platanum R. Paraná, Corrientes, _ARG UP 54 14m+18sm+12st+10a 98 Sanchez and Fenocchio (2005)

Hemisorubim

H. platyrhynchos R. Paraná, PR UP 56 22m+18sm+6st+10a (96) Martins-Santos et al. (1996)

H. platyrhynchos R. Araguai, Barra do _{Garças, MT} PA 56 20m/sm+8st/a Faria et al. (2000)

H. platyrhynchos R. Araguai, Barra do _{Garças, MT} PA 56 Silva et al. (2004)

Pseudoplatystoma

P. coruscans R. Paraná, PR UP 56 18m+16sm+10st+12a (90) Martins-Santos et al. (1996)

P. coruscans Coxim, MS PA 56 42m/sm+14st/a (98) Souza et al. (1992)

P. coruscans R. Mogi-Guaçu,SP UP 56 18m+18sm+10st+10a 92* Bigoni et al (1992)

P. coruscans D. Tres Marias, MG SF 56 20m+12sm+12st+12a (88) Fenocchio (1993)

P. corruscans R. Paraguai, MS PA 56 20m+16sm+8st+12a (92) Swarça et al. (2005b)

P. corruscans R. Paraná, Jupiá/SP; R. _{Paraná, PR} UP 56 26m+10sm+6st+14 a (92) Swarça et al. (2005b)

P. fasciatum R. Solimões, AM AM 56 18m+14sm+10st+14a (88) Fenocchio and Bertollo (1992)

P. tigrinum R. Solimões, AM AM 56 18m+16sm+8st+14a (90) Fenocchio and Bertollo (1992)

Paulicea/Zungaro

P. luetkeni R. Paraná,PR UP 56 26m+10sm+6st+14a (92) Martins-Santos et al. (1996)

Z. zungaro R. Paraná, Jupiá, SP UP 56 32m+6sm+8st+10a (94) Swarça et al. (2001a)

Sorubim Silva et al. (2004)

S. lima R. Solimões, AM AM 56 18m+12sm+14st+12a 86 Fenocchio and Bertollo (1992)

S. lima R. Paraná,PR UP 56 20m+14sm+10st+12a (90) Martins-Santos et al. (1996)

S. lima R. Araguai, Barra do _{Garças, MT} PA 56

Steindachneridion

S. scriptum R. Paranapanema, R. _{Tibagi, PR} UP 56 24m+20sm+4st+8a (100) Swarça et al. (2005a)

S. melanodermatum R. Iguaçu,PR UP 56 20m+24sm+2st+10a _{21m+23sm+2st+10a} (100) XX/_XY Swarça et al. (2006)

For abbreviations see legend to table III.

Table 2. Cytogenetic data of the Heptapteridae Family

Genera, Species Locality HB n 2n 3n Karyotype FN Bs Size B HS References Pimelodella

P. kronei Iporanga, SP UP 58 54m/sm+4st (112) 1 micro Almeida-Toledo et al. ₍₁₉₉₂₎

P. transitoria Iporanga, SP UP 58 54m/sm+4st (112) Almeida-Toledo et al. ₍₁₉₉₂₎

P. sp R. Mogi-Guaçu, SP UP 46 40m/sm+6st/a 86 XX/_XY Dias and Foresti (1993)

P. sp R. Mogi-Guaçu, R. _{Pardo, SP} UP 46 28m+10sm+8a 84 Toledo and Ferrari (1976a)

P. sp R. Araquá;R. Capivara, _SP UP 46 32m,sm+14st/a 78* Braga (1989)

P. sp Argentina PA 46 Fenocchio et al. (1994)

P. sp R. Tibagi, PR UP 46 34m/sm+12st/a 80* Silva et al. (1996)

P. sp1 R. Paraná, PR UP 46 20m+20sm+6st (86) _{Martins-Santos (2000)}Vasconcelos and

P. sp2 R. Paraná, PR UP 52 22m+22sm+8st (96) _{Martins-Santos (2000)}Vasconcelos and

P. avanhandavae R. Araquá; R. Capivara, _SP UP 46 20m+20 sm+6 st 86 Vissotto et al. (1999a)

P. avanhandavae R. Machado, MG 58 48m+10st Pereira and Maistro (2002)

P. aff.

avanhandavae R. Tibagi, PR UP 52 30m+22sm (82) Swarça et al. (2003)

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Genera, Species Locality HB n 2n 3n Karyotype FN Bs Size B HS References

P. meeki Couro do Boi;R. Tibagi, R.

R. Gabriel da Cunha, PR UP 46 30m+12sm+4st (88) Vidotto et al. (2004)

P. cristata R. Araguai, Barra do _{Garças, MT} PA 52/46 36m/sm+20st/a 92* Faria et al. (2000)

P. gracilis R. Paraguai, MS PA 52 Mazzuchelli et al. (2004)

Rhamdella

R.sp Itaetê, BA SF 56 26m/sm+30st/a 82* Souza et al. (1994)

R. microcephala R. Machado, MG SF 56 18m+30sm+8st 104 Fonseca et al. (2003)

Cetopsorhamdia

C. iheringi D. Três Marias, MG;_{R. das Marrecas, SP} SF; _UP 58 22m+16sm+10st+10a 96* Fenocchio et al. (2003a)

C. iheringi R. Capivara, R. Pardo, SP UP 58 28m+24sm+6st 110 Vissotto et al. (1999a)

C. sp b. Canta Galo, SP UP 58 22m+16sm+10st+10a 96* Fenocchio et al. (2003a)

Rhamdia

R. hilarii R. Onça, SP UP 63 116 Toledo and Ferrari (1976a)

R. hilarii D. Monjolinho, SP UP 58 >100 0-5 small Fenocchio and Bertollo ₍₁₉₉₀₎

R. hilarii D. Lobo, SP UP 58 >100 0-3 small Fenocchio et al (2000)

R. hilarii D. 29, SP UP 58 >100 0-5 small Fenocchio et al. (2000)

R. hilarii R. Mogi-Guaçu, SP UP 58 >100 Fenocchio et al. (2000)

R. hilarii R. São Francisco, MG SF 58 >100 0-2 small Fenocchio et al. (2000)

R. hilarii R. Aguapey, Corrientes, _ARG PA 58 26m+16sm+8st+8a (100) Fenocchio et al. (2003a)

R. hilarii L. Nova; L. Jataí, SP UP 58 >100 0-2 small Fenocchio (1993)

R. hilarii R. Mogi-Guaçu, SP UP 58 58m/sm 116* 0-2 medium Maistro et al. (2002)

R. cf. hilarii S. Hortelã, S. Quinta, R. Araquá, b. Jacutingá, R.

Pardo, S. Jurumirim, SP UP 58 30m+18sm+10st 106 0-3 metacentric Vissotto et al. (1999b)

