a checklist of chromosome numbers and karyotypes of...

A checklist of chromosome numbers and karyotypes of Amazonian freshwater jishes

Jorge Ivan Rebelo PORTO (l), Eliana FELDBERG (1), Céleste Mutuko NAKAYAMA (l),

José das Neves FALCAO (2)

RÉSUMÉ

CATALOGUE DES NOMBRES DE CHROMOSOMES ET CARYOTYPES DES POISSONS DU BASSIN AMAZONIEN

Une liste des nombres de chromosomes de 211 espèces de poissons du bassin amazonien est présenfée. Ces espèces, appartenant à 5 ordres, sont caractérisées par une grande variabilité du nombre de chromosomes (2n = 22 à 2n = 134), par la présence de chromosomes sexuels et de polymorphisme chromosomique chez certaines espèces, et enfin par une grande spécificité des bandes C et des sites des régions de l’organisafeur du nucléole.

MOTS CLÉS : Chromosomes - Poissons - Eaux douces - Bassin amazonien.

ABSTRACT

A CHECKLIST OF CHROMOSOME NUMBERS AND KARYOTYPES OF AMAZONIAN FRESHWATER FISHES

A checklist of chromosome numbers of Amazonian freshlvater fishes is presented. 211 nominal species belonging fo 5 orders have had their haploidldiploid number listed. These species are characterized by a high karyotypic diversity including a wide chromosome number range (2n=22 to 2n=134), d ff i erent sex chromosomal mechanisms, chromosomal polymorphisms and nearly always a species-specific pattern of C-banding and nucleolar organizer regions .

KEY WORDS : Chromosomes - Fishes - Freshwaters - Amazon Basin.

RESUMEN

LISTA DE LOS NUMEROS CROMOS~MICOS Y CARIOTIPOS DE PECES AMAZONICOS DE AGUA DULCE

Es presentada una lista con 10s datos cromo&micos de peces de la cuenca amazonica. 211 especies pertenecienfes a 5 ordenes ya fueron estudiados citogeneticamente, determinandose por 10 menos su numero cromosomico haploideldiploide. La variaci0n del numero diploide va desde 2n=22 hasfa 2n=134. Tanbien fueron observado mecanismos cromo&micos sexuales y polimorfismos cromoscimicos en algunas especies, seguido de padrones casi siempre especie-specificos de la banda-C y de las regiones organizadoras del nucleolo.

PALABRAS ~LAVES : Cromosomos - Peces - Agua dulce - Cuenca amazonica.

(1) Instituto National da Pesquisas da Amazônia, Coordenaçüo de Pesquisas em Biologia Aqudtica, Cx Postal 478, Manaus-AM, Brasil, CEP 69083.

(2) Unioersidade do Amazonas, Instituto de Biociências, Deparfamenio de Biologia, Estrada do Contorno S/N, Manaus-AM, Brasil, CEP 69000.

Rerr. Hydrobiol. trop. 25 (4) : 2X7-299 (1992).

288 J. 1. R. PORTO,

INTRODUCTION

E. FELDBERG, C. M. NAKAYAMA, J. N. FALCAO

In the neotropics, chromosomal data have contri- buted t.o t.he studies of the biology, genetics and syst.ematics of fishes. This is particularly true with respect to species characterization and diagnosis (BERTOLLO, 1978; ALMEIDA-TOLEDO, 1978), detec- tion of inter and intraspecific polymorphism (Gru- LIANO-CAETANO and BERTOLLO, 1988; OLIVEIRA et al., 199Oa), data on sex chromosome systems (Mo- REIRA FILHO et al., 1980; GALETTI Jr. et al., 1981; BERTOLLO et al., 1983; GALETTI Jr. and FORESTI, 1986 ; FELDBERG et al., 1987 ; FALCÂO, 1988), supernumerary chromosomes (FALCAO et al., 1984 ; PAUL~, 1985; VENERE and GALETTI Jr., 1985; OLIVEIRA et al., 1988b; FORESTI ef al., 1989; ERDTMAN et al., 1990) and natural triplody (ALMEIDA-TOLEDO et al., 1985; VENERE and GALETTI Jr., 1985; GIULIANO- CAETANO and BERTOLLO, 1990). Al1 of t.he above mentioned findings are important for the understan- ding of flsh or even animal chromosomal evolution.

Checklists cont.aining chromosome data of fish have been published since 1971 (GYLDENHOLM and SCHEEL, 1971; CHIARELLI and CAPANNA, 1973; DENTON, 1973; PARK, 1974; OJIMA et al., 1976; GOLD ef al., 1980; SOLA et al., 1981 ; OLIVEIRA et al., 1988a). SOLA et al. (1981) stated that the elaboration of flsh chromosome lists is important. because they allow for easy access to condensed information and identification of the cytogenet.ic characters important for the elaboration of evolutionary and phylogenetic models.

The Amazon basin is an excellent field for ichthyogenetic study (ALMEIDA-VAL et al., 1991) and we believe that a checklist of Amazonian fish chromosome data Will c.ontribute to a better under- standing of the diversit,y of fish species.

CYTOGENETIC FEATURES OF AMAZONIAN FISHES

In 1988, OLIVEIRA et al. listed the chromosome formular of 433 neotropical fish species. In this study t.he origin of 55 O,& of the fish is unknown, some of them undoubtedly obtained from aquarium dea- lers.

In elaborat.ing our list, we attempted to identify, among the species of unknown origin, the Amazo- nian freshwater species, and we updated the taxono- mit status of many species. Thus, we have listed species belonging to the Amazon basin based on the compilation of OLIVEIRA et al. (OP. cif.), and new data obtained by us (Table 1). For the taxonomy and origin of t.he species with diploid/haploid numbers

Heu. Hydrobiol. irop. 2.5 (4) : .ZV-299 (1992).

already determined, several papers were consult.ed : GOLDSTEIN (1973), GÉRY (1977), NIJSSEN and Is- BRUCKER (1980), KULLANDER (1983, 1986), ORTEGA and VARI (1986), GOULDING et al. (1988), BURGESS (1989). The classification of the higher taxa of the listed species is arranged according to GREENWOOD et al. (1966) and LAUDER and LIEM (1983) with the exception of the Serrasalmidae which is considered at a family level, according to GÉRY (1977).

