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A checklist of chromosome numbers and
karyotypes of Amazonian freshwater jishes
Jorge Ivan
Rebelo PORTO (l), Eliana FELDBERG
Céleste Mutuko NAKAYAMA (l),
José das Neves FALCAO (2)
(1),
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 i ff 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.
determinandose
por 10 menos
a 5 ordenes ya fueron estudiados citogeneticamente,
haploideldiploide.
La variaci0n del numero diploide va desde 2n=22 hasfa 2n=134.
mecanismos cromo&micos
sexuales y polimorfismos
cromoscimicos en algunas especies,
siempre especie-specificos de la banda-C y de las regiones organizadoras
del nucleolo.
PALABRAS ~LAVES : Cromosomos -
(1) Instituto National
CEP 69083.
(2) Unioersidade
CEP 69000.
Peces -
da Pesquisas da Amazônia,
Agua dulce -
211 especies pertenecienfes
su numero cromosomico
Tanbien fueron observado
seguido de padrones casi
Cuenca amazonica.
Coordenaçüo de Pesquisas em Biologia
Aqudtica,
Cx Postal 478, Manaus-AM,
Brasil,
do Amazonas, Instituto
Rerr. Hydrobiol. trop. 25 (4) : 2X7-299(1992).
de Biociências,
Deparfamenio
de Biologia,
Estrada do Contorno S/N, Manaus-AM,
Brasil,
288
J. 1. R. PORTO, E. FELDBERG, C. M. NAKAYAMA,
INTRODUCTION
In the neotropics, chromosomal data have contributed 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), detection of inter and intraspecific polymorphism (GruLIANO-CAETANO and BERTOLLO, 1988; OLIVEIRA et
al., 199Oa), data on sex chromosome systems (MoREIRA 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; GIULIANOCAETANO and BERTOLLO, 1990). Al1 of t.he above
mentioned findings are important for the understanding 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 understanding of the diversit,y of fish species.
CYTOGENETIC
FISHES
FEATURES
OF AMAZONIAN
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 dealers.
In elaborat.ing our list, we attempted to identify,
among the species of unknown origin, the Amazonian freshwater species, and we updated the taxonomit 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).
J. N. FALCAO
already determined, several papers were consult.ed :
GOLDSTEIN (1973), GÉRY (1977), NIJSSEN and IsBRUCKER (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
unifasciatus)
to
widely : 2n = 22 (Nannostomus
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 lepidosireniforms (lungfishes).
In the characiforms, the diploid numbers vary
from 2n = 22 to 2n = 102. However, in the families
Anostomidae, Curimatidae, Prochilodidae, Hemiodidae 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, Characidae and Serrasalmidae each of which shows diversification in t.heir diploid numbers, karyotype morphologies and multiple NORs. Erythrinidae,
Lebiasinidae, 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
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 Callychthyidae (Siluroidei) and Sternopygidae (Gymnotoidei). It
should be pointed out t,hat the karyotypic diversity
detected in Callichthyidae is relat.ed to gene duplication 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 a11species present 2n=48, dominated 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, Pimelodidae, 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 Cbands at, or around, the cent.romeres (centrolpericentromeric 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 CRev. Hydrobiol.
trop. 25 (4) : 287-299 (1992).
289
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 geneSemaprochilodus
and Eigenmannia)
ra (Triporfheus,
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 heterochromatinization
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., respectivelY)In the Amazon basin two cases of natural triploidy
SP. and
have been reported in fish : Eigenmannia
Hopleryfhinus
unifaeniafus.
In both cases diploid and
triploid forms were found. The fertilizat.ion of a nonreduced ovule by a haploid spermatozoon was
considered to be the most probable origin of the
et af., 1985;
triploids observed (ALMEIDA-TOLEDO
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 substantial 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,
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
t,hat better elucidate the taxonomie and evolutionary problems.
J. N. FALCAO
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, Eletronort.e and Orstom.