R. sp Sapucaí, D. Furnas, MG SF 58 0-3 Andrade et al (1998)

R. sp R. Iguaçu, Usina de _{Salto Segredo, PR} UP 58 36m+14sm+4st+4a (108) 0-2 small Abucarma et al. (2001)

R. sp S. Grande, SP UP 58 46m/sm+12st 104* 0-4 metacentric Garcia et al. (2003)

R. sp S. Grande, SP UP 87 69m/sm+18st 156* Garcia et al. (2003)

R. sp R. São João, PR UP 58 26m+10sm+10st+12a 94* Roman et al. (2001)

R branneri R. Iguaçu, Usina de Salto _{Segredo, PR} UP 58 36m+14sm+4st+4a (108) 0-4 medium Abucarma et al. (2001)

R. branneri R. iguaçu, PR UP 58 30m+10sm+14st+4a 98* 0-4 medium Roman et al. (2002a)

R. voulezi R. Iguaçu, Usina de Salto _{Segredo, PR} UP 58 36m+14sm+4st+4a (108) 0-2 medium Abucarma et al. (2001)

R. voulezi R. Iguaçu, Quedas do _{Iguaçu, PR} UP 58 30m+10sm+14st+4a 98* Roman et al. (2001)

R. quelen R. Iguaçu, PR UP 58 32m+16sm+ 6st+4a (106) 0-1 small Fenocchio et al. (2003b)

R. quelen R. Paraná, Posadas, ARG PA 58 26m+16sm+8st+8a (100) Fenocchio et al. (2003a)

R quelen L. Nova and Jataí, SP UP 58 0-4 small Fenocchio (1993)

R. quelen _{L. dos Quadros, RS}R. Guaíba; PA 58 52m,sm,st+6a 110 0-4 small Hochberg and Erdtmann ₍₁₉₈₈₎

R. quelen R. Iguaçu, Porto União, _SC UP 58 0-1 Fenocchio et al. (2000)

R. quelen R. Tibagi, PR UP 58 0-4 Carvalho and Dias (2001)

R. quelen R. Taquarussu, SP UP 58 26m+20sm+6st+6a 104* 1-4 medium/_large _{Martins-Santos (2004)}Stivari and

R. quelen S. Maringá, PR UP 58 26m+22sm+6st+4a_26m+24sm+8st 106*_108* _{Martins-Santos (2004)}Stivari and

R. quelen Canal São Gonçalo, B. B. Sarandi, L. Fragata,

Santa Izabel, RS 58 0-3 Born (2002)

R. quelen S. Lindóia, Londrina, PR UP 58 30m+14sm+10st+4a (102) Tsuda (2005)

R. quelen S. Lindóia, Londrina, PR UP 87 45m+21sm+30st+12a 174* Tsuda (2005)

R. quelen R. São Francisco, MG SF 58 40m+6sm+8st+4a (104) 0-4 meta/micro Garcia (2005)

R. quelen R. Uberabinha, MG SF 58 1-4 Guilherme et al. (2004)

R. quelen R. Aguas das Pedras, PR UP 58 Mattanó and Dias (2004)

R. sapo Buenos Aires, Arg PA 58 44m/sm+14st/a 102 0-1 medium Valcarcel et al. (1993)

R. sapo Uruguai PA 28 56 Gonzales (1994)

Heptapterus

H. longicauda S. Quinta, SP UP 52 22m+26sm+4st 100 Vissotto et al. (1999a)

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and arm rates (L

EVAN

et al. 1964). In order to

determine the NF we considered as monoarmed

only the acrocentric ones.

RESULTS AND DISCUSSION

Along the last thirty years, from the ﬁrst

de-scription of the chromosomal number of a

Pime-lodidae specie, in 1973 by S

CHEEL

, until 2006,

were analyzed cytogenetically twenty four genera

and ﬁfty three species (48 species valid) of

Pime-lodidae, Heptapteridae and Pseudopimelodidae

families. These numbers were increased in four

genera and seventeen species from the last

revi-sion made by S

WARÇA

et al. (2000) and at now

the diploid number varies from 2n=42 to 2n=58,

not considering the diploid number of 2n=62 in

Rhamdia hilarii described by T

OLEDO

and F

ERRARI

(1976a). Additionally have been found three cases

of triploids with 3n=87 in Rhamdia branneri (R

O

-MAN

et al. 2002) Rhamdia sp (G

ARCIA

et al. 2003)

and Rhamdia quelen (T

SUDA

2005). In the present

study to compare the fundamental number (FN)

of all species some of them were recalculated (in

parentheses) and the others were calculated for

the ﬁrst time (asterisk), considering M and SM

with two arms and ST-A with one arm. The three

families have a relatively high FN (from 74 to

116), due to the predominance of biarmed

chro-mosomes and not considering the triploids and

supernumerary chromosomes (Tables 1, 2, 3).

In the family Pimelodidae 23 of the 27

kary-otyped species, have a diploid number of 2n=

56 chromosomes, being exceptions Calophysus

macropterus. Luciopimelodus plati and

Piniram-pus pirinampu with 2n=50 and Megalonema

pla-tanum with 2n=54, which seems to be share

an-other characteristics (Table 1). The species with

56 chromosomes could be divided in two groups,

the “Pimelodus group” and the “Sorubiminae

group” (=Sorubinae), according to their

particu-lar cytogenetic attributes, i.e., the NORs localized

on one single chromosome pair but in the ﬁrst one

in terminal position in long arm, and in the

sec-ond one in terminal position of short arms (F

EN

-OCCHIO

et al. 2003a) (Fig. 1).

