See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/262707033
Molecular diagnosis of the arowanas
Osteoglossum ferreirai Kanazawa, 1966 and O.
bicirrhossum (Cuvier, 1829) from the...
Article in Neotropical Ichthyology · June 2013
DOI: 10.1590/S1679-62252013000200011
CITATION
READS
1
45
5 authors, including:
Maria Doris Escobar Lizarazo
Donald Charles Taphorn
9 PUBLICATIONS 11 CITATIONS
202 PUBLICATIONS 1,496 CITATIONS
Federal University of Amazonas
SEE PROFILE
Royal Ontario Museum
SEE PROFILE
Miguel Angel Landines
Tomas Hrbek
16 PUBLICATIONS 71 CITATIONS
321 PUBLICATIONS 1,603 CITATIONS
National University of Colombia
SEE PROFILE
Federal University of Amazonas
SEE PROFILE
Some of the authors of this publication are also working on these related projects:
MANEJO COMUNITARIO DE QUELÔNIOS - PROJETO PÉ-DE-PINCHA View project
Guide to the Freshwater Fishes of Guyana View project
All content following this page was uploaded by Miguel Angel Landines on 22 July 2014.
The user has requested enhancement of the downloaded file. All in-text references underlined in blue are added to the original document
and are linked to publications on ResearchGate, letting you access and read them immediately.
Neotropical Ichthyology, 11(2):335-340, 2013
Copyright © 2013 Sociedade Brasileira de Ictiologia
Molecular diagnosis of the arowanas Osteoglossum ferreirai Kanazawa, 1966
and O. bicirrhossum (Cuvier, 1829) from the Orinoco and
Amazon River basins
María Doris Escobar L.1, Izeni P. Farias1, Donald C. Taphorn B.2, Miguel Landines3
and Tomas Hrbek1
The arowanas, fishes of Gondwanan origin, are represented in South America by the genus Osteoglossum. All species were
initially reported as being exclusive to the Amazon region, with O. ferreirai restricted to the Negro River basin and O.
bicirrhosum to the Amazon and Essequibo Rivers basin. Starting in the mid 1970’s it was reported that O. ferreirai also occurs
in the Orinoco River basin. In all regions the arowanas assumed significant socio-economic importance due to their popularity
in the international ornamental fish trade, leading to over-exploitation of both species in some areas. The Orinoco populations
are particularly heavily exploited, and thus conservation and management measures are needed. Both depend on the clarification
of taxonomic status, and phylogenetic distinctness of the Orinoco populations. With the goal of molecularly characterizing
the two species of Osteoglossum, and comparing populations of Osteoglossum from the Orinoco and Amazon basins, we
characterized individuals sampled from eight localities, one in the Orinoco River basin and seven in the Amazon River basin.
We sampled 39 individuals, obtaining 1004 base pairs, of which 79 were synapomorphies. Genetic distance between the two
species calculated using the HKY + G model of molecular evolution was 8.94%. Intraspecific distances ranged from 0.42% in
O. bicirrhosum to 0.10% in O. ferreirai. The genetic characterization confirmed the taxonomic status of O. ferreirai in the
Orinoco basin, and suggested that its distribution in the Orinoco basin is unlikely to be the result of vicariance or natural
dispersal, but rather an anthropic introduction.
Os aruanãs, peixes de origem Gondwana, são representados na América do Sul pelo gênero Osteoglossum. Todas as espécies
foram inicialmente reportadas como sendo exclusivamente da região Amazônica, com O. ferreirai restrito a bacia do Rio Negro e
O. bicirrhosum para a bacia Amazônica e rio Essequibo. No meio dos anos 70 foi reportado que O. bicirrhosum também ocorre
na bacia do rio Orinoco. Em todas as regiões os aruanãs são de significante importância socio-econômica devido a sua popularidade
no comércio internacional de peixes ornamentais, levando a sobre-exploração de ambas as espécies em algumas áreas. As
populações no rio Orinoco são particularmente muito exploradas, e assim medidas de conservação e manejo são necessárias.
Ambas as medidas dependem de clarificações do estatus taxonômico, e distinções filogenéticas das populações do Orinoco.