211’ nominal species belonging to t,he Amazonian ichthyofauna have had their diploid/haploid number determined. However, few of these species have had their karyotypes described. These listed species correspond to 49 o/. of the neotropical fish species listed by OLIVEIRA et al. (OP. cit.). We also det,ected that a vast amount of cytogenetic data is available in abstracts of Brazilian scientiflc meetings and others not readily available sources.

Diploid numbers

Diploid numbers (2n) of Amazonian fishes range widely : 2n = 22 (Nannostomus unifasciatus) to 2n = 134 (Corydoras aeneus). Also, species differ in having single or multiple nucleolar organizer regions (NORs) as well as different sex chromosomal mechanisms, including multiple systems. This karyotype diversity is apparently c.orrelated with the rich specific diversity of Amazonian ichthyofauna.

Karyologicaly, the characiforms are the most studied fish group of the Amazon, followed by siluriforms (including the gymnotoids - electric fishes), perciforms (cichlids) and then on a minor scale, osteoglossiforms (bonytongue fishes) and lepi- dosireniforms (lungfishes).

In the characiforms, the diploid numbers vary from 2n = 22 to 2n = 102. However, in the families Anostomidae, Curimatidae, Prochilodidae, Hemiodi- dae and Chilodidae the chromosome numbers and karyotype morphology are very similar, usually with 2n = 54, and met,a-submetacentric (M-SM) chromosomes in their karyotypes and single NORs. On the other hand, there are karyotypicaly divergent. families such as the Erythrinidae, Lebiasinidae, Characi- dae and Serrasalmidae each of which shows diversifi- cation in t.heir diploid numbers, karyotype morphologies and multiple NORs. Erythrinidae, Lebiasini- dae, Characidae plus CXenoluciidae are characterized by diploid numbers smaller than 2n = 54 (considered primitive in characiforms). Serrasalmidae, on the other hand, contains species with diploid numbers equal to or great,er than 2n = 54. This cyt.ogenetic feature of serrasalmids (2n = 54) has lead AREFJEV (1989) and PORTO et al. (1989, 1991) to consider them a distinct group at a family level, and not a

IURYOTYPES OF AMAZONIAN FRESHWATER FISHES 289

subfamily of Characidae, as postulated by some authors.

In the siluriforms, different from characiforms, there are few cytogenetic studies, although this group also has a large number of species in t,he Amazon basin. Diploid numbers of siluriforms vary from 2n=24 to 2n= 134. The available data indicate that they are characterized by high karyotypic diversity, especially the families Callychthyi- dae (Siluroidei) and Sternopygidae (Gymnotoidei). It should be pointed out t,hat the karyotypic diversity detected in Callichthyidae is relat.ed to gene duplica- tion and/or polyploid event.s (OLIVEIRA et al., op. cif.).

In the perciforms, especially the cichlids, the diploid numbers vary from 2n = 38 (n = 19) to 2n = 60. Almost a11 species present 2n=48, domina- ted by subtelo-acrocentic (ST-A) chromosomes, although there atie some different diploid numbers and karyotype morphologies in this group, for example 2n=60 predominating M-SM chromosomes.

Finally, in the osteoglossiforms, three species of osteoglossids were karyotyped and their diploid numbers vary from 2n=54 t.o 2n=56. In lepidosireniformes, a single spec.ies was karyotyped and a small diploid number (2n=38) was detected, with enormous chromosomes and an extremely high DNA value.

Nucleolar organizer region (NOR)

The NOR chromosome data from Amazonian freswater fish families presented in Table 1 makes it, possible to detect three groups in terms of patterns of specific localization of NORs sites : 1) single NORs, 2) multiple NORs, and 3) both single and multiple NORs. In the first group, t.he following families cari be listed : Osteoglossidae, Anostomidae, Curimatidae, Prochilodidae, Hemiodidae, Pimelodi- dae, Apteronotidae, St.ernopygidae and Cichlidae ; in the second group : Serrasalmidae and Lebiasinidae; and in the third group : Erhythrinidae, Characidae and Callichthyidae.

C-banding

Most of the species analyzed thus far present C- bands at, or around, the cent.romeres (centrolperi- centromeric band). Less frequently they occur at the chromosome tips (telomeric band). Positive C-bands cari also be found associated with NOR regions (in almost a11 of the serrasalmid species) or along chromosome arms (int,erstitial band) or as entirely heterochromatic short or long chromosome arms (particularly the sex chromosomes). Al1 types of C-

Rev. Hydrobiol. trop. 25 (4) : 287-299 (1992).

bands cari occur in the same species. Thus, when different species are compared, aimost always a species-speciflc pattern of constitutive heterochromatin distribution is apparent.

!3ex chromosomes

Based on cytological heteromorphy, heterologous sex chromosomes (ZZ/ZW and XX/XY1Y2 types) have been described for 6 Amazonian species, corresponding to 2.8 0/0 of the total number of species list.ed. The occurrence of t,he same kind of sex chromosome mechanisms (ZZ/ZW) in different genera (Triporfheus, Semaprochilodus and Eigenmannia) shows that this kind of heterogamety has evolved several times among the Amazonian Ostariophysi fishes. FALCAO (1988) suggests that in Triporfheus, the W chromosome differenciated through a hete- rochromatinization process, followed by deletions.

J3ibridization and Triplody

Artificial crossing has shown that serrasalmids species, Mylossoma durivenfris X Colossoma macropomum and Piaracfus brachypomus X Colossoma macropomum, have a capacity to hybridize even when the taxa belongs to distinct genera (KOSSOWSKI et al., 1983 and NAKAYAWA et al., pers. Comm., respective- lY)-

In the Amazon basin two cases of natural triploidy have been reported in fish : Eigenmannia SP. and Hopleryfhinus unifaeniafus. In both cases diploid and triploid forms were found. The fertilizat.ion of a non- reduced ovule by a haploid spermatozoon was considered to be the most probable origin of the triploids observed (ALMEIDA-TOLEDO et af., 1985; GIULIANO-CAETANO and BERTOLLO, 1990).