Karyotypes with 56 chromosomes represent a

conservated character for this family, reinforcing

previous observations of L

E

G

RANDE

(1981) and

Genera, Species Locality HB n 2n 3n Karyotype FN Bs Size B HS References Imparﬁnis

I. piperatus R. São João, SP UP 56 56m,sm 112* Vicente et al. (1994)

I. cf. piperatus R. Juquiá, SP UP 56 24m+12Ssm+20st (92) Fenocchio et al. (2003a)

I. cf. piperatus R. Juquiá, SP UP 56 22m+26sm+4st+4a 104 Vissotto et al. (2001)

I. piperatus R. Araras, SP UP 58 32m+26sm 116 Vissotto et al. (2001)

I. piperatus R. Grande, SP UP 58 26m+22sm+8st+2a 106 Vissotto et al. (2001)

I. aff. schubarti b. Canta Galo, SP UP 58 22m+16sm+10st+10a (96) Fenocchio et al. (2003a)

I. aff. schubarti R. Tibagi, PR UP 58 28m+28sm+2st (114) Stolf et al. (2004)

I. mirini b. Jacutingá, S. Quinta, _SP UP 58 24m+34sm 116 ZZ/_ZW Vissotto et al. (1997)

Pariolius

P. cf. longicaudus R. Tibagi, PR UP 52 22m+16sm+4st+10a 90* Garcia et al. (2004)

P. hollandi R. Iguaçu, PR UP 42 22m+10sm+4st+6a 74* Roman et al. (2002b)

For abbreviations see legend to Table 3.

Table 3. Cytogenetic data of the Pseudopimelodidae Family

Genera, Species Locality HB n 2n Karyotype FN Bs Size B HS References Lophiosilurus

L. alexandri D. Três Marias, MG 54 54m/sm/st/a Marques et al. (2002)

Microglanis

M. garavelloi r. Araquá, R. Capivara, SP UP 54 22m+20sm+12st 96* Vissotto et al. (1999b)

Pseudopimelodus

P. bufonius R. Capim, Pará 54 18m+22sm+6st+8a 94* Souza et al. (2003b)

P. mangurus R. Mogi-Guaçu, SP 54 6m+26sm+12st+10a 86* Martinez et al. (2004)

P. zungaru R. São Francisco, MG SF 54 30m+14sm+10a (98) Garcia (2005)

HB = Hydrograﬁc basin; n = haploid number; 2n = diploid number; FN = fundamental number; Bs = supernumerary chro-mosome; HS = heterogametic Sex; m = metacentrics; sm = submetacentrics; st = subtelocentrics; a = acrocentrics; R. = river; b = brook; L. = lagoon; D = Dam; PA = Paraguay; UP = Upper Paraná; SF = São Francisco; AM = Amazon; MG, AM, SP, PR, RS, MS, MT, BA = Brazilian states; () FN recalculated; * FN calculated

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K

LINKHARDT

(1998). O

LIVEIRA

and G

OSZTONYII

(2000) reported that in thirteen genera of

Sil-uriformes have been reported 2n=56, suggesting

that this could be the basal chromosome number

for the Order. However, as in some groups have

been few available cytogenetic data,

generaliza-tions should be carefully revised.

The variability in chromosome number is

greater among species of the family

Heptapteri-dae ranging from 2n=42 in Pariolius to 2n=58 in

Rhamdia hilarii (Table 2) and this feature suggests

that extensive chromosomal rearrangements were

involved in speciation within this group. In this

family were analyzed twenty-two species, twelve

having 2n= 58 (more than 50%).

The family Pseudopimelodidae is smaller than

Pimelodidae and Heptapteridae with only forty

species and cytogenetic data on this group of ﬁsh

are scarce and relatively hard to collect, where

only three genera and ﬁve species were analysed

all of them with 2n=54 chromosomes (Table 3).

Mistakes and changes in speciﬁc names are

a frequent trouble in this group, i. e. in a recent

taxonomic revision of the genus Steindachneridion

was described “Steindachneridion

melanoderma-tum” (G

ARAVELLO

2005), endemic from the Iguaçu

river (Paraná State, Brazil), this specie was

cytoge-netically characterized by S

WARÇA

et al. (2006),

being named by the authors as Steindachneridion

sp. Another case is Microglanis garavelloi from

upper Paraná river, identiﬁed as new species by

S

HIBATTA

and B

ENINE

(2005) but that had been

katyotypically studied as M. cottoides by V

ISSOTO

et al. (1999). The genus Rhamdia also shows cases

of synonymy, being reported cytogenetic studies

about R. voulezzi and R. branneri, that are

consid-ered synonymous of R. quelen (S

ILFVERGRIP

1996).

Have been reported cases with two diploid

num-bers Pimelodus blochii of Solimões River and

Ara-guaia River with 2n=58 and 56 respectively, and

Pimelodus fur with 2n=54 and 56 chromosomes

(Table 1), but the taxonomic identiﬁcation of these

species as well as the numbers should be revised.

In the Pimelodidae family, B chromosomes

have been identiﬁed in Bergiaria westermanni,

Iheringichthys labrosus, Pimelodus ortmanni,

Pimelodus sp (Table 1), and in the

Heptapteri-dae family in Pimelodella kronei, Rhamdia hilarii,

Rhamdia sp, R. branneri, R. voulezi, R. quelen, R.

sapo (Table 2). This extra chromosome is highly

Fig. 1. — Hypothetical phylogenetic tree modiﬁed from LUNDBERG et al. (1988; 1991) and FENOCCHIO et al. (2003a);

based in new available cytogenetic data. The interrogation signs suggest probable dichotomy ways of branch. As-terisks indicate cytogenetically studied species.

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conserved in the genus Rhamdia being found in

several populations of these species along a wide

geographic area.

Systems of sexual chromosomes have been

reported only in Steindachneridion

melanoderma-tum (Pimelodidae) (Table 1) and Pimelodella sp

(Heptapteridae) (Table 2) where the male is the

heterogametic sex, possessing a system of the

type XX/XY. V

ISSOTTO

et al. (1997) described in

Imparﬁnis mirini a ZZ/ZW system, but later the

same authors rectiﬁed the ﬁrst information (V

IS

-SOTTO

2000). At now the only confirmed sex

chro-mosome system in Pimelodidae corresponds to S.

melanodermatum (S

WARÇA

et al. 2006).

Among all the species analyzed (Tables 1 and

2), Pimelodus maculatus, Rhamdia hilarii and

Rhamdia quelen, are the most studied until now,

demonstrating, a very wide geographic

distribu-tion of these species along the South American

river basins associated with a very high facility of

capture.

According to R

EIS

et al. (2003) the

Heptapteri-dae family is composed by 238 species (valid and

undescribed), the Pimelodidae by 128 and

Pseu-dopimelodidae by 40, but were analyzed

citoge-netically only around ﬁfty species in these three

families of Siluriformes, this fact demonstrates

that more species must be studied to increase the

number of cytogenetic data for a better

under-standing of the species relationships and

karyo-typic evolution in this ﬁsh group. In spite of this,

could be suggested a preliminary version of a

phylogenetic tree, on the basis of available

phy-logenetic data and the chromosome features

sum-marized in the present paper (Fig. 1).

REFERENCES

ABUCARMA M. and MARTINS-SANTOS I.C., 1996 —

Cara-cterização cromossômica de duas espécies da família Pimelodidae (Pisces, Siluriformes). In: Proceedings

of VI Simpósio de Citogenética Evolutiva Aplicada a Peixes Neotropicais, São Carlos SP. pp. 73. —, 2001 — Karyotype and B Chromosome of Rhamdia

species (Pisces, Pimelodidae) Endemic in the River Iguaçu Basin. Cytologia, 66: 299-306.

ALMEIDA-TOLEDO L.F., FORESTI F., TRAJANO E. and TO -LEDO FILHO S.A., 1992 — Cytogenetic analysis of the

Brazilian blind catﬁsh Pimelodella kronei and of its presumed ancestor Pimelodella transitoria.

Caryolo-gia, 45: 255-262.

ANDRADE S.F., MAISTRO E.L., OLIVEIRA C. and FAUSTO

F., 1998 — Caracterização cromossômica da espécie

Rhamdia sp. (Pisces, Pimelodidae), proveniente do Rio Sapucaí, Represa de Furnas, MG. In:

Proceed-ings of 44º Congresso Brasileiro de Genética, Águas de Lindóia, SP. pp. 66.

BIGONI A.P.V., ALMEIDA-TOLEDO L.F. and TOLEDO FIL -HO S.A., 1992 — Estudos citogenéticos em

Pseudo-platystoma corruscans (Pimelodidae, Sorubiminae) do rio Mogi-Guaçu, SP. In: Proceedings of IX

Sim-pósio de Citogenética Peixes Neotropicais pp. 32. BOCKMANN F.A. and GUAZZELI G.M., 2003 — Family

Heptapteridae (Heptapterids). Pp. 406-431. In: R.E.

REIS, S.O. KULLANDER and C.J. FERRARIS JR (eds).

Check list of the freshwater ﬁshes of South and Central America. Edipucrs, Porto Alegre.

BORIN L.A. and MARTINS-SANTOS I.C., 2002 —

Cytoge-netic aspects in species of the genus Pimelodus (Pis-ces, Siluriformes, Pimelodidae) of the river Paraná Basin. Cytologia, 67: 199-204.

—, 2004 — Study on karyotype and occurrence of B

chromosomes in two endemic species of the genus Pimelodus (Siluriformes, Pimelodidae) from the Riv-er Iguaçu. HRiv-ereditas, 140: 201-209.

BORN G.G., 2002 — Análise citogenética de Rhamdia

quelen (Pisces, Rhamdiidae) da região sul do Rio Grande do Sul. In: Proceedings of IX Simpósio de

Citogenética e Genética de Peixes, Maringá, PR pp. 79.

BRAGA M.A., 1989 — Estudos citogenéticos em

Pime-lodella sp. (Pisces, Siluriformes). Monograﬁa de

Ba-charelado, Instituto de Biociências, UNESP Botu-catu, SP.

CARVALHO R.A. AND DIAS A.L., 2001 — Caracterização

citogenética de Rhamdia quelen (Pisces, Rhamdiidae) proveniente da bacia do Rio Tibagi/PR. In:

Proceed-ings of Congresso Nacional de Ictiologia, p. 73. CARVALHO R.A., GIULIANO-CAETANO L., DIAS A.L., 2004

— Cytogenetic Analysis of A- and B- Chromosomes

of Iheringichthys labrosus (Pisces, Pimelodidae) from the Tibagi River, Paraná, Brazil. Cytologia, 69(4):

381-385.

COSTA L.J. and REGGI R., 1986 — Estudos

prelim-inares de duas espécies da família Pimelodidae: Pimelodus maculatus e Parapimelodus valenciennes do rio Guaiba, RS. In: Proceedings of I Simpósio

de Citogenética Evolutiva Aplicada em Peixes Neo-tropicais, São Carlos, SP. pp. 34.

DELLA-ROSA V.A., BERTOLLO L.A.C., FERRARI I., TAKA -HASHI C.S., MOREIRA FILHO O. and FORESTI F., 1980

— Estudos citogenéticos de peixes da Amazônia. II.

Ordem Siluriformes. Ciênc. Cult. 32:735

(proceed-ings).

DIAS A.L. and FORESTI F., 1990 — Algumas

consider-ações a respeito do cariótipo de Iheringichthys labro-sus (Siluriformes, Pimelodidae) do rio Mogi-Guaçu.

In: Proceedings of III Simpósio de Citogenética Evolutiva Aplicada em Peixes Neotropicais., Botu-catu, SP. pp. 32.

—, 1993 — Cytogenetic studies on ﬁshes of the family

Pimelodidae (Siluroidei). Rev. Bras. Genet., 16:

585-600.

FARIA A.A., BRITO J.G. and VENERE P.C., 2000 —

(8)

cromo-ssômica de Pimelodus blochii, Pimelodella cristata e Hemisorubim platyrhynchos (Siluriformes) do mé-dio Araguaia. In: Proceedings of VIII Simpósio de

Citogenética e Genética de Peixes, Manaus, AM. pp 92.

FENOCCHIO A.S., 1993 — Cromossomos

supranumer-ários no gênero Rhamdia (Pisces). Caracterização cromossômica e considerações sobre a evolução cari-otípica nos Siluroidei. PhD Thesis. Faculdade de

Medicina de Ribeirão Preto, Universidade de São Paulo.

FENOCCHIO A.S. and BERTOLLO L.A.C., 1988 — A

sim-ple method for fresh-water ﬁsh lymphocyte culture.

Brazilian Journal of Genetics, 11(4):847-852. FENOCCHIO A.S. and BERTOLLO L.A.C., 1990 —

Super-numerary chromosomes in a Rhamdia hilarii popula-tion (Pisces, Pimelodidae). Genetica, 81:193-198.

FENOCCHIO A.S. and BERTOLLO L.A.C., 1992 —

Karyo-type similarities among Pimelodidae (Pisces, Sil-uriformes) from the Brazilian Amazon region.

Cyto-bios, 69: 41-46.

FENOCCHIO A.S., VENERE P.C., CESAR A.C.G., DIAS A.L.

and BERTOLLO L.A.C., 1991 — Short termculture

from solid tissues of ﬁshes. Caryologia, 44:161-166.

FENOCCHIO A.S., PASTORI M.C., LOPEZ P.A., SANCHEZ

S., ALBERDI A.J., BORDENAVE S. and DIB M.C., 1994

— Levantamento citogenético em peixes de

água-doce da Argentina: resumo das espécies estudadas.

In: Proceedings of V Simpósio de Citogenética Ev-olutiva Aplicada a Peixes Neotropicais, Botucatu SP, pp. 8.

FENOCCHIO A.S., BERTOLLO L.A.C., TAKAHASHI C.S. and

CAMACHO J.P., 2000 — Considerations on

B-Chro-mosome origin and distribution in ﬁsh populations of the genus Rhamdia (Siluriformes, Pimelodidae).

Folia Biologica, 48: 105-109.

FENOCCHIO A.S., BERTOLLO L.A.C., TAKAHASHI C.S.,

DIAS, A.L. and SWARÇA A.C., 2003a — Cytogenetic

Studies and Correlated Considerations on Rhamdii-nae Relationships (Pisces, Siluroidei, Pimelodidae).

Cytologia, 68(4): 363-368.

FENOCCHIO A.S., SWARÇA A.C., CESTARI M.M. and DIAS

A.L., 2003b — Karyotypic characterization and NOR

analysis by different banding techniques of Rhamdia quelen (Pisces, Pimelodidade) from the ﬁrst plateau of the Iguaçu River. Folia Biologica, 51(3-4):

219-222.

FERRARIS C.J. JR., 2007 — Checklist of catﬁshes, recent

and fossil (Osteichthyes: Siluriformes), and catalogue of siluriform primary types. Zootaxa, 1418, 1-628.

FONSECA Y.M., OLIVEIRA C., FORESTI F. and MAISTRO

E.L., 2003 — First cytogenetic description of the

spe-cies Rhamdela microcephala (Pisces, Heptapteridae).

Cytologia, 68(1): 31-34.

GARAVELLO J.C., 2005 — Revision of genus

Steindachne-ridion (Siluriformes: Pimelodidae). In: All catﬁshe

Species Inventory. Neotropical Ichthyology, 3(4): 607-623.

GARCIA R.M.G., SACHETE S. and MARTINS-SANTOS I.C.,

1990 — Aspectos citogenéticos de Iheringichthys

la-brosus (Pisces, Pimelodidae) do rio Paraná, Região de Porto Rico, PR. In: Proceedings of III Simpósio

de Citogenética Evolutiva Aplicada em Peixes Neo-tropicais., Botucatu, SP. pp. 32.

GARCIA C., 2005 — Contribuição aos estudos

citogenéti-cos em algumas espécies de 5 famílias de Siluriformes do rio São Francisco. Master Thesis. Universidade

Federal de São Carlos, SP. P104.

GARCIA C., MOREIRA-FILHO O., BERTOLLO L.A.C. and

CENTOFANTE L., 2003 — B Chromosomes and

Natu-ral Triploidy in Rhamdia sp. (Pisces, Siluriformes, Heptapteridae). Cytologia, 68(4): 403-411.

GARCIA C. and MOREIRA-FILHO O., 2005 —

Cytoge-netical analyses in three ﬁsh species of the genus Pimelodus (Siluriformes: Pimelodidae) from Rio São Francisco: Considerations about the karyotypi-cal evolution in the genus. Neotropikaryotypi-cal Ichthyology,

3(2): 285-289.

GARCIA J.B.F., DIAS A.L., FENOCCHIO AS.F., 2004 —

Caracterização cromossômica de Pariolius cf. longi-caudus (Pisces, Pimelodidae) do Rio Tibagi (PR).

In: Proceedings of X Simpósio de Citogenética e Genética de Peixes, Natal RN pp. 144.

GONZALES A., 1994 — Estudio preliminar de tres

espe-cies pertenecientes al orden Siluriformes (Pisces). In:

Proceedings of V Simpósio de Citogenética Evolu-tiva Aplicada a Peixes Neotropicais, Botucatu SP. pp. 21.

GUILHERME L.C., MARIANO A.R.T., MOLINA R.A.S.,

PAULA A.T., BERNARDES L.S. and MORELLI S., 2004

— Presença de cromossomos B em uma população de

Rhamdia quelen (Pisces, Rhamdiidae) do Rio Uber-abinha no Município de Uberlândia-MG. In:

Pro-ceedings of X Simpósio de Citogenética e Genética de Peixes, Natal RN pp. 146.

HERAS M.P. and MENDOZA N.R., 2002 — Analísis

cari-otípico de Pimelodus maculatus (LACÉPÈDE, 1803) (Pisces, Siluriformes, Pimelodidae) provenientes del delta Paranaense. In: Proceedings of XXXI

Con-greso Argentino de Genetica, La Plata, Argentina. pp. 95

HOCHBERG V.B.M. and ERDTMANN B., 1988 —

Cytoge-netical and morphological considerations on Rham-dia quelen (Pisces, Pimelodidae) – the occurrence of B chromosomes and polymorphic NOR regions. Rev.