Com o objetivo de caracterizar molecularmente as duas espécies de Osteoglossum, e comparar as populações de Osteoglossum
das bacias do Orinoco e Amazonas, nós caracterizamos indivíduos amostrados de oito localidades, uma da bacia do rio Orinoco
e sete da bacia Amazônica. Nós amostramos 39 indivíduos, obtendo 1004 pares de bases, dos quais 97 foram sinapomorfias. A
distância genética entre as duas espécies, calculadas usando-se o modelo HKY+G de evolução molecular foi de 8.94%. Distâncias
intraespecíficas variaram de 0.42% em O. bicirrhosum a 0.10% em O. ferreirai. A caracterização genética confirmou o estatus
taxonômico de O. ferreirai na bacia do rio Orinoco, e sugere que sua distribuição na bacia do rio Orinoco é improvavel de ser o
resultado de um evento de vicariância ou dispersão natural, sendo melhor explicada como uma introdução antrópica.
Key words: Aruana, Biological introduction, Black arowana, Silver arowana.
1
Laboratório de Evolução e Genética Animal, Instituto de Ciências Biológicas, Universidade Federal do Amazonas, Manaus, AM,
Brazil.tomas@evoamazon.net (TH)
2
Aquatic Experts, 1822 North Charles Street, Belleville, IL, 62221, USA.
3
Laboratorio de Fisiología de Peces, Universidad Nacional de Colombia, Bogotá, Colombia.
335
336
Molecular diagnosis of the arowanas Osteoglossum ferreirai and O. bicirrhossum
Introduction
The osteoglosids, known as the bony tongue fishes, are
ancient creatures whose diversification is associated with
the breakup of Gondwana (Carroll, 1988; Kumazawa & Nishida,
2000). Currently, ten species are known with distribution in
the following regions: three in South America, one in Africa,
four in Asia and two in Australia (Nelson, 2006). In South
America, osteoglossids are represented by Arapaima gigas
of the family Arapaimidae, and Osteoglossum bicirrhosum
and Osteoglossum ferreirai of the family Osteoglossidae,
distributed in the Amazonas, Essequibo and Orinoco basins
(Reis, et al., 2003; Watkins, et al., 2004; Maldonado-Ocampo,
et al., 2008). The existence of arowanas in South America was
reported first by the Portuguese naturalist Alexandre
Rodriguez Ferreira during expeditions in the Brazilian Amazon
between the years 1783 to 1792. Based on specimens he
collected and his illustrations of the white arowana, Cuvier
erected the genus and described the species O. bicirrhosum.
It was Wallace during his 1845 expedition to the Negro River
(Brazil) who found and illustrated for the first time the black
arowana, however, this individual was lost during his return
trip to England. The black arowana was described in 1966 by
Kanazawa as Osteoglossum ferreirai, and its distribution was
restricted to The Negro River basin, with rest of the Amazon
and Essequibo basin being occupied by O. bicirrhossum.
Based on an ichthyological survey performed by Cala (1973),
the range of O. ferreirai was expanded to the lower Tomo
River in the Colombian Llanos. By the turn of the millennium,
O. ferreirai was being reported in the Bita, Tomo and Tuparro
Vichada Rivers by Mojica (1999), Lasso et al. (2005) and
Maldonado-Ocampo et al. (2006; 2008).
The arowanas from the Bita River appear to differ in their
reproductive biology from those of the Negro River. A study
of Gutiérrez et al. (2009) showed that the Bita River population
had lower fecundity, producing 38 to 127 oocites per female
and maturing at 62 cm TL, while in the Negro River populations
females produced an average of 136 oocites, and matured
sexually at 70 cm TL (Rabello-Neto, 1999; 2002). These
differences were viewed by the authors as indication of
population or even species level differences.
The two South American species of arowanas began to
be exported in large numbers for the aquarium trade in the
1970’s when they started to replace the Asian arowana
Scleropages formosus, also known as dragon fish, after S.
formosus was listed as endangered by CITES (Yue, et al.,
2003). Starting in the 1970’s, over-exploitation of South
American arowanas turned these species threatened with
extinction in Colombia (Alvarez-León, 2002a; 2002b). In 2005,
Traffic and WWF reunited prominent ichthyologists in the
“International Workshop of ornamental fishes from northern
South America”. They discussed the high vulnerability of
both Osteoglossum species, and proposed to include them in
Appendix II of CITES. They considered as first action the
necessity to clarify the taxonomic status of O. ferreirai in the
Orinoco basin, its divergence from the Negro basin
populations, and an assessment of genetic variability of O.
bicirrhossum in its area of distribution.