Manuscrit
accepté par le Comité de rédacfion
le 15 février
1993
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Cytotaxonomy
of 41 species of
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URUSHIDO
PORTO (J. 1. R.), FELDBERG
POST
293
TABLE
ORDER
FAMILY
Species
I(1)
NOR
CBA
n
2n
-
56
27
-
56
56
54
27
27
27
27
27
-
54
54
54
54
54
54
54
54
64
54
54
54
28M+26SM
54M-SM
-
50
50
50
20M+22SM&ST
20M+22SM+8ST
20M+22SM+8ST
-
54
52
32M+22ST
44MSM+8ST-T
25
24
26
-
26
-
Scheel(l973)
Scheel (1973)
21
24
-
Scheel (1973)
Post (1965)
26
16
25
24
26
52
36
32
50
-
2.5
-
Scheel(lQ73)
25
-
Scheel (1973)
25
25
26
27
-
48
KF
numb
pair
type
atm
location
f
P
T
HSex
reference
OSTEOGLOSSIFORMES
ARAPAIMIDAE
Ampaima gigas
Urushido
et a/.
(1975)
OSTEOGLOSSIDAE
Osteoglossum bicirrhosum
0. bicirrhosum
0. bicirrhosum
0. ferreirai
lSM+lST+64A
3ST+53A
2M+4SM+14ST+34A
2
ST,A
TEFIM
Hinegardner % Rosen (1972)
uyeno (1973)
Suzuki et a/. (1982)
Suzuki et a/. (1982)
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.
El&!-SM
54M-SM
MM-SM
54MsM
54M-SM
32Md2SM
2
M
TERM.
12
M
TERM.
ST
ST
ST
TERM.
TERM.
TERM.
54M-SM
Scheel(lQ73)
Hinegardner & Rosen (1972)
Scheel(lQ73)
Ojima et a/ (1976)
Cestari et a/. (1990)
Scheel (1973)
Venere (1988)
Venere & Galetti Jr.(1986b)
Galetti Jr. et a/. (1991)
Venere (1988)
Bertollo et a/. (1980)
Me&ner
& Galeti Jr.(i 987)
Galatti Jr. et a/. (1991)
Venere & Galetti Jr. (1986a)
CHAFXIDAE
BRYCONINAE
Brycon cf.cephalus
B. cf. erythropterum
B. cf. pesu
Santos et a/. (198.5)
Santos et a/. (1985)
Feldberg et a/., unpubl.
CHALCIDIINAE
Chalceus macrolepidotus
C. macrolepidotus
Muramoto et a/. (1968)
Ojima et a/. (1976)
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
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)
IGUANODECTINAE
Iguanodectes spilurus
STETHAPRIONINAE
Poptella compressa 3
TETRAGONOPTERINAE
Bfyconella palidifrons 4
Ctenobrycon aff. hauxwellianus
Exodon paradoxus
Hemigrammus anales
H. erythrozonus
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
25
26
24
19
24
25
25
25
25
26
25
26
24
24
25
24
26
26
26
24
24
24
25
24
25
24
26
25
23
24
25
25
24
48
52
48
52
52
50
52
52
-
26
52
52
52
52
-
27
-
54
-
36
26
23
51
-
54
54
54
56
54
54
54
54
102
56
54
54
54
-
52
48
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
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)
22M-SMt30ST-A
-
Gyldenholm & Scheel(lQ71)
Scheel (1973)
Scheel (1973)
Arefjev (1989)
Scheel(l973)
Scheel (1973)
Scheel (1973)
Post (1965)
12Mt26SM+14ST-A
,
(1970)
-
TRIPORTHEINAE
Triportheus albus
T. cuiter
T. elongatus
T. flavus
T. pictos
14
6,11
ST
SUB,TER
X
X
l-4
i-4
331
ST
M,ST
TERM.
SUB,INT
X
X
X
X
6,7
FalCao (1988)
FalCao (1988)
FalcZio (1988)
FalCao (1988)
Scheel(l973)
CHILODIDAE
Chilodus punctatus
C. punctatus
Scheel (1973)
Cestari et a/. (1990)
54M,SM
CTENOLUCIIDAE
Boulengerella sp
Porto et a/., unpubl.
CURIMATIDAE
Curimata cyprinoides
C. knerl
C. inornata
C. ocellata
C. vittata
Cutimatella albuma
C. meyeri
Curimtopsis afl. macdepis (CYT 4
C. aff. macrolepis (CYT B)
Cyphocharax cf. spilura
Potamorhina altamazonica
P. latior
P. pristigaster
Psectrogaster rutiloides
Steindachnerina leuciscus
4lMtlOSM
4OMt12SM+2ST
4OM+14SM
4OMt16SM
42M+12SM
48M+8SM
46M+kSM
2
2
2
2
1
2
2
3
15
4
22
9
14
9
M
ST
SM
SM
SM
M
M
TERM.
TERM.
INTERST.
INTERST.
TERM.
TERM.
TERM.
2M+2SM+Q8A
58M+2SM+2ST
42M+12SM
42M+12SM
48M+6SM
2
2
2
2
2
2
10
5
27
16
11
16
M
A
M
!SA
M
M
TERM.
TERM.
TERM.
TERM.
TERM.
TERM.
1-8
-
_
M
TERM.
48MSM
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Feldberg et a/. (1992)
Felciberg et a/. (1992)
Felclberg 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)
Fekiberg et a/. (1992)
ERYTHRINIDAE
Erythrinus erythrinus
Hopleryhtrinus unitaeniatus
Bertdlo (1988)
Giuliano-Caetano
(1986)
TABLE
H. unitaeniatus
H. unitaeniafus
(CYT A)
(CYT B)
H. unitaeniafus (CYT C)
H. unitaeniatus (CYT D)
H. unitaeniatus (Tryploid)
Hoplias malabaricus
Hoplias sp.
Hoplias sp.
1
(3)
4EM-SM
24
24
24
40
40
48
24
g
48
72
40
41
-
40
50
B
24
27
-
-
54
54
54
24M+26SM+4ST
20M+28SM+6ST
24M+24SM+6ST
2
2
2
-
54
50MSMt4ST
2
27
27
27
54
54
54
54
22M+26SM&ST
20M+30SM+4ST
26M+24SMt4ST
26Mt24SMt4S.T
2
2
2
2
21
22
22
18
22
18
23
21
19
15
12
20
11
-
42
34
42
42
27
27
54
54
54
27
31
-
60
54
54
54
54
62
62
62
58
58
58
58
54
54
54
54
58
34
2
46NSMt2ST-A
46NSMt2ST-A
47MSM+lST-A
69MSM+3ST-A
4
3
29MtllSMtlA
30MclOSM
MA
M
MA
M-SM
INTERST.
TERJNT.
INTERST.
TERM.
y2
(M)
xx
(F)
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)
GASTEROPELECIDAE
Camegiella stfigata
C. strigata
Gasteropslscus sternica
Scheel (1973)
Scheel(1973)
Scheel (1973)
HEMIODIDAE
ANODINAE
Anodus elongatus
Anodus melanopogon
Anodus sp.
26
25
25
ST
ST
ST
TERM.
TERM.
TERM.
Porto (1992)
Porto (1992)
Porto (1992)
TERNI.
Porto, unpubl.
TERM.
TERM.
SUBT.
TERM.
Porto
P&o
Porto
Porto
BIVIBRANCHIINAE
Argonectes scapularis
HEMIODINAE
Hemiodus immaculatus
H. cf. microlepis
H. ocellatus
H. unimaculatus
25
15
17
ST
SM
SM
SM
(1992)
(1992)
(1992)
(1992)
LEBIASINIDAE
Copeina guttata
Copella amokli (CM A)
Copella amokii FVr B)
Copella nattereri
Nannostomus beckfordi (CYT A)
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
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)
2M+4OA
34A
2M+2SMt38ST-A
2-4
PRCCHILODIDAE
P. nigricans
Semaprochilodus insignis
S. taeniurus
4OM+14SM
4OM+14SM
4QM+14SM
2
2
2
2
3
3
M
M
M
INTERST.
INTERST.
INTERST.
m
Pauls & berIoll0 (1990)
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
18Mt36SM
20Mt34SM
26M+28SM
30M+30SM+2A
‘XIM-SMI~ST
64MSM+8ST-A
40M-SMt18ST-A
l-4
14
34
6-12
TERM.
TERJNT
814
ST-A
ST-A
ST-A
ST-A
ST-A
M,SM
INTERST.
TERM.
TERM.
TERM.
TERM.
TERM.
614
612
M,SM
ST-A
TERM.
TERM.
5-9
5-a
42M-SM+16ST-A
54M-SM
5OM-SMt4T
18M+34SM+2A
54M-SM
42M-SM+l GST-A
M
M
5-8
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)
Piaractus brachypomus
Pristobrycon eigenmanni
P. serrulatus
P. stri0latu.s
Pristobrycon sp.