CONCLUSION

There still are too few cytogenetic data on Amazonian fishes. The lack of published karyograms or metaphase photographs as well as the problem of taxonomie determinat.ion and origin of analyzed material constitute the main problems in the cytogenetic study of Amazonian fishes, especially in the characiforms whose cytogenetic data forms a subs- tantial part of the checklist. Thus, these data must be considered with tare.

Considering the existence of approximately 3 000 species in the Amazon Basin and the high karyotypic diversity detected SO far, cyt,ogenetic studies of

290 J. 1. R. PORTO, E. FELDBERG, C. M. NAKAYAMA, J. N. FALCAO

Amazonian fishes should be carried out preferably on monophyletic groups of fishes, taking into account their biogeography and the utilization of high resolution chromosome banding. In this way, it Will be possible to determine the karyotypic. characters

ACKNOWLEDGEMENTS

We thank Dr. James PATTON for his comments on the manuscript and Dr. Michel JÉGU for his French translation. This research was supported by CNPq-Pig, Inpa/SCT, Eletro- nort.e and Orstom.

t,hat better elucidate the taxonomie and evolutionary problems. Manuscrit accepté par le Comité de rédacfion le 15 février 1993

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MOREIRA FILHO (0.). BERTOLLO (L. A. Cl.), FERRARI (1.). TAXAHASHI (C. S.) and FORESTI (F.), 1980. - Est.udos citogon&icos em peixes da regiao amazônica. III. Ordem Perciformrs. Ciência e Culfura (Supl.), 32 : 73-1.

TV~URAMOTO (J.), OHNO (S.) and ATKIN (M. B.), 1968. - On t.he diploid st.ate of the fish order Ostariophysi. Chromoso- ma, 24 : W-66.

NAK.~Y~%~A (c. M.), .JÉGu (111.) and FELDBERG (E.), 1986. - Cararteriza+o citotaxondmica de algumas espécies do gènero Serrasalmus (Serrasalmidae) da Amazônia. Res. ‘1111 Congresso Brasileiro de Zoologia : 335.

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NAKAYAM~ (CM.), FELDBERG (E.) and PORTO (J. 1. R.), 1988a. - Variabilidade de NORs e ntimero cromossômico no gènero Serrasalmus (Serrasalmidae, Characiformes). Res. .YF’ Congresso Brasileiro de Zoologia : 370.

NAKAYAMA (C. ;CI.), FELDBERG (E.) and PORTO (J. 1. R.), 1988 h. - Padrao de handa C em 4 espbcies do gênero Serrasnlmus (Characiformes, Serralmidae) de bacia Amazônica. Res. II Simp. Citog. Euol. e =Iplic. Peixes Neofrop. : 25.

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NIJSYEN (H.) and ISBRUCI~ER (1. J. H.), 1980. - A review of the genus Corydoras (Lacepède, 1803) (Pisces, Silurifor- mes, Callichthyidae), Bijdragen Tof de Dierkunde, 50 (1) : 190-220.

OHNO (S.) and ATICIN (N. B.), 1966. - Comparat.ive DNA values and chromosome comp1ement.s in eight, species of flshes. Chromosoma, 18 : 455-456.

OJIMA (Y.), UENO (L.) and HAYASHI (M.), 1976. -A review of t.he chromosome numbers in fishes. La Kromosomo II (1) : 19-47.

OLIVEIRA (C.), 1987. - Esfudos cifogenéficos no gênero Corydo- ras (Pisees, Siluriformes, Callichthyidae). Disserta@0 de Mestrado. Departamento de Biologia, Instituto de Biociéncias. Universidade de Sao Paulo. 154 p.

OLIVEIRA (C.), z4L~%EIDA-TOLEDo (L. F.), FORESTI (F.), BRITSKI (H.) and TOLEDO-FILH~ (S. A.), 1988a. - Chromosome formulae of Neot.ropical freshwater flshes. Reu. Brasil. Genet., 11 (3) : 577-624.

OLIVEIRA (f;.), ALMEIDA-TOLEDO (L. F.). FORESTI (F.) and TOLEDO-FILHO (S. A.), 1988 b. - Supernumerary chromosomes, Robert.sonian rearrangement and multiple NORs in Corydoras aeneus (Pisces, Siluriformes, Callich- thyidae). Caryologia, 41 (3-4) : 227-236.

OLIVEIRA (C.), ALMEIDA-TOLEDO (L. F.) and TOLEDO-FILHO (S. A.), 1990a. - Comparative cytogenet.ic analysis of t,hree cytotypes of Corydoras naftereri (Pisces, Silurifor- mes, Callichthyidae). Cyfologia, 55 : 21-26.

OLIVBIR.~ (C.), ALYEIDA-TOLEDO (L. F.) and TOLEDO-FILHO (S.A.), 199Ob. - Estudos citogenéticos na familia Callichthyidae (Pisces, Siluriformes) : os gèneros Diane- ma e Brochis. Res. III Simp. Cifog. Euol. e Aplic. Peixes Neofrop. : 27,

OLIVEIRA (M. 1. B.) AND FALCAO (J. N.), 1992. - Caracteriza- çao cariotipica em algumas espécies de peixes dos igarapés do municipio de Manaus. Res. I Congresso de Iniciaçdo Cienfifica do .-lmazonas : 99.

ORTEGA (H.) and Va~1 (R.), 1986. - Annotated checklist of t.he freshwater fishes of Peru. Smifhsonian Contributions fo Zoology, 437 : 25 p.

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PAULS (E.), 1985. - Considerac$es sobre a evoluçtio cromos- shmica e sistema de cromossomos supranumerarios em espécies do gènero Prochilodus (Pisces, Prochilodonti- dae). Tese de Doutorado. Departamento de Ciências Biologicas, Universidade Federal de Sao Carlos, Sao Paulo, Brasil. 156 p.

KARYOTYPES OF AMAZONIAN FRESHWATER FISHES 293

PAULS (E.) and BERTOLLO (L.A.C.), 1990. - Distribution of a super numerary chromosome system and aspects of karyotypic evolution in the genus Prochilodus (Pisces, Prochilodontidae). Genetica, 81 : 117-123.