Material and Methods
We obtained and analyzed fishes caught by artisanal
fishermen at these locations: the Orinoco River basin in the
Bita River (Puerto Careño, n = 5), the Amazon River basin: the
Negro River (Barcelos, n = 7), the Putumayo River (Puerto
Leguizamo, n = 8), the Juruá River (Eirunepé, n = 5), the Madeira
River (Humaita, n = 4) and the Solimões-Amazonas (Tabatinga,
n = 4; Tefé, n = 3; Santarém, n = 3) Fig.1.
All tissue samples were preserved in absolute alcohol and
processed in the Laboratório de Evolução e Genética Animal
(LEGAL) at the Universidade Federal do Amazonas (UFAM),
Manaus, Brazil. DNA extraction was performed with the CTAB
method (Doyle & Doyle, 1987), and quantified by means of
electrophoresis in a 0.8% agarose gel against a known quantity
of a molecular size standard (Fermentas).
Subsequently we amplified via the PCR (Polymerase Chain
Reaction) a segment of mitochondrial DNA between the 3' end
of cytochrome oxidase subunit 2 and the 5’ end of the
cytochrome oxidase subunit 3, including the tRNA Lysine, and
ATPase subunits 6 and 8. This region was amplified using the
primers L8106 5’-TGGGTGTTAAAATAGATGC-3’ and H9264
5’-GAGGAGAGCRGCRGATGCCCC-3’ and sequenced with
L8106, L8537 5’-TGAAACTGACCATGACACTAAG-3’ and
H8516 5’-CTTGTGTCATGGTCAGTTTCA-3’ developed by
Hrbek et al. (2005) for Arapaima gigas.
The PCR had a total volume of 15ìl, and included 5.6 µl of
ddH2O, 1.2 µl of 25 mM MgCl2, 1.2 µl of 10X buffer (75 mM Tris
HCl, 50 mM KCl, 20 mM (NH4)2SO4), 1.5 µl of 10 mM dNTP, 1.5
Fig. 1. Sampling localities in the Amazon and Orinoco basins
of O. bicirrhossum and O. ferreirai . Base map was obtained
from Online Map Creation (Geomar) currently available as
Planiglobe (http://www.planiglobe.com/).
M. D. Escobar L., I. P. Farias, D. C. Taphorn B., M. Landines & T. Hrbek
µl of each primer (2 µM), 1.0 µl nuclease-free BSA (2.5 mg / ml),
0.5 µl of Fermentas Taq DNA polymerase (1 U/µl) and 1.0 µl
DNA (~20 ng/µl).
The PCR reactions were tested on 1% agarose gel.
Sequencing reactions were performed using the BigDye
Terminator Kit V.3.1. according to the manufacturer’s
guidelines and the product was analyzed on an ABI 3130xl
automated sequencer. The sequences obtained were viewed
and edited by using the program BioEdit (Hall, 1999). The
dataset was aligned in Crustal W (implemented within Bioedit)
and then adjusted manually (Thompson, et al., 1996). The
composition of nucleotide bases, identification of variable
sites, stop codons, transversions/transitions and genetic
distances was estimated in the program MEGA5 (Tamura, et
al., 2011). The number and frequency of haplotypes was
determined in the program DnaSP v.5 (Librado & Rozas, 2009).
The program Modeltest 3.7 was used to test for the best
molecular evolutionary model fitting the dataset (Posada &
Crandall, 1998). Phylogenetic reconstruction was performed
using the maximum likelihood method (ML) implemented in
the program Treefinder (Jobb, et al., 2004) and support for
relationships was assessed via 1000 bootstrap pseudoreplications. We used Scleropages formosus and Scleropages
leichardti as outgroups (Genbank No. DQ023143, FJ890319).
Phylogenetic species were identified using the Population
Aggregation Analysis algorithm, that involves a search for
fixed differences between local populations, followed by
successive rounds of aggregation of populations and
previously aggregated population groups that are not distinct
from each other (Davis & Nixon, 1992), and complementary
analytical procedures described by Cook et al. (2010).
Results
The final aligned sequences had a length of 1004 bp and
79 of which were diagnostic for the two species. The 79
synapomorphies were divided between 73 transitions and 6
transversions. The base pair composition of the data set was
A = 27.2%, T = 29.1%, C = 30.2%, G = 13.4%. In total eleven
haplotypes were identified for both species, seven in O.
bicirrhosum and four in O. ferreirai. In both species a single
haplotype was the most frequent (online Supplement 1-3).
Haplotypes are available under the GenBank accession
numbers JQ436740 to JQ436750.