Bras. Genet., 11: 563-576.

KLINKHARDT M.B., 1998 — Some aspects of

karyoevolu-tion in ﬁshes. Animal Res. Development, 47: 7-36.

LEGRANDE W.H., 1981 — Chromosomal evolution in

North American Catﬁshes (Siluriformes, Ictaluridae) with particular emphasis on the madtoms, Noturus.

Copeia, 1: 33-52.

LUNDBERG J.G., LINARES O.J., ANTONIO M.E. and NASS

P., 1988 — Phractocephalus hemiliopterus

(Pimelodi-dae, Siluriformes) from the upper Miocene Urumaco Formation, Venezuela: a further case of evolutionary stasis and local extinction among South American ﬁshes. J. Vert. Paleo, 8: 131-138.

LUNDBERG J.G., BORNBUSH A.H. and MAGO LECCIA

(9)

Ecuador with diagnoses of the subfamilies Rham-diinae Bleeker and Pseudopimelodidae N. Subf. (Sil-uriformes, Pimelodidae). Copeia, 1: 190-209.

LUNDBERG J.G. and LITTMANN M.W., 2003 — Family

Pimelodidae (Long-whiskered catﬁshes). Pp.

432-446. In: R.E. REIS, S.O. KULLANDERAND C.J. FER -RARIS JR (eds). Check list of the freshwater fishes of

South and Central America. Edipucrs, Porto Ale-gre, 729p.

MAISTRO L.M., OLIVEIRA C. and FORESTI F., 2002 —

Cy-togenetic Analysis of A- and B- Chromossomes of Rhamdia hilarii (Teleostei, Pimelodidae): C-Band-ing, Silver Nitrate and CMA3 Staining and Restric-tion Endonuclases Banding. Cytologia, 67: 25-31.

MARQUES S., MAISTRO E.L., OLIVEIRA C. and FORESTI F.,

1998 — Estudos cariotípicos na espécie Pimelodus

maculatus (Pisces, Pimelodidae) coletada no rio Sapucaí, represa de Furnas, MG. In: Proceedings

of 44º Congresso Brasileiro de Genética, Águas de Lindóia SP pp. 59.

MARQUES M.B., GARCIA C. and MOREIRA FILHO O.,

2002 — Estudos Citogenéticos de duas espécies de

Siluriformes endêmicas do rio São Francisco. In:

Pro-ceedings of IX Simpósio de Citogenética e Genética de Peixes, Maringá PR, pp. 89.

MARTINEZ E.R.M., OLIVEIRA C. and FORESTI F., 2004

— Cytogenetic Analyses of Pseudopimelodus

man-gurus (Teleostei: Siluriformes: Pseudopimelodidae).

Cytologia, 69(4): 419-424.

MARTINS-SANTOS I.C., JULIO JR H.F. and BURIN I., 1996

— Karyotypic studies of four species of the

Sorubimi-nae subfamily (Pisces, Siluriformes). Caryologia,

49(1): 73-80.

MATTANÓ H. and DIAS A.L., 2004 — Dados

cromossômi-cos preliminares em Rhamdia quelen (Siluriformes, Pimelodiade) da Bacia do Rio Tibagi-PR. In:

Pro-ceedings of X Simpósio de Citogenética e Genética de Peixes, Natal, RN, pp. 131.

MAZZUCHELLI J., VIDOTTO A.P. and DIAS A.L., 2004

— Estudos cirtogenéticos preliminares em

Pime-lodella gracilis (Siluriformes, Pimelodidae) da Bacia do Rio Paraguai. In: Proceedings of X Simpósio

de Citogenética e Genética de Peixes, Natal RN, pp. 139.

MAZZUCHELLI J., DIAS A.L., FENOCCHIO A.S. and

SWARÇA A.C., 2006 — Caracterização

cromossômi-ca de Pimelodus maculatus (Pisces, Pimelodidae).

XXVI Congresso Brasileiro de Zoologia, Londrina Pr. CD-Rom.

MAZZUCHELLI J. and DIAS A.L., 2006 — Contribuição

aos estudos cromossômicos de Pimelodus maculatus (Pisces, Pimelodidae) coletados no Ribeirão Con-gonhas, Paraná. XXVI Congresso Brasileiro de

Zoologia, Londrina Pr. CD-Rom.

MOLINA R.A.S. and MORELLI S., 2004 — Análise

Citogenética de uma população de Pinirampus piri-nampu (Pisces, Siluriformes) da Bacia do Rio Ara-guari (Uberlândia, MG). In: Proceedings of X

Sim-pósio de Citogenética e Genética de Peixes, Natal RN, pp. 143.

MORAES V.P.O. and DIAS A.L., 2006 — Análise

citogenética comparativa em quatro populações de Rhamdia quelen (Pisces, Heptapteridae). XXVI

Congresso Brasileiro de Zoologia, Londrina, Pr. CD-Rom.

MOREIRA R.G., MORELLI-SHIMIZU L., MORELLI S., 2004

— Aspectos citogenéticos de duas populações do

mandi amarelo Pimelodus maculatus (Siluriformes, Pimelodidae). In: Proceedings of X Simpósio de

Citogenética e Genética de Peixes, Natal, RN, pp. 145.

OLIVEIRA C. and GOSZTONYI A.E., 2000 — A

cytoge-netic study of Diplomystes mesembrinus (Teleostei, Siluriformes, Diplomystidae) with a discussion of chromosome evolution in siluriforms. Caryologia,

53(1): 31-37.

PEREIRA M.G.A.G. and MAISTRO E.L., 2002 —

Caracteri-zação citogenética da espécie Pimelodella avanhanda-vae (Pisces, Pimelodidae) provenientes da nascente do Rio Machado, MG. 48º Congresso Brasileiro de

Genética, Águas de Lindóia SP. CD-ROM.