P. sp.
Pygocentrus nattereri
Serïasalmus affuvei
S. elongatus
S. hollandi
S. manuelli
S. rhombeus
Serrasalmus sp.1
Serrasalmus 4x2
S. spilopleura
TABLE
1 (4)
3-6
-
M,SM
PJl
TERM.
_
_
_
_
_
ST-A
ST-A
ST-A
ST-A
ST-A
P
P
P
P
P
TERM.
TERM.
TERM.
BOM-SMtlOST-A
BOM-SMtlOST-A
46M-SMt14ST-A
4OM-SM+20ST-A
e-10
67
6-10
8-8
68
612
ST-A
P
TERM
44M-SMtlGsT-A
44M-SMt16ST-A
46M-SM+12ST-A
46M-SM+14ST-A
44M-SMtWST-A
_
810
_
4%
610
_
_
_
-
ST-A
P
TERM.
ST,A
P
TERM.
ST-A
P
TERM.
34M+2OSM
44M-SMtl GST-A
44MSMt16ST-A
46M-SM+1 GST-A
48M-SM+1 PST-A
x
x
-
x
x
x
x
x
x
-
x
_
x
-
_
x
x
x
x
x
x
_
x
_
x
x
x
x
x
_
_
_
-
Porto et a/. (1989; 1991)
Porto et a/. (1989; 1991)
Porto et a/. (1989; 1991)
Nakayama et a/. (1968a)
-
54
60
60
62
60
60
60
60
60
64
60
60
58
60
60
-
56
54
Fennocchio
Fennocchio
& EWlollo (1987)
& Bertollo (1987)
-
58
Fennocchio
& Berlollo (1987)
49
66
60
29
49
23
25
25
23
24
;t
100
52
54
58
132
58
134
56
98
46
50
46
_
29
23
-
102
58
74
46
46
60
60
62
62
60
64
66
62
-
58
58
-
52’
62
TERM.
TERM.
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/. (198813).
Porto et a/. ( 1991)
SILURIFORMES
SILUROIDEI
AGENEIOSIDAE
Ageneiosus brevifilis
Ageneiosus sp.
AUCHENIPTERIDAE
Parauchenipierus d. galeatus
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.
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
4
2
3
SM,ST
SM,ST
2
2-3
SM,ST
q
P
P
TERM.
TERM.
Scheel (1973)
Oliveira et a/. (1990b)
Porto & Feldberg (1992b)
TERM.
Porto & Feldberg (1968)
Scheel et a/. (1972)
Scheel(l973)
Scheel (1973)
Turner et a/. (1992)
Turner et a/. (1992)
Scheel(l973)
Sd-ml et af. (1972)
Scheel(l973)
Scheel(l973)
Scheel (1973)
SM
TERM.
A
TERM.
M
TERM.
Oliveira (1987)
Scheel(l973)
Oliveira (1987)
Scheel(l973)
Oliveira (1987)
Hudson (1976)
Marcon et a/. (1992)
Oliveira et a/. (199Ob)
Marccn etal. (1992)
Porto & Feldberg (1992a)
Porto & Feklberg (1992a)
.sM
A
ST
A
A
INTERST.
TERM.
TERM.
TERM.
TERM
MSM
TERM.
DORADIDAE
Opsodoras humeralis
Pseudodoras niger
SBM-SMST-A
Della-Rosa et a/. (1980)
Venere (1968)
LORICARIIDAE
Pterygoplich thys mulfiradia tus
Loricaria sp.
Della-Rosa
Della-Rosa
et a/. (1980)
et a/. (1980)
TABLE
1 (5)
PIMELODIDAE
50
Callophysus macrop&rus
Pimelodus blochii
Pseudoplatystoma fasciatum
P. &winum
Sorubim lima
58
56
56
56
2
2
2
2
-
_
-
ST-A
ST-A
ST-A
ST-A
p
p
p
p
TERM
_
_
-
-
_
-
_
_
_
_
_
_
_
_
_
Ftamirez-Gil&
Della-Rasa
Fennocchii
Fennocchio
Fennocchii
-
-
_
-
Scheel(l973)
Feklberg,
et
&
&
&
1992
a/. (1980)
EWtollo (1992)
Bertollo (1992)
Bertollo (1992)
TRICHOMYCTERIDAE
Vandelia cirrhosa
32
GYMNOTOIDEI
APTERONOTIDAE
Apteronotus albifrons
A. albifrons
A. albifrons
11
24
24
2
4
SM
q
Howell(l972)
Almeida-Tokdo
et a/. (1981)
INTERST.
GYMNOTIDAE
Gymnotus carapo
G. carapa
G. carapo
42
48
32MSM+lOST-A
34M-SM+14ST-A
36
30
50
4M+2SM+8ST+22A
32MSM+6ST-A
16M+20SM+lOST+4A
52
38M+lOSM+4ST
Eigenmannia humboldti
Eigenmannia sp. (Triploid)
Eigenmannia sp. (Cm-l)
40
46
311
8MSM+32ST-A
20M-SM+26ST-A
12/13M-SM+1&2OST-A
2
3
2
3
4
4
M-SM
M-SM
M-SM
P
P
P
TERM.
TERM.
TERM.
Eigenmannia sp. (CYT-2)
&
W8M-SM+22/24ST-A
2
11
ST-A
9
INTERST.
31/
El
1lHPM-SM+1 WOST-A
2
4
M-SM
P
TERM.
1 PWSM+18/20ST-A
2
4
M-SM
P
TERM.
3”o
12M-SM+18ST-A
14M-SM+14ST-A
17M-SM+2lST-A
(F)
16M-SM+22ST-A
(M)
mA-SM
2
2
2
2
2
4
3
6
6
1
M-SM
ST-A
M-SM
M-SM
M-SM
P
q
P
P
9
TERM.
TERM.
TERM.
TERM.
INTERST.
Foresti et a/. (1984)
Foresti et a/. (1984)
19
HYPOPOMIDAE
Hypopomus brevirostris
H. artedi
Hypopigus lepturus
Almeida-Toledo
(1978)
Oliveira ef a/. (1988a)
Almeida-Toledo
(1978)
RHAMPHICHTHYDAE
Ramphichthys cf. pantherinus
Almeida-Toledo
(1978)
Foresti (1987)
Almeida-Toledo
Aimeida-Toledo
et a/. (1985)
et a/. (1985)
STERNOPYGIDAE
Efgenmannia sp. (CYT-1)
Eigenmannia sp. (CYT-3)
Eigenmannia sp. (CYT-B)
28
38
38
46
46
Sternopygus macrurus
S. macrurus
Foresti (1987)
‘
Foresti (1987)
Foresti (1987)
X
X
X
Foresti (1987)
Foresti (1987)
zvv
22
Foresti (1987)
Foresti (1987)
Bertdlo et a/. (1980)
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
S. aequifasciatus
48
6WSM+42ST-T
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
48
48
4MSM+44ST-A
4M-SM+44ST-A
40
48
4MSM+44ST-T
6MSM+42ST-T
48
48
60
-
4MSM+44ST-T
20
19
24
24
23
24
M-SM
P
INTERST.
2
2
M-SM
MSM
P
P
INTERST.
INTERST.
12M-.SM+34ST-T
24
24
19
24
24
30
2
44M+16A
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)
TABLE
S. aequifasciatus
Uaru amphiacanthoides
1
(6)
-
60
46
5@Jm+2sT-T
m$M+3&ST-T
_
_
_
_
_
_
_
_
_
_
_
_
_
_
_
_
_
_
Thompson (1979)
Tholllpsoil(l979)
-
38
38
38M
38wsM-sT
_
_
-
_
_
_
_
-
_
Ohno & Atfdn (1966)
Ofiieira et al. (1988a)
LEPIDOSIRENIFORMES
LEPIDCSIRENIDAE
Lepidosiren paradoxa
L. paradoxa
1 = Cheirodon axelrodi
5 = Hemigrammus
gracilis
9 = P. harrfsoni
13 = Utiarftichthys
sp
17 = Dianema urostriata
21 = Geophagus surinamensis
25 = Geophagus jurupari
R~U. Hydrobiol.
2 = Hyphessobrycon
simulans
6 = Hemigrammus
stictus
10 = P. unifasciatus
14 = Brochis coeruleus
18 = Apistograma
pertense
22 = Cichlasoma
severum
26 = Symphysodon
aequifasciata
frop. 25 (Y) : 287-299 (1992).
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

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