The maximum likelihood topology using the HKY+G model
of molecular evolution (Posada & Crandall, 1998) showed the
formation of two evolutionary lineages representing the two
species. The clade representing O. bicirrhosum split into two
sub-clades. The first comprised the majority of individuals from
the Putumayo (Puerto Leguizamo), Jurua (Eirunepé) and
Solimões (Leticia) Rivers and the second was formed by majority
of individuals in the Madeira (Humaitá), Solimões (Tefé) and
Amazon (Santarém) Rivers. The clade representing O. ferreirai
indicated the existence of a single evolutionary lineage
consisting of individuals from the Bita (Orinoco basin) and
from the Negro (Amazon basin) Rivers (Fig 2). Genetic
337
divergence between the two species of Osteoglossum averaged
at 8.94%, while intraspecific divergence in O. bicirrhosum was
0.42% and O. ferreirai was 0.10% (Tables 1-2).
Population Aggregation Analysis (online Supplement) also
resulted in the identification and diagnosis of two species:
Osteoglossum bicirrhosum (Cuvier, 1829)
DNA Diagnosis. This species can be distinguished from O.
ferreirai by combinations of the following nucleotide
characters: COII region: 528T, 561C, 567C, 582A, 585G, 603C,
606C, 612T, 618C, 648G. Lysine tRNA region: 22T, 41C, 45G.
ATP8 region: 6A, 12C, 15T, 27G, 30A, 33G, 37T, 54G, 72C, 84C;
112A. ATP6 region: 87C, 96C, 111C, 114C, 138A, 150T, 192G,
198C, 213T, 217T, 219T, 226G, 249C, 258G, 265T, 267A, 282C,
285A, 294G, 300G, 312G, 313T, 318G, 324A, 330G, 333C, 339T,
351T, 363A, 393T, 394G, 410C, 429C, 447C, 448C, 450C, 474G,
489T, 495C, 508T, 528C, 552T, 576G, 583A, 585T, 586T, 591T,
600A, 607C, 625G, 627T, 633A, 654G, 657A.
Morphological features (data from Kanazawa, 1966). Tip of
snout to anus 45.5 to 53.2; snout to origin of dorsal fin 55.9 to
61.9; snout length 3.7 to 6.5; eye diameter 3.1 to 8.7; head length
19.6 to 26.9; body depth 14.6 to 20.7. These data are given in
percentages with respect to standard length. Branched dorsal
fin rays 42 to 50; unbranched anal fin rays 49 to 58; pectoral
rays i, 6; scales along the lateral line to base of caudal fin 30 to
37; pre-dorsal scales 16 to 19. Adult color is varied and can be
silver, yellow, or dark greenish, paler ventrally.
Distribution. Essequibo River basin; Amazon River basin not
including the Negro River basin except its Branco River
affluent.
Osteoglossum ferreirai Kanazawa, 1966
DNA Diagnosis. This species can be distinguished from O.
bicirrhosum combinations of the following nucleotide
characters: COII region: 528C, 561T, 567T, 582G, 585A, 603T,
606T, 612C, 618T, 648A. Lysine tRNA region: 22C, 41T, 45A.
ATP8 region: 6G, 12T, 15C, 27A, 30G, 33A, 37C, 54A, 72T, 84T,
112G. ATP6 region: 87T, 96C, 111T, 114T, 138G, 150C, 192A,
198T, 213C, 217C, 219A, 226A, 249T, 258A, 265C, 267G, 282T,
285C, 294A, 300A, 312A, 313C, 318A, 324C, 330A, 333T, 339C,
351G, 363G, 393A, 394A, 410T, 429T, 447T, 448T, 450A, 474T,
489C, 495T, 508C, 528T, 552C, 576A, 583G, 585C, 586C, 591C,
600G, 607T, 625A, 627C, 633G, 654A, 657G.
Morphological features (data from Kanazawa, 1966). Tip of
snout to anus 48.5; snout to origin of dorsal fin 53.6; snout
length 6.4; eye diameter 7.0; head length 22.6; body depth
13.9. These data are given in percentages with respect to
standard length. Branched dorsal fin rays 52 to 57;
unbranched anal fin rays 61 to 66; pectoral rays i, 6; scales
along the lateral line to base of caudal fin 37 to 40; pre-dorsal
scales 15 to 19. Adult color is dark bluish, paler ventrally.