DE PINNA M.C.C., 1998 — Phylogenetic relationship of

Neotropical Siluriformes (Teleostei, Ostariophisy): Historical overview and synthesis of hypotheses.

Pp. 279-330. In: L.R. MALABARBA, R.E. REIS, R.P.

VARI, Z.M. LUCENA and C.A.S. LUCENA (eds).

Phyl-ogeny and Classiﬁcation of Neotropical ﬁshes. Ed-ipucrs, Porto Alegre, 603p.

RAMIREZ-GIL H., FELDEBERG E., ALMEIDA-VAL V.M.F.

and VAL A.L., 1998 — Karyological, biochemical,

and physiological aspects os Callophysus macropterus (Siluriformes, Pimelodidae) from the Solimões and Negro Rivers (Central Amazon). Braz. J. Med. Biol.

Res., 31(11): 1449-1458.

REIS R.E., KULLANDER S.O. and FERRARIS JR C.J., 2003

— Check list of the freshwater ﬁshes of South and

Central America. Edipucrs Porto Alegre, 729 p.

ROMAN M.P, PEGORARO J.L. and MARGARIDO V.P., 2001

— Análises citogenéticas em 4 espécies de Siluriformes

(Pisces) com ênfase ao padrão de distribuição da het-erocromatina constitutiva. 47º Congresso Nacional

de Genética, Águas de Lindóia SP. CD-ROM ROMAN M.P., MOREIRA-FILHO O. and MARGARIDO V.P.,

2002a — O cariótipo de Rhamdia branneri (Pisces,

Siluriformes, Pimelodidae) do rio Iguaçu e um caso de triploidia natural. In: Proceedings of IX

Simpó-sio de Citogenética e Genética de Peixes, Maringá PR pp. 76.

—, 2002b — Análise Citogenética em Pariolius hollandi

(Pisces, Siluriformes, Pimelodidae) do Rio Iguaçu.

In: Proceedings of IX Simpósio de Citogenética e Genética de Peixes, Maringá PR, pp. 75.

SANCHEZ S., 2006 — Estudios Citogeneticos en Peces

de la Família Pimelodidae (Pisces, Siluriformes) de la Cuenca del Rio Paraná, Argentina. PhD Thesis

Ciencias Biológicas. Universid Nacional de Cór-doba, Argentina.

SCHELL J.J., 1973 — Fish Chromosomes and their

Evo-lution. Interval Report of Danmarks Akvarium,

(10)

SHIBATTA O.A., 2003 — Family Pimelodidae

(Bumble-bee catﬁshes, dwarf marbled catﬁshes). Pp. 401-405.

In: R.E. REIS, S.O. KULLANDER and C.J. FERRARIS JR

(eds). Check list of the freshwater ﬁshes of South and Central America. Edipucrs, Porto Alegre. SHIBATTA A.O. and BENINE R.C., 2005 — A new species

of Microglanis (Siluriformes: Pimelodidae) from up-per rio Paraná basin, Brazil. In: All catﬁshes Species

Inventory. Neotropical Ichthyology, 3(4): 579-585. SILFVERGRIP A.M.C., 1996 — A Systematic revision of

the Neotropical catﬁsh genus Rhamdia (Teleostei, Pimelodidae). Departament of Zoology, Stockholm

University and Departament of Vertebrate Zoology, Swedish Museum of Natural History, Stockholm, pp. 95-99.

SILVA C.T., DIAS A.L. and GIULIANO-CAETANO L., 1996

— Análise citogenética em Iheringhthys labrosus

e Pimelodella sp. (Pisces, Pimelodidae) da bacia do rio Tibagi. In: Proceedings of VI Simpósio de

Citogenética Evolutiva Aplicada a Peixes Neotrop-icais, São Carlos, SP. pp 76.

SILVA, L.V.B., ABREU, M.F., SOUZA, I.L. and VENERE,

P.C., 2004 — Caracterização Cromossômica de

Hem-isorubin platyrhynchos, Sorubim lima e Pimelodus blochii (Pisces, Siluriformes, Pimelodidae) do Mé-dio Araguaia. In: Proceedings of X Simpósio de

Citogenética e Genética de Peixes, Natal, RN pp. 132.

SOUZA, A.B., FONSECA, C.G., PINHEIRO, L.E.L. and RIBEI -RO, L.P., 1992 — Estudos citogenéticos preliminares

em Pseudoplatystoma corruscans (Siluriformes, Pime-lodidae) da bacia do rio Paraguai. In: Proceedings of

IV Simpósio de Citogenética Evolutiva Aplicada a Peixes Neotropicais, Rio de Janeiro, RJ. Pp. 28. SOUZA I.L., MOREIRA-FILHO O. and CASTRO R.M.C.,

1994 — Informações citogenéticas preliminares de

uma nova espécie de Pimelodidae cego de cavernas (Itaête, BA). In: Proceedings of V Simpósio de

Citogenética Evolutiva Aplicada a Peixes Neotrop-icais, Botucatu SP, pp 18.

SOUZA A.C.P., NAGAMACHI C.Y., RISSINO J.D., JUNIOR

J.R.C., BARROS R.M.S. and PIECZARKA J.C., 2000

— Descrição cariotípica de Pimelodus cf. maculatus

(Siluriformes, Pimelodidae). In: Proceedings of VIII

Simpósio de Citogenética e Genética de Peixes, Manaus AM, pp. 90.

SOUZA L., GIULIANO-CAETANO L. and DIAS A.L., 2003a

— Karyotipic Study of three species of Pimelodus

(Pisces, Pimelodidae) from the Paraguai River Basin.

Cytologia, 68(4): 345-350.

SOUZA L., GIULIANO-CAETANO L. and SHIBATTA O.,

2003b — Citogenética de Pseudopimelodus bufonius

(Siluriformes, Pseudopimelodidae). 49º Congresso

Nacional de Genética, Águas de Lindóia, São Pau-lo, Brazil. CD-Rom.

SOUZA L., GIULIANO-CAETANO L. and DIAS A.L., 2004

— Banding chromosome pattern of two species of

Pimelodus (Siluriformes, Pimelodidae) from the Paraná River Basin of Brazil. Folia Biologica,

52(3-4): 165-169.

STIVARI M.K. and MARTINS-SANTOS I.C., 2004 —

Karyo-type Diversity in two Populations of Rhamdia quelen (Pisces, Heptapteridae). Cytologia, 69(1): 25-34.

STOLF R., SWARÇA A.C., GIULIANO-CAETANO L. and DIAS

A.L., 2004 — Analyses of karyotype and nucleolus

organizer regions of Imparﬁnis aff. schubarti (Sil-uriformes, Pimelodidae) of the Tibagi River basin, Paraná, Brazil. Caryologia, 57(4): 348-352.

SWARÇA A.C., GIULIANO-CAETANO L. and DIAS A.L.,

1999 — Cytogenetic characterization through

chro-mosomic banding of Pinirampus pirinampus (Pisces, Pimelodidae) from the Tibagi River basin PR/Brazil.

Caryologia, 52(1-2): 31-35.

—, 2000 — Cytogenetics of species of the families

Pime-lodidae and Rhamdiidae (Siluriformes). Gen. Mol.

Res., 23(3): 589-593.

SWARÇA A.C., CESTARI M.M., GIULIANO-CAETANO L.

and DIAS A.L., 2001a — Cytogenetic

characteriza-tion of the large south American Siluriform ﬁsh spe-cies Zungaro zungaro (Pisces, Pimelodidae). Chrom.

Science, 5: 51-55.

SWARÇA A.C., GIULIANO-CAETANO L. and DIAS A.L.,

2001b — Analyses of nucleolus organizer regions

and heterochromatin of Pimelodus maculatus (Pis-ces, Pimelodidae). Genetica, 110: 97-100.

SWARÇA A.C., VIDOTTO A.P. and DIAS A.L., 2003 —

Cy-togenetic characterization of Pimelodella aff. avan-handavae (Siluriformes, Pimelodidae) from Tibagi River (Paraná State, Brazil). Caryologia, 56(4):

421-425.

SWARÇA A.C., FENOCCHIO A.S., CESTARI M.M. and DIAS

A.L., 2005a — First chromosome data on

Stein-dachneridion scripta (Pisces, Siluriformes, Pimelodi-dae) from Brazilian Rivers: Giemsa, CBG, G- and RE banding. Genetic Mol. Res., 4(4): 734-741.

—, 2005b — Chromosomal divergence among

popula-tions of Pseudoplatystoma corruscans (Pisces, Pime-lodidae): I. Evidences obtained from conventional giemsa staining, C- and G- banding. Ichthol. Expl.

Freshwaters, 16(4): 325-330.

—, 2006 — First report of a sex chromosome

sys-tem in a Sorubiminae ﬁsh, Steindachneridion sp. (Pimelodidae), with an uncommon large Y chro-mosome. Mitotic and Meiotic analysis. Cytogen.

Gen. Res., 112(3-4): 325-328.

TOLEDO V. and FERRARI I., 1976a — Estudo citogenético

de Pimelodella sp. e Rhamdia hilarii (Pimelodinae, Pimelodidae, Pisces): Cromossomo marcador.

Cientí-ﬁca, 4: 120-123.

—, 1976b — Estudos citogenéticos de três espécies do

gênero Pimelodus (Pisces, Pimelodidae). Cientíﬁca,

4:101-106.

TRECO F.R., RUBERT M., DIAS A.L., GIULIANO-CAETANO

L. and MARGARIDO V.P., 2006 -Estudos

Citogenéti-cos em Pimelodus paranaensis (Pimelodidae) coleta-dos no Rio Piquiri, Novas Larangeiras-PR. XXVI

Congresso Brasileiro de Zoologia, Londrina, Pr. CD-Room.

TSUDA J.R., 2005 — Análise citogenética em Rhamdia

(11)

ocorrência de triploidia natural. Monograﬁa de

Ba-charelado, Universidade Estadual de Londrina, Londrina, Paraná.

VALCARCEL A., BRUNNER, P. and MAGGESE M.C., 1993

— B-Chromosome polymorphism in South American

catﬁsh, Rhamdia sapo. Aquaculture, 110: 111-118.

VARI R.P., 1992 — Systematics of Neotropical

Characi-form genus Cyphocharax fowler (Pisces, Ostariophy-si). Smithson Contib. Zool. P., 529.

VASCONCELOS C. and MARTINS-SANTOS I.C., 2000 —

Chromosome polymorphism in species of the Pime-lodidae family (Pisces, Siluriformes). Hereditas, 132:

103-109.

VICENTE M.R.R., MARGARIDO V.P. and GALETTI JR P.M.,

1994 — Estudos cromossômicos em Imparﬁnis

pip-eratus (Pisces, Siluriformes) do Riacho São João (Bacia do Leste). In: Proceedings of V Simpósio de

Citogenética Evolutiva Aplicada a Peixes Neotrop-icais, Botucatu SP, pp. 17.

VIDOTTO A.P., SWARÇA A.C., FENOCCHIO A.S. and DIAS

A.L., 2004 — Cytogenetic studies in three

Pime-lodella meeki populations (Pisces, Pimelodidae)

from Tibagi River Basin (Brazil). Journal of

Hered-ity, 95(6): 517-520.

VISSOTTO P.C., 2000 — Análise citogenética

compara-tiva do sistema de determinação sexual em espécies do gênero Imparﬁnis (Pisces, Siluriformes, Pimelo-didae). Unpublished Ph.D. Thesis. UNESP,

Botu-catu, SP.

VISSOTTO P.C., FORESTI F. and OLIVEIRA C., 1997 — A

ZZ/ZW sex chromosome system in Imparﬁnis mirini (Pisces, Siluriformes). Cytologia, 62: 61-66.

—, 1999a — Karyotype description of ﬁve species of

Pimelodidae (Teleostei, Siluriformes). Chrom.

Sci-ence, 3: 1-7.

—, 1999b — Supernumerary chromosomes in two species

of the family Pimelodidae (Teleostei, Siluriformes).

Chrom. Science, 3: 9-13.

—, 2001 — Karyotypic characterization of two species

of the genus Imparﬁnis (Teleostei, Siluriformes, Hep-tapteridae). Chrom. Science, 5: 97-103.