Porto (J. 1. R.), 1992. - Estudos citotaxondmicos em peixes da familia Hemiodidae (Ostariophysi, Characiformes). da AmazOnia central. Dissertaçao de Mest,rado. Programa de Pas-Gradua@0 INPA-FUA. Manaus-AM. 103 p.

PORTO (J. 1. R.) and FELDBERG (E.), 1988. - Consideraçoes citotaxonômicas em Callichthys callichthys (Siluriformes, Callichthyidae) da Amazonia central. Res. II Simp. Ciiog. Eool. Aplic. Peixes Neotrop. : 8.

PORTO (J. 1. R.) and FELDBERG (E.), 1992a. - Ocorrência de novos citotipos nos gêneros Hoplosternum e Callichfhys (Siluriformes, Callichthyidae) da bacia amazonica. Reu. Brasil. Genet., 15 (1) Supplement 2 : 81.

PORTO (J. 1. R.) and FELDBERG (E.), 1992b. - Comparative cytogenet.ic study of the armored catfishes of the genus Hoplosternum (Siluriformes, Callicht.hyidae). Reo. Bra- si/. Genet., 15 (2) : 359-367.

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SCHEEL (J. J.), 1972. - The chromosomes of the third neon tetras. Trop. Fish Hobbyisf, 20 (4) : 60-65.

SCHEEL (J. J.), 1973. - Fish chromosome and their euolzztion. Interna1 reports of Danmarks Akvarium. Charlotten- lund, Danmark, 22 p.

SCHEEL (J. J.) and CHRIRTRNSEN (B.), 1970. - The chromosomes of thr two common neon t.etras. Trop. Fish Hobbyist, 19 (7) : 24-31.

SCHEEL (.J. J.), SIMONSEN (V.) and GYLDENHOLM (A. O.), 1972. - The karyotypes and some electrophoretic patterns of fourteen species of the genus Corydoras. Z. ZOO~. Sysf. Euol. Forsh., 10 : 144-152.

SOLA (L.), CATAUDELLA (S.) and CA~~NNA (E.), 1981. - New developmenm in vertebrate cytot.axonomy. III. Karyo- logy of bony lishes : a review. Genetica, 54 : 285-328.

SUZUKI (A.). TAKI (Y.) and URUSHIDO (T.), 1982. - Karyoty- pes of two species of Arowana, Osteoglossum bicirrhosum and 0. ferreirai. Japan. J. Ichfhyol., 29 (2) : 220-222.

THOM~SON (K. W.), 1979. - Cytotaxonomy of 41 species of Neotropical Cichlidae. Copeia, 4 : 679-691.

TURNER (B. J.), DIFFOOT (N.), RASCH (EM.), 1992. - The callichthyid catllsh Corydoraas aenezzs is an unresolved diploid-t.etraploid sibling species complex. Zchthyol. Explor. Freshzoaters, 3 (1) : 17-23.

URUSHIDO (T.), TAKAHASHI (E.) and TAKI (Y.), 1975. - Karyotypes of three species of lishes in the order Osteoglossiformes. Chromosome Information Seroice, 18 : 20-22.

UYENO (T.), 1973. - A comparat.ive study of chromosomes in the teleostean fish order Osteoglossiformes. In Suzuki A., Taki Y. and IJrushido T., 1982. - Karyot.ypes of two species of Arowana. Osteoglossum bicirhosum and 0. ferreirai. Japan. J. Ichthyol., 29 (2) : 220-222.

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VENERE (P. C.) and GALETTI Jr. (P. M.), 1985. - Natural triploidy and chromosome B in t,he fish Czzrimafa modesfa (Curimatidae, Characiformes). Reo. Brasil. Ge- nef., 7 (4) : 681-687.

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VENERE (P. C.) and GALETTI .Jr. (P. M.), 1986b. - Estudos cromossomicos em Leporinus (Anostomidae) de diferen- tes bacias hidrograflcas. Res. I Simp. Cifog. Evol. Aplic. Peixes Neofrop. : 62.

VENERE (P. C.) and GALETTI .Jr. (P. M.), 1989. - Chromosome relationships of somr Neotropical Characiformes of the family Curimatidae. Rro. Brasil. Genet., 12 (1) : 17-25.

Reo. Hydrohiol. trop. 25 (4) : 287-299 (1992).

ORDER

FAMILY

Species

OSTEOGLOSSIFORMES

TABLE I(1)

NOR CBA

n 2n KF numb pair type atm location f P T HSex reference

ARAPAIMIDAE Ampaima gigas

OSTEOGLOSSIDAE Osteoglossum bicirrhosum 0. bicirrhosum 0. bicirrhosum 0. ferreirai

CHARACIFORMES

ANOSTOMIDAE

Abramites hypselonotus Anostomus anostomus A. anostomus A. anostomus A. anostomus A. temstzi Leporinuç affinis L. brunneus L. cy/indfiformes 1. friderici L. ortomaculatus L. tigfinus Rhyticdus microlepis Schizodon fasciatus S. fasciatus Schizodon sp.

CHAFXIDAE BRYCONINAE

Brycon cf.cephalus B. cf. erythropterum B. cf. pesu

CHALCIDIINAE

Chalceus macrolepidotus C. macrolepidotus

CHEIRODONTINAE

C. troemneri Megalomphodus megalopterus M. megalopterus M. sweglesi Microschemobrycon casiquiare Paracheirodon axelrodi P. axelmdi 1 P. axelrodi P. innesi P. innesi P. innesi P. simulans 2 P. simulans Pristella riddlei P. riddlei

IGUANODECTINAE

Iguanodectes spilurus

STETHAPRIONINAE

Poptella compressa 3

TETRAGONOPTERINAE

Bfyconella palidifrons 4 Ctenobrycon aff. hauxwellianus Exodon paradoxus Hemigrammus anales H. erythrozonus

- 56

27 - - 56 - 56 - 54

lSM+lST+64A 3ST+53A

2M+4SM+14ST+34A

27 - 27 - 27 - 27 54 - 54

27 - - 54 - 54 - 54 - 54 - 54 - 54 - 64 - 54 - 54 - 54

El&!-SM 54M-SM

MM-SM 54MsM 54M-SM

32Md2SM

54M-SM 28M+26SM

54M-SM

- 50 20M+22SM&ST

- 50 20M+22SM+8ST

- 50 20M+22SM+8ST

- 54 32M+22ST - 52 44MSM+8ST-T

25 - 24 -

26 -

26 -

21 - 24 -

- 52 26 -

- 36 - 32

16 - 25 - - 50

24 - 26 -

2.5 -

25 -

25 - 25 - 26 - 27 - - 48

2

2

12

ST,A

M

M

ST

ST

ST

TEFIM

TERM.

TERM.

TERM.

TERM.

TERM.

Urushido et a/. (1975)

Hinegardner % Rosen (1972) uyeno (1973) Suzuki et a/. (1982) Suzuki et a/. (1982)

Scheel(lQ73) Hinegardner & Rosen (1972) Scheel(lQ73) Ojima et a/ (1976) Cestari et a/. (1990) Scheel (1973) Venere (1988) Venere & Galetti Jr.(1986b) Venere & Galetti Jr.(1986b) Galetti Jr. et a/. (1991) Venere & Galetti Jr.(1986b) Venere (1988) Bertollo et a/. (1980) Me&ner & Galeti Jr.(i 987) Galatti Jr. et a/. (1991) Venere & Galetti Jr. (1986a)

Santos et a/. (198.5)

Santos et a/. (1985) Feldberg et a/., unpubl.

Muramoto et a/. (1968) Ojima et a/. (1976)

Scheel (1973) Post (1965)

Scheel(l973)

Scheel (1973)

Scheel (1973) Post (1965)

Scheel & Christensen (1970) Scheel(l973)

Lueken & Foerster (1969) Scheel & Christensen (1970) Scheel (1973) Scheel (1973) Schael(i972) Post (1965) Schael (1973)

Scheel(lQ73)

Scheel (1973)

Scheel (1973) Scheel(l973) Scheel (1973) Schael(1973) Lueken & Foerster (1989)

TABLE 1 (2)

H. erythrozonus H. gracilis 5 H. hyanuary H. marginatus H. micropterus H. ocellifer H. ocellifer H. pulcher H. pulcher H. rhodostomus H. aff. rhodostomus H. aff. rodwayi H. aff. schmardae H. aff. schmardae H. stictus s H. stictus H. unilineatus H. vordetwinkleri Hyphessobrycon agulha H. aff. agulha H. belloti H. bentosi bentosi H. aff. copelandi H. erythmstigma 7 H. haraldschultzi H. heterorhabdus H. loretoensis H. peruvianus H. pulchripinnis H. pulchripinnis H. scholzei H. setpae H. serpae H. stictus H. stictus H. tropis Inpaichthys kerri Moenkhausia colle fi M. oligolepis Thayeria boehkei T. obliqua

26 - 24 48 - 52

25 - 26 - 24 48 19 - 24 - 25 - 25 - 25 - 25 - - 52

26 -

- 52 25 - 26 - 24 - 24 - 25 - 24 - 26 - 26 - 26 - 24 - 24 - 24 - 25 - 24 - 25 - - 50

24 - 26 -

- 52 25 - 23 - - 52

24 - 25 - 25 - 24 -

TRIPORTHEINAE

Triportheus albus T. cuiter T. elongatus T. flavus T. pictos

- 52

- 52 - 52 - 52

26 -

CHILODIDAE

Chilodus punctatus C. punctatus

27 - - 54

CTENOLUCIIDAE

Boulengerella sp - 36

CURIMATIDAE

Curimata cyprinoides C. knerl C. inornata C. ocellata C. vittata Cutimatella albuma C. meyeri Curimtopsis afl. macdepis (CYT

C. aff. macrolepis (CYT B) Cyphocharax cf. spilura Potamorhina altamazonica P. latior P. pristigaster Psectrogaster rutiloides Steindachnerina leuciscus

- 54 - 54 - 54 - 56 - 54 - 54 - 54

4 26 - 23 - - 54

51 102 - 56 - 54 - 54 - 54

ERYTHRINIDAE

Erythrinus erythrinus Hopleryhtrinus unitaeniatus

- 52 - 48

22M-SMt30ST-A

-

12Mt26SM+14ST-A

54M,SM

4lMtlOSM 4OMt12SM+2ST

4OM+14SM 4OMt16SM 42M+12SM 48M+8SM 46M+kSM

2M+2SM+Q8A 58M+2SM+2ST

42M+12SM 42M+12SM 48M+6SM

48MSM

14

l-4 i-4

2 2 2 2 1 2 2

2 2 2 2 2 2

, -

6,11 ST

331 ST 6,7 M,ST

3 M 15 ST 4 SM

22 SM 9 SM

14 M 9 M

10 5

27 16 11 16

M A M

!SA M M

M 1-8 - - _

SUB,TER

TERM. SUB,INT

TERM. TERM.

INTERST. INTERST.

TERM. TERM. TERM.

TERM. TERM. TERM. TERM. TERM. TERM.

TERM.

X X

X

X

X X

X X X

X X

X X X X

X

X X

X X X

X X

X X

X

Scheel(lQ73) Post (1965) Arefjev(l990b) Scheel(l973) Scheel (1973) Post (1965) Scheel (1973) Post (1965) Scheel (1973) Scheel (1973) S&el (1973) Scheel(l973) Scheel (1973) S&el (1973)

Soheel8 Christensen (1970) Scheel (1973) Scheel (1973) Scheel(l973) Scheel (1973) Scheel (1973) Scheel (1973) Scheel (1973) Scheel (1973) Scheel (1973) Post (1965) Scheel (1973) Scheel(lQ73) Scheel(lQ73) Post (1965) Scheel(l973) Arefjev (199Ob) Post (1965) Scheel (1973)

Gyldenholm & Scheel(lQ71) Scheel (1973) Scheel (1973) Arefjev (1989) Scheel(l973) Scheel (1973) Scheel (1973) Post (1965)

FalCao (1988)

FalCao (1988) FalcZio (1988) FalCao (1988) Scheel(l973)

Scheel (1973) Cestari et a/. (1990)

Porto et a/., unpubl.

Feldberg et a/. (1992) Felciberg et a/. (1992) Feldberg et a/. (1992) Feldberg et a/. (1992) Feldberg et a/. (1992) Felclberg et a/. (1992) Feldberg et a/. (1992) Scheel (1973) Scheel (1973) Venere & Galetti Jr.(1989) Fekfberg et a/. (in press) FeMberg et a/. (in press) Feklberg et a/. (in press) Feldberg et a/. (1992) Fekiberg et a/. (1992)

Bertdlo (1988) Giuliano-Caetano (1986)

TABLE 1 (3)

H. unitaeniatus (CYT A) H. unitaeniafus (CYT B) H. unitaeniafus (CYT C) H. unitaeniatus (CYT D) H. unitaeniatus (Tryploid) Hoplias malabaricus Hoplias sp.

Hoplias sp.

GASTEROPELECIDAE

Camegiella stfigata

C. strigata Gasteropslscus sternica

HEMIODIDAE ANODINAE

Anodus elongatus Anodus melanopogon Anodus sp.

BIVIBRANCHIINAE

Argonectes scapularis

HEMIODINAE

Hemiodus immaculatus H. cf. microlepis H. ocellatus H. unimaculatus

LEBIASINIDAE

Copeina guttata Copella amokli (CM A) Copella amokii FVr B) Copella nattereri

21 - 22 - 22 - 18 -

Nannostomus beckfordi (CYT A) 22 - N. bedrfordi (CYT B) N. beckfordi (CYT C) N. erythrurus N. marginatus N. tdfasciatus (CYT A) N. trifasciatus (CYT B) N. trifasciatus (CYT C) N. eques 8 N. harrisonig N. unifasciatus 10 Pyrhulina sp Pyrhulfna sp

18 - - 42

23 - 21 - 19 - 15 - 12 - - 34

20 - 11 - - 42 - 42

24 40 24 40 24 48

24 48 - 72 - 40

g 41

- 40 - 50

4EM-SM 46NSMt2ST-A 46NSMt2ST-A

47MSM+lST-A 69MSM+3ST-A

29MtllSMtlA (M)

30MclOSM (F)

B - 24 - 27 -

- 54 24M+26SM+4ST - 54 20M+28SM+6ST - 54 24M+24SM+6ST

- 54 50MSMt4ST

27 54 22M+26SM&ST - 54 20M+30SM+4ST

27 54 26M+24SMt4ST 27 54 26Mt24SMt4S.T

PRCCHILODIDAE P. nigricans Semaprochilodus insignis S. taeniurus

SERRASALMIDAE

Catoprion ment0 Colossoma macropomum 11 C. macropomum C. macropomum C. mawopomum C. maffopomum Metynnis argenteus M. lippincottianus M. schreitmuelleri 12 M. sp. Mylesinus paraschomburgkii Myleus pacu M. rubripinnis M. shomburgkii Mylossoma aureum M. duriventtis M. dutiven tris M. duriventris n.gen. 1 13

2M+4OA

34A

2M+2SMt38ST-A

- 54 4OM+14SM 27 54 4OM+14SM 27 54 4QM+14SM

- 60 27 - - 54 - 54 - 54 - 54

31 - - 62 - 62 - 62 - 58 - 58 - 58 - 58 - 54 - 54 - 54 - 54 - 58

18Mt36SM 20Mt34SM 26M+28SM

30M+30SM+2A ‘XIM-SMI~ST

64MSM+8ST-A

40M-SMt18ST-A

42M-SM+16ST-A 54M-SM

5OM-SMt4T 18M+34SM+2A

54M-SM 42M-SM+l GST-A

34

2

4 3

2 2 2

2

2 2 2 2

2-4

2 2 2

l-4 14

34 6-12 5-9

5-a 5-8

814

614 612

MA M

MA M-SM

26 ST 25 ST 25 ST

25

15 17

ST SM SM SM

2 3 3

M M M

M M

ST-A ST-A ST-A ST-A ST-A M,SM

M,SM ST-A

INTERST. TERJNT.

INTERST. TERM.

TERM. TERM. TERM.

TERNI.

TERM. TERM. SUBT. TERM.

INTERST. INTERST. INTERST.

TERM. TERJNT

INTERST. TERM. TERM. TERM. TERM. TERM.

TERM. TERM.

y2

xx

m

Giuliano-Caetano (1986) Giuliano-Caetano (1986) Giuliano-Caetano (1986)

Giuliano-Caetano (1986) Giuliano-Caetano (1986) Berldlo et a/ (1980) Bertdlo et a/ (1983)

Bertollo et a/. (1983) Bertollo & Moreira F” (1983)

Scheel (1973)

Scheel(1973) Scheel (1973)

Porto (1992) Porto (1992) Porto (1992)

Porto, unpubl.

Porto (1992) P&o (1992) Porto (1992) Porto (1992)

Scheel (1973) Scheel (1973) Scheel(l973) Scheel (1973) Scheel(l973) Scheel(l973) Arefjev (199Oa) Soheel (1973) Scheel(l973) Scheel(l973) Scheel (1973) Scheel (1973) Arefjev (199Oa) Scheel(l973) Scheel (1973) Oliveira & Fal&o (1992) Sangüino & FalCao (1992)

Pauls & berIoll0 (1990) Feldberg et a/. (1987) Feldberg et a/. (1987)

Porto et a/., unpubl. Scheel(l973) Berlollo et a/. (1980) Kossowski et a/. (1983) Almeida-Toledo et a/ (1987) Nakayama et a/. (1990) Scheel (1973) Arefjev (1989) Ojima et a/. (1976) Porto et a/. (1989;1991) Porto et a/. (1989; 1991) Porto et a/ (1989; 1991) Porto et a/ (1989; 1991) Porto et a/ (1989: 1991) Porto et a/. (1989; 1991) Ojima et a/. (1976) Kossowsld et a/ (1983) Porto et a/. (1969; 1991) Porto et a/ (196% 1991)

TABLE 1 (4)

3-6 - M,SM - _

e-10 _ ST-A 67 _ ST-A

6-10 - ST-A 8-8 - ST-A 68 - ST-A

- _ 612 _ - ST-A

_ - 810 _ ST-A

_ _ 4% _ ST,A

- - 610 - ST-A

Piaractus brachypomus - 54

Pristobrycon eigenmanni - 60 P. serrulatus - 60 P. stri0latu.s - 62 Pristobrycon sp. - 60

P. sp. - 60 Pygocentrus nattereri - 60 Serïasalmus affuvei - 60 S. elongatus - 60 S. hollandi - 64 S. manuelli - 60 S. rhombeus - 60 Serrasalmus sp.1 - 58 Serrasalmus 4x2 - 60 S. spilopleura - 60

SILURIFORMES SILUROIDEI

AGENEIOSIDAE

Ageneiosus brevifilis Ageneiosus sp.

- 56 - 54

AUCHENIPTERIDAE

Parauchenipierus d. galeatus - 58

CALLICHTHYIDAE

Brochis splendens 14 B. splendens Callichthys cakhthys C. callichthys C. callichthys Corydoras aeneus 15 C. aeneus Corydoms aeneus C. aeneus 16 C. aeneus C. aeneus C. aeneus C. agassizii C. agassizii C. arcuatus C. arcuatus C. elegans C. elegans C. elegans C. melanistus C. melanistus C. melanistus C. punctatus C. aff. punctatus C. rabauti C. rabauti C. reticulatus C. schwartzi C. schwartzi C. sch wartzi Dianema longibatbis D. longibarbis D. urostriatum 17 D. urostriatum H. littorale H. thoracatum H. ail. thoracatum H. sp.

49 - - 100 - 52

- 54 - 58 - 132

66 - 60 - - 58

29 - - 134 - 56 - 98

49 - - 46

23 - - 50

25 - 25 - - 46

23 - 24 - ;t _

- 102 - 58

29 -

- 74 - 46

23 - - 46 - 60 - 60 - 62 - 62 - 60 - 64 - 66 - 62

DORADIDAE

Opsodoras humeralis Pseudodoras niger

- 58 - 58

LORICARIIDAE

Pterygoplich thys mulfiradia tus Loricaria sp.

- 52’ - 62

34M+2OSM

44M-SMtl GST-A 44MSMt16ST-A 46M-SM+1 GST-A 48M-SM+1 PST-A

BOM-SMtlOST-A BOM-SMtlOST-A 46M-SMt14ST-A 4OM-SM+20ST-A

44M-SMtlGsT-A 44M-SMt16ST-A 46M-SM+12ST-A 46M-SM+14ST-A 44M-SMtWST-A

lEM+iESM+20ST+44A

46M-.SMt6ST-A

32Mcl4SM

8MSMt52ST-A 8M+4SMi&T+46A

6MSM+56ST-A 8MSMt52ST-A

30M-SM+34ST-A

8MSM+54ST

SBM-SMST-A

4 2

3

2

2-3

SM,ST q TERM. SM,ST P TERM.

SM,ST P TERM.

SM TERM.

A TERM.

M TERM.

.sM A

ST A A

INTERST. TERM. TERM. TERM. TERM

MSM TERM.

PJl

P P P

P P

P

P

P

P

TERM.

TERM. TERM. TERM.

TERM. TERM.

TERM

TERM.

TERM.

TERM.

x x x - x x x - - x x -

- _ _ _ x x x - _ - _ _ - x x - - - - _ - x x - - - - - x x x - - x x - - x x -

Nakayama et a/. (1990) Porto et a/. (1989; 1991) Porto et a/. (1989; 1991) Porto et a/. (1989; 1991) Nakayama et a/. (1968a)

Nakayama et a/. (1965a) Nakayama et a/. (1986a;b) Nakayama et a/., unpubl. Nakayamaetd(1986,1988b)

Muramoto et a/. (1968) Porlo et a/. (1991) Nakayama et a/. (1992) Nakayama et a/. (1992) Nakayama et a/. (198813). Porto et a/. ( 1991)

Fennocchio & EWlollo (1987) Fennocchio & Bertollo (1987)

Fennocchio & Berlollo (1987)

Scheel (1973) Oliveira et a/. (1990b) Porto & Feldberg (1992b)

Porto & Feldberg (1968) Porto & Feldberg (1992b) Scheel et a/. (1972) Scheel(l973) Hinegardner & Rosen (1972) Scheel et a/. (1972) Scheel (1973) Turner et a/. (1992) Turner et a/. (1992) Scheel et a/. (1972) Scheel(l973) Sd-ml et af. (1972) Scheel(l973) Scheel et a/. (1972) Scheel(l973) Hinegardner & Rosen (1972) Scheel et a/. (1972) Scheel (1973) Hinegardner & Rosen (1972) Hinegardner & Rosen (1972)

Oliveira (1987) Scheel et a/. (1972) Scheel(l973)

Oliveira (1987) Scheel et a/. (1972) Scheel(l973) Oliveira (1987) Hudson (1976) Marcon et a/. (1992) Oliveira et a/. (199Ob) Marccn etal. (1992) Porto & Feldberg (1992b) Porto & Feldberg (1992b) Porto & Feldberg (1992a) Porto & Feklberg (1992a)

Della-Rosa et a/. (1980) Venere (1968)

Della-Rosa et a/. (1980) Della-Rosa et a/. (1980)

PIMELODIDAE

Callophysus macrop&rus Pimelodus blochii Pseudoplatystoma fasciatum P. &winum Sorubim lima

TRICHOMYCTERIDAE

Vandelia cirrhosa

GYMNOTOIDEI

APTERONOTIDAE

Apteronotus albifrons A. albifrons A. albifrons

GYMNOTIDAE

Gymnotus carapo G. carapa G. carapo

HYPOPOMIDAE

Hypopomus brevirostris H. artedi Hypopigus lepturus

RHAMPHICHTHYDAE

Ramphichthys cf. pantherinus

STERNOPYGIDAE

Eigenmannia humboldti Eigenmannia sp. (Triploid) Eigenmannia sp. (Cm-l)

Eigenmannia sp. (CYT-2)

Eigenmannia sp. (Cm-l)

Efgenmannia sp. (CYT-1)

Eigenmannia sp. (Cm-l) Eigenmannia sp. (CYT-3) Eigenmannia sp. (CYT-B)

Sternopygus macrurus S. macrurus

PERCIFORMES

CICHLIDAE Acarichihys heckeli Apistograma agassizi A. agassizi Apistograma ortmanni A. orimanni A. pettensis 18 Astronotus ocellatus Cichla temensis Cichla sp. Crenicichla strigata C. lucius Dicrossus filamentosus 19 D. maculata 20 Geophagus altifrons 21 Guinacara sp. Heros sevems 22 H. severus H. severus Hypselacara coryphaenoides 23 Laetacara cutviceps24 Pterophyllum eimeksi P. scalare P. scalare Saianoperca jurupari 25 Symphysodon aequifasciafus 26

50 58 56 56 56

2 - ST-A p TERM - - - - Ftamirez-Gil& Feklberg, 1992 - - - - _ - Della-Rasa et a/. (1980)

2 - ST-A p - - _ _ - Fennocchii & EWtollo (1992) 2 - ST-A p _ - _ _ _ Fennocchio & Bertollo (1992) 2 - ST-A p _ _ _ _ _ Fennocchii & Bertollo (1992)

32 _ - - - _ - Scheel(l973)

11 24 24 2 4 SM q INTERST.

Hinegardner & Rosen (1972) Howell(l972) Almeida-Tokdo et a/. (1981)

19

42 32MSM+lOST-A 48 34M-SM+14ST-A

Foresti et a/. (1984) Foresti et a/. (1984) Hinegardner & Rosen (1972)

36 4M+2SM+8ST+22A 30 32MSM+6ST-A 50 16M+20SM+lOST+4A

Almeida-Toledo (1978) Oliveira ef a/. (1988a) Almeida-Toledo (1978)

52 38M+lOSM+4ST Almeida-Toledo (1978)

40 46 311

&

31/ El

3”o 28 38 38 46 46

8MSM+32ST-A 20M-SM+26ST-A

12/13M-SM+1&2OST-A

W8M-SM+22/24ST-A

1 lHPM-SM+1 WOST-A

1 PWSM+18/20ST-A

12M-SM+18ST-A 14M-SM+14ST-A

17M-SM+2lST-A (F) 16M-SM+22ST-A (M)

mA-SM

2 3 2

2

2

2

2 2 2 2 2

2

2 2

3 M-SM 4 M-SM 4 M-SM

11 ST-A

4 M-SM

4 M-SM

4 M-SM 3 ST-A 6 M-SM 6 M-SM 1 M-SM

48 6WSM+42ST-T 20

19

24 24

46 24MSM+22ST-T

46 24MSM+22ST-T

48 48 48 40 48 4s

12M-SM+36ST-A 48ST-T 48ST-T

6MSM+42ST-T

M-SM

12M-.SM+34ST-T 23 24 48 4MSM+44ST-A

48 4M-SM+44ST-A M-SM MSM

P P P

9

P

P

P q P P 9

P

P P

TERM. TERM. TERM.

INTERST.

TERM.

TERM.

TERM. TERM. TERM. TERM.

INTERST.

Foresti (1987) Almeida-Toledo et a/. (1985) Aimeida-Toledo et a/. (1985)

Foresti (1987)

Foresti (1987)

Foresti (1987)

Foresti (1987) Foresti (1987)

Foresti (1987) Foresti (1987) Bertdlo et a/. (1980)

INTERST.

INTERST. INTERST.

24 24

19 24 24

40 4MSM+44ST-T 48 6MSM+42ST-T

48 48 60

4MSM+44ST-T

44M+16A

X X X

‘

zvv 22

Thompson (1979) Scheel(i973) Thompson (1979) Scheel (1973) Thompsorl(l979) Post (1965) Feldberg B Bertdlo (1985ab) Thompsoil(l979) Venere (1988) Thompson (1979) Thompson (1979) Thompsom (1979) Scheel (1973) Feldberg & f3ertdlo (1985ab) Feldberg et a/. ( 1990) Post (1965) Scheel (1973) Thompson (1979) Thompson (1979) Scheel(l973) Post (1965) Scheel(l973) Thompson (1979) Moreira F” et a/. (198Oa) Ohno & Atkin (1966) Scheel(l973) S. aequifasciatus 30 -

TABLE 1 (5)

TABLE 1 (6)

S. aequifasciatus Uaru amphiacanthoides

LEPIDOSIRENIFORMES LEPIDCSIRENIDAE

Lepidosiren paradoxa L. paradoxa

- 60 - 46

- 38 - 38

5@Jm+2sT-T _ _ _ _ _ _ _ _ _ Thompson (1979) m$M+3&ST-T _ _ _ _ _ _ _ _ _ Tholllpsoil(l979)

38M - - - - - - Ohno & Atfdn (1966) 38wsM-sT _ _ - _ _ _ _ - _ Ofiieira et al. (1988a)

1 = Cheirodon axelrodi 2 = Hyphessobrycon simulans 5 = Hemigrammus gracilis 6 = Hemigrammus stictus 9 = P. harrfsoni 10 = P. unifasciatus 13 = Utiarftichthys sp 14 = Brochis coeruleus 17 = Dianema urostriata 21 = Geophagus surinamensis

18 = Apistograma pertense 22 = Cichlasoma severum

25 = Geophagus jurupari 26 = Symphysodon aequifasciata

3 = Poptella orbicularis 7 = Hyphessobrycon erythrostigma 11 = Piaractus nigripinnis 15 = Corydoras aneus 19 = Crenicara filamentosa 23 = Cichlasoma coryphaenoides

4 = Bryconella palifrons 8 = Poecilobrycon eques 12 = Metynnis hypsauchen 16 = Corydoras schuftzies 20 = Crenicara macufatus 24 = Aequidens curviceps

R~U. Hydrobiol. frop. 25 (Y) : 287-299 (1992).

a checklist of chromosome numbers and karyotypes of...

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