338
Molecular diagnosis of the arowanas Osteoglossum ferreirai and O. bicirrhossum
Fig. 2. Maximum likelihood phylogenetic hypothesis of relationships of Osteoglossum individuals representing the species O.
bicirrhossum and O. ferreirai.
Distribution: Negro River basin, including the affluent Branco
River; Orinoco River basin.
Discussion
Phylogenetic and Population Aggregation Analysis
clearly separate and diagnose the two species of
Osteoglossum, and place the Colombian populations within
the species O. ferreirai. This diagnosis confirms that of Cala
(1973). There were no unique haplotypes in the Orinoco basin
samples of O. ferreirai, indicating a very recent divergence
of the populations of the Orinoco and Negro River basins.
We suggest that the lack of divergence of the Orinoco basin
samples is most likely due to a recent introduction from the
Negro to the Orinoco river basin. Based on ichthyofaunal
surveys of areas that can potentially serve as biological
corridors and links between the two basins such as the
Casiquiare channel and the floodplain of the Inirida and
Atabapo Rivers, neither O. ferreirai nor O. bicirrhossum is
Table 1. Table of K2P distances (Kimura, 1980) between the
two species of Osteoglossum and the Negro and Orinoco
River populations of O. ferreirai.
O. ferreirai Negro
O. ferreirai Orinoco
O. bicirrhossum
0.0892
0.0897
O. ferreirai Negro
0.0009
M. D. Escobar L., I. P. Farias, D. C. Taphorn B., M. Landines & T. Hrbek
Table 2. Table of intraspecific and intragroup K2P distances
(Kimura, 1980) observed in the two species of Osteoglossum
and the Negro and Orinoco River populations of O. ferreirai.
O. bicirrhossum
O. ferreirai Negro
O. ferreirai Orinoco
0.0042
0.0010
0.0010
present in these areas (Maldonado-Ocampo, et al., 2006;
Maldonado-Ocampo, et al., 2008; Winemiller et al., 2008;
Lasso, et al., 2009; Miller-Hurtado, et al., 2009; Winemiller &
Willis, 2011). Additionally, based on fauna surveys (Cala, 1973;
Mojica, 1999; Lasso et al., 2005; Maldonado-Ocampo et al.,
2006; 2008) the expansion in the distribution of O. ferreirai in
Colombia appears to have proceeded from north to south, i.e.
towards the Negro River basin. Therefore, we consider it
highly unlikely that the occupation of the Orinoco basin would
proceed via geographically intermediate areas between the
Orinoco and Negro basins. Natural long distance dispersal
and colonization also appears an unlikely mechanism.
Although we could not find any reference or report that would
suggest that O. ferreirai in the Orinoco basin was or could
have been introduced, we consider an introduction the most
likely explanation. The area of origin of O. ferreirai from the
Orinoco basin is unclear, however, and will require a detailed
characterization of O. ferreirai from the Negro basin.
The possibility and what we consider to be the most likely
scenario that O. ferreirai has been introduced in the Orinoco
basin, constitutes a management and conservation dilemma.
The Orinoco O. ferreirai have genetic diversity comparable
to that of the fishes found in the Negro River basin, and
appear to have expanded their geographic distribution. Thus
they appear likely to increase in density and expand
geographically if an ecological opportunity presents itself.
On one hand, O. ferreirai makes an important contribution to
the local economy, and is a highly desirable export species in
the aquarium trade. In the Orinoco basin it appears to be
over-exploited (Alvarez-León, 2002b; CCI, 2009), hence
implementing a management and/or conservation program
may result in not only its recovery, but also its expansion to
other areas where it is currently not found. Being a top of the
pyramid predator, and predating not only aquatic but also
terrestrial vertebrates, could have a major impact on population
ecology of the areas where densities O. ferreirai will increase.
On the other hand, conservation programs normally try to
eliminate introduced species. However, it seems unlikely that
a program to eradicate O. ferreirai would succeed due to the
area already colonized, and the remoteness of these areas. A
third option is to do nothing. Under this scenario, the global
aquarium market will largely determine if the species increases
in density and colonizes new areas, remains at status quo, or
its densities will decrease and possibly will even experience
local extinctions. Ultimately, however, conservation and
management decisions will be mandated by governmental
agencies and will likely tend to balance socioeconomic needs
with environmental protection.
339
Acknowledgments
We thank the Colombian Institute for Rural Development
(INCODER), under-institute of Fisheries and Aquiculture, and
the National University of Colombia with the support of the
project “Evaluación biológico-pesquera de la arawana azul y
la sapuara”. Financial support was provