Os Biólogos:
Sven O. Kullander
- Department of Vertebrate Zoology, Swedish Museum of Natural History, POB 50007, SE-104 05 Stockholm, Sweden.
E-mail: sven.kullander@nrm.se


Efrem J. G. Ferreira - Centro de Pesquisas de Biologia Acuatica, INPA, CP 478, 69083-000 Manaus, AM, Brazil.- E-mail: efrem@inpa.gov.br

São os responsáveis pela publicação feita em dez-06 do periódico abaixo:

"Ichthyological Exploration of Freshwaters
An international journal for fi eld-orientated ichthyology
Volume 17 • Number 4 • December 2006
pages 289-398, 93 fi gs., 27 tabs.

ISSN 0936-9902 - Printed in Germany
Verlag Dr. Friedrich Pfeil, Wolfratshauser Str. 27, D-81379 München, Germany
Tel. + 49 - (0)89 - 74 28 27 0 – Fax + 49 - (0)89 - 72 42 77 2 – E-mail: info@pfeil-verlag.de – www.pfeil-verlag.de

A review of the South American cichlid genus Cichla,
with descriptions of nine new species
(Teleostei: Cichlidae)."

Trata-se da conclusão de exaustivo trabalho de coleta e pesquisa em campo e laboratório, sobre o nosso querido TUCUNARÉ.
Neste trabalho, os Biólogos consolidam a existência de 15 diferentes espécies de "TUCUNAS" (9 espécies novas) e apontam para a possibilidade de existirem pelo menos mais 30 outras não catalogadas.

Segue, parte deste maravilhoso trabalho; Para os apaixonados pelo majestoso peixe, resta adquirir o periódico conforme acima.

Por se tratar de um trabalho científico, só é permitida a tradução por tradutor juramentado.

Cichla, with the junior synonym Acharnes, is widely distributed in the Amazon, Tocantins, and Orinoco river basins, and in the smaller rivers draining the Guianas to the Atlantic Ocean. Within South America transplantations are recorded from the Paraná and Paraguay river drainages in Paraguay and Brazil, and the Paraíba do Sul and Paraguaçu rivers in Brazil. The genus comprises 15 species recognized by external characters of which colour pattern and meristics are most significant. In six species juveniles possess three dark blotches on the side and a dark band connecting the posterior blotch to the dark blotch at the caudal-fin base: Cichla ocellaris is known from the Guianas, including the Marowijne, Suriname, Corantijn, Demerara, and Essequibo river drainages, and also the upper Rio Branco in Brazil. Cichla orinocensis is known from the Negro and Orinoco river drainages in Brazil, Colombia, and Venezuela. Cichla monoculus is widespread in the floodplains of the Amazon basin, in Colombia, Peru, and Brazil, and also collected from rivers of Amapá in Brazil, and the lower Oyapock River on the border between Brazil and French Guiana. Cichla nigromaculata is known from the upper Rio Orinoco in Venezuela and, tentatively, the middle Rio Negro in Brazil. Cichla kelberi, new species, is restricted to the Tocantins river basin, but also found transplanted in the Paraná and Paraíba do Sul river drainages and reported from the Nordeste region of Brazil. Cichla pleiozona, new species, occurs in the Madre de Dios, Beni, and Guaporé river drainages in Bolivia and Brazil, and in the Rio Jamari in Brazil. A lectotype is fixed for Cychla toucounarai which is a synonym of Cichla monoculus. Juveniles and young of the remaining nine species, in addition to the three midlateral blotches, possess a dark horizontal band extending from the head to the dark blotch at the caudal-fin base: Cichla mirianae, new species, is restricted to the upper Tapajós river drainage, in the Juruena and Teles Pires rivers, and the upper Xingu river drainage in Brazil. Cichla melaniae, new species, is restricted to the lower Xingu river drainage in Brazil. Cichla piquiti, new species, is restricted to the Tocantins river basin, but transplanted in the Paraná river basin in Brazil and Paraguay. Cichla thyrorus, new species, occurs in the Rio Trombetas in Brazil, upstream from the Cachoeira Porteira. Cichla jariina, new species, occurs in the Rio Jari in Brazil, where it is so far recorded only from the region of the Santo Antonio rapids. Cichla pinima, new species, occurs in the lower parts of southern tributaries of the Rio Amazonas in Brazil (Tapajós, Curuá-Una, Xingu), and the lower Tocantins and Capim rivers. Tentatively identified specimens are recorded from the Amapá, Araguari, and Canumã rivers in Brazil. Cichla pinima occurs translocated in the Rio Paraguaçu in southeastern Brazil, and is reported as translocated from the northeast of Brazil. Cichla vazzoleri, new species, occurs in the Uatumã and lower Trombetas rivers in Brazil. Cichla temensis is known from the Negro and Orinoco river drainages in Brazil, Colombia, and Venezuela. It is also recorded from blackwater rivers along the Rio Solimões-Amazonas in Brazil (Tefé, Rio Puraquequara, Rio Uatumã, and Silves). Cichla intermedia is restricted to the Casiquiare and Orinoco river drainages in Venezuela.

A phylogenetic analysis suggests that C. intermedia, C. piquiti, and C. melaniae are successive basal species, whereas an unresolved group composed of C. jariina, C. pinima, C. temensis, C. thyrorus, and C. vazzoleri is the sister group of (C. mirianae,(C. orinocensis,((C. ocellaris, C. nigromaculata),(C. monoculus, C. kelberi, C. pleiozona))).

Nominal species of Cichla in chronological order of description.


combination author

year type locality (river basin)

current name

Cichla ocellaris

Schneider, 1801

East India, erroneous

Cichla ocellaris
Cichla temensis Humboldt, 1821 Negro River Cichla temensis
Cichla atabapensis  Humboldt, 1821 Orinoco River Cichla temensis
Cichla argus Valenciennes, 1821 Unknown Cichla orinocensis
Cichla orinocensis Humboldt, 1821 Orinoco and Negro Rivers Cichla orinocensis
Cichla monoculus Agassiz, 1831 Brazilian sea -Amazon basin Cichla monoculus
Cichla Tucunare Heckel, 1840 Branco River Cichla temensis
Cichla flavo-maculata Jardine, 1843 Negro River Cichla temensis
Cichla nigro-maculata Jardine, 1843 Negro River Cichla nigromaculata
Cichla trifasciata Jardine, 1843 Negro River Cichla temensis
Acharnes speciosus Müller & Troschel, 1849 Guyana coast, Essequibo River Cichla ocellaris
Cycla toucounarai Castelnau, 1855 Amazon basin Cichla monoculus
Cichla unitaeniatus Magalhães, 1931 Purus and Negro Rivers Cichla temensis
Cichla bilineatus Nakashima, 1941 Amazon River Cichla monoculus
Cichla intermedia Machado-Allison, 1971 Casiquiare River Cichla intermedia
Cichla kelberi new Tocantns River Cichla kelberi
Cichla pleiozona new Guaporé River Cichla pleiozona
Cichla mirianae new Tapajós Cichla mirianae
Cichla melaniae new Xingu Cichla melaniae
Cichla piquiti new Tocantins Cichla piquiti
Cichla thyrorus new Trombetas Cichla thyrorus
Cichla jariina new Jari Cichla jariina
Cichla pinima new Curuá-Una Cichla pinima
Cichla vazzoleri new Trombetas Cichla vazzoleri

Cichla Ocellaris

Geographical distribution:
French Guiana, Surinam, Guyana, and Brazil, in the Marowijne, Suriname, Saramacca, Nickerie, Corantijn, Berbice, Essequibo, and Branco drainages.
Keith et al. (2000: 65) plotted many records from
along most of the main channel of the Marowijne,
except the mouth and distant headwaters.




Local names:

Cichla Orinocensis

Geographical distribution:
In the Amazon basin along the Rio Negro from near San Carlos de Rio Negro and from the Rio Uaupés near Mitú downstream to Manaus, and also in the lower Rio Branco, the Rio Puraquequara slightly east of the Rio Negro, and the Rio Casiquiare. Widely distributed in the Orinoco basin including the Inírida, Atabapo, Guarrojo, Guaviare, Meta, Aguaro, and Caroní rivers. Most localities are referable to
blackwaters, as suggested by the restriction to Rio Negro and Rio Puraquequara in the Amazon basin, and most collecting sites in the Orinoco drainage are in blackwater rivers.

Cichla orinocensis can be distinguished from all other Cichla species by the adult colour pattern, which typically includes three prominent black, ocellated blotches on the side, but no additional blotches on the dorsum. The ocellated blotches develop gradually and directly from dark blotches along the midline of the side, and vary considerably in appearance in adults. Cichla orinocensis never shows black blotches posterior to the pectoral-fin base as in C. ocellaris,
C. monoculus, C. kelberi, or C. pleiozona.

Local names: Tucunaré, Pávon.

Cichla nigromaculata

Geographical distribution:
Upper Orinoco and Casiquiare tributaries and the middle Rio Negro.

Local names: Tucunaré

Cichla Monoculus

Geographical distribution:
Rio Solimões-Amazonas along the main channel and lower courses of tributaries; in Peru from Yarina Cocha
on the Ucayali north to the lower Napo in Peru and Ecuador; in Colombia at Leticia, in Brazil from Tabatinga to the Marajó island, including lower parts of the Tefé, Trombetas and Tapajós rivers. Also present in the Araguari and lower Oyapock rivers north of the Amazon. The species is probably much more widespread in the lowland
Amazon basin than our records show.

Live colouration:
Adults dull olivaceous or yellowish to golden or dark greenish with golden sheen on side, white ventrally. Narrow orange to yellow band from mouth angle to lower caudal fin base marking ventral extent of yellow/golden side. Caudal ocellus ringed with golden. Pelvic, anal, and lower half of caudal fin orange to dark red. Iris yellowish red. Vertical bars dark grey, not contrasted, or deep black,
concentrated to dorsal blotches in large specimens and much more contrasted in living than in preserved specimens. Juveniles olivaceous with faint brassy sheen, ventrally white.

Local names: Tucunaré

Chicla Kelberi (new)

Geographical distribution:
Rio Araguaia drainage
and the lower Rio Tocantins drainage  Introduced in reservoirs in Rio Grande do Norte, Minas Gerais and Ceará (Chellappa et al., 2003, as C. monoculus; Fontenele, 1948, as C. ocellaris), in the Rio Paraíba do Sul (State of Rio de
Janeiro), and the Rio Paraná.


Cichla kelberi has long been confused with C. monoculus (e.g., Fontenele, 1958, as C. ocellaris), which it resembles in shape and general colour pattern. We have not found any diagnostic character other than the light-spotted pelvic, anal and caudal fins, to separate the two species, but since this character state is unique in the genus, we are confident about species distinctness. The lateral scale count is within the higher range of C. monoculus, and below the range of C. pleiozona.

Local names: Tucunaré Amarelo(yellow).

Cichla pleiozona (new)

Geographical distribution:The Bolivian Amazon basin including the Rio  Madre de Dios, Beni, Mamoré, and Guaporé river drainages in Bolivia and Brazil, and in the Rio Jamari, which is a tributary of the Rio Madeira.

Named pleiozona for reason of the additional vertical bar on the caudal peduncle, in contrast to C. monoculus and C. kelberi, recognizing also the usually distinct occipital bar; from the Greek pleios, more, and zona, girdle. To be treated as a noun in apposition.

Almost all specimens of C. pleiozona possess a distinct dark vertical bar on the caudal peduncle, which is very rarely seen in C. monoculus, and C. kelberi. Out of 22 specimens of C. pleiozona, the fourth bar is missing only in the 109 mm specimen (NRM 19131) and one adult specimen (INPA 24069) from Lago Fortaleza. Three other specimens from Lago Fortaleza (INPA 24074) however, possess the fourth bar. There are no morphometric differences between C. monoculus, C. pleiozona, and C. kelberi.

Local names: Tucunaré.


Cichla mirianae (new)

Geographical distribution:
Known from the upper Rio Tapajós drainage, in the Juruena and Teles Pires rivers, and from the middle and upper Rio Xingu drainage, in the Fresco, Batovi, Culuene
and Suiá-Missu rivers..

Named for Mirian Leal-Carvalho, who participated in the collection of part of the type series.

Small specimens of C. mirianae display an indistinct continuous lateral band from snout to caudal peduncle (Fig. 37), and retain this band as adults displaying as irregular blackish blotches arranged in a narrow band connecting the three lateral ocellar blotches and continuing onto the caudal peduncle. This development of the colour pattern is similar to that of C. intermedia, but in
that species, the juvenile lateral band is wider and more prominent, and whitish spots covering body and head of C. mirianae are absent. Cichla intermedia lacks the prominent lateral ocelli of C. mirianae and has about 6 narrow vertical bars across the side. Adult C. mirianae are similar to C. orinocensis in possession of three prominent ocellated blotches along the side, but juvenile C. orinocensis do
not possess a continuous horizontal band along the side.

Local names: Tucunaré.

Cichla melaniae (new)

Geographical distribution:
Only three localities in the lower Rio Xingu drainage are verified by preserved material. The Balneário do Pedral is located on the Rio Xingu about 8 km from Altamira. The type locality, Cachoeira do Espelho is located 13 km south of Altamira. The third locality, Gorotire, is located on the lower course of the Rio Fresco.

Local names:

Cichla piquiti (new)

The species epithet piquiti is a Tupi-Guarani word meaning striped, and is here used as a noun in apposition.

Local names: Recorded as tucunaré by Costi et al. (1977) and as tucunaré azul by Kelber (1999).

Cichla piquiti is known as tucunaré azul among Brazilian recreational fishermen (Kelber, 1999), with reference to an overall light bluish impression of living specimens.

Cichla piquiti bears some resemblance to C. ocellaris and C. nigromaculata in the presence of five wide vertical bars on the side, but ocellar markings are absent from the side, and the vertical bars reach more ventrally on the side. Cichla piquiti can be distinguished from the sympatric C. kelberi by the absence of light spots on the anal fin, presence of bars 1a and 2a, absence of abdominal blotches, and absence of occipital bar. It may be confused at small sizes with C. pinima, which also occurs in the lower Rio Tocantins, but the two species appear to be only narrowly parapatric.


Geographical distribution:
Natural occurrences are recorded from the lower (Rio Itacaiunas, Marabá, Itupiranga), and upper Rio Tocantins (Rio Paranã), as well as in the Araguaia drainage (Cocalinho and Rio das Mortes).
Cichla piquiti is extensively stocked in reservoirs in the Paraná river drainage, and the States of Minas Gerais and São Paulo. Kelber (1999) lists tucunaré azul from the Itaipu, Porto Primavera, Jupiá, Três Irmãos, Ilha Solteria, São Simão, Jaguara, Estreito, Peixoto, Furnas, Avanhandava, and Promissão dams in the Rio Paraná drainage, the Tres Marias dam in the Rio São Francisco drainage, and the Serra da Mesa dam in the Rio Tocantins drainage, as well as from the Rio Piquiri in the upper Rio Paraguay drainage. The tucunaré azul corresponds to C. piquiti, according to photographs provided by D. Kelber. Photos of large adults were made available by D. Kelber from Tres Irmãos (Rio Tietê), collected in 1995, and Itumbiara (Rio Paranaíba). Preserved material is available from Embarcação and Bocaina, also in the Rio Paranaíba drainage.

Cichla thyrorus (new)

Geographical distribution:
Rio Trombetas near Cachoeira Porteira, only upstream of the falls.

The species epithet thyrorus is from the Greek noun thyroros, meaning doorkeeper or porter, in reference to the geographical distribution in the Cachoeira Porteira, the Portuguese word porteira meaning gate or entrance.

The Rio Trombetas apparently has a distinctive endemic fauna from the Cachoeira Porteira upstream. Kullander & Ferreira (2005) listed seven cichlid species that are known only from that region, and examples from other fish families.

They concluded that the lower rapids of the Trombetas may constitute a distribution barrier to some species, but at least three species are rheophilic. The lower rapids certainly are a barrier to upstream dispersal, as lowland Amazonian cichlid species are absent from the Rio Trombetas upstream from the Cachoeira Porteira, but present immediately below, e.g., Apistogramma pertensis listed by Kullander & Ferreira (2005). Among species of Cichla, this geographical separation is evident in C. thyrorus being present upstream and C. vazzoleri only in and downstream of the Cachoeira Porteira.

Local names: Tucunaré.

Cichla jariina (new)

Geographical distribution:
Known only from the Rio Jari, in the region of Cachoeira Itacari and Cachoeira Santo Antonio.




Named with reference to the type locality, the Rio Jari; an adjective with alternative endings -inus and -inum.

Local names: Tucunaré.

Cichla pinima (new)

Geographical distribution:
Lower Rio Tapajós, Rio Curuá-Una, lower Rio Xingu, lower Rio Tocantins, and Rio Capim; uncertain localities in southern Amapá and Rio Arapiuns, introduced in Rio Paraguaçu in eastern Brazil, and probably elsewhere in northeastern Brazil.




The species epithet pinima is a Tupi-Guarani adjective meaning spotted with white, and is here used as a noun in apposition.

Local names: We propose tucunaré pinima as Brazilian name for this species, selecting a name already in use.

Cichla vazzoleri (new)

Geographical distribution:
Cichla vazzoleri is recorded from the middle Rio Uatumã and
lower Rio Trombetas including but not above the Cachoeira Porteira, to the Oriximiná region at the mouth of the Rio Cuminá.




Named in honour of Gelso Vazzoler (1929-1987), collector of part of the type series, former head of the Departamento de Biologia Acuática, INPA, Manaus.

Cichla vazzoleri is most similar to C. pinima. Refer to comparison under that species. Jégu et al. (1989) described Mylesinus paraschomburgkii, a rheophilic characid from the Rio Trombetas (Cachoeira Vira-Mundo and upstream) and Rio Uatumã (Cachoeira do Miriti and upstream).

This species appears to be restricted to rapids, and thus presents different dispersal options than the Cichla species. Although the distribution pattern of M. paraschomburgkii is similar to that of C. vazzoleri, which is present in both the
Uatumã and the Trombetas, C. vazzoleri, although collected from rapids in both rivers, is present in the lower Rio Trombetas chiefly in lowland conditions.

We are unaware of other fish species restricted to the Uatumã and lower Trombetas rivers, but we also cannot conclude that this is the total distribution of C. vazzoleri.

Cichla temensis

Geographical distribution:
Restricted to blackwater rivers and their tributaries, recorded from many localities along the Rio Negro in Brazil and Venezuela, many localities in the Rio Orinoco drainage, a few localities only in the Rio Branco drainage. Along the Rio Solimões-Amazonas recorded from Tefé, Rio Puraquequara, lower Rio Uatumã, Rio Preto da Eva, and Lago Saracá, all black-water habitats. Reported by Lowe-McConnell (1969) from the Rio Branco drainage in Guyana.
An adult specimen from the Rio Jamari (INPA 3493) and a small specimen from Lago Genipapo (INPA uncat.) on the Rio Aripuanã are questionably referred to C. temensis, but regrettably there is nearly no other Cichla material available from the Brazilian portion of the Madeira drainage to permit an understanding of the distribution of C. temensis in this region.

Local names:
Sarabiana (Natterer, in Heckel, 1840: 413, Rio Negro); tucunaré (Natterer in Heckel, 1840: 409, Boa Vista area); pintado (Schomburgk in Jardine, 1843:145, Rio Negro); Lucanari Grande (Schomburgk, in Jardine, 1843: 151, Rio
Negro); tucunaré sorubiana (Magalhães, 1931: 225, errata, Rio Negro); pavón pintado, pavón trucha, pavón venado (Román, 1981: 76, Venezuela); Pavón cinchado (large specimens; Román, 1981: 82).

Both the appelation pintado and sorubiana refer to the pimelodid catfish species Pseudoplatystoma fasciatum, known as surubi (Tupi-Guarani) or bagre pintado (Spanish).

Cichla temensis is the most elongate species in the genus, manifest above all at larger sizes, where it overlaps with C. vazzoleri but is more slender than C. pinima and C. thyrorus. The colour pattern and the ontogenetic development of principal dark markings, distinguish it from
all other species of Cichla.

Cichla intermedia

Geographical distribution:
Specimens at hand are from the Rio Cinaruco in Apure state, and from the Rio Casiquiare (Rio Cunucuna, Rio Cataniapo), reflecting the fragmented range given by Machado-Allison (1971: fig. 13, also including the Rio Siapa drainage) (Fig. 9). Winemiller (2001: fig. 2) also included the lower Rio Caura in the distribution. There are no records from Colombia, although the species is probably
present also in that country.




Local names: Pavón real.

Species diversity. Despite being one of the most colourful, well-known, and most commercial of the genera of cichlids, of importance to artisanal as well as tourism based sport fishing, and among the largest fishes in the Neotropics, the taxonomic confusion in Cichla has remained considerable.
A major source of confusion has been the obvious acceptance in species of Cichla of much more than modest variability in colour pattern, both ontogenetic and between individuals, as well as a traditional recognition of just a few species. This has been possible because most authors have
only dealt with very few specimens or a restricted geographical region. In this paper, covering a wider geographic area, we have been able to organise phenotypic variation within the genus into spatially clustered units representing 15 distinct phylogenetic species.

The species diversity of Cichla is thus demonstrated to be much greater than indicated by current literature. Nevertheless, we believe the work on a revision of Cichla has just started. Our species accounts include references to samples that cannot be satisfactorily assigned, and some
species are still known only from a few specimens. Huge areas within the geographical range of the genus have not been sampled for fishes. We expect that there may likely exist between 20 and 30 species of Cichla, which require large series of specimens and more extensive sampling to be located and diagnosed. Hopefully, this revision can inspire more efforts in further revising the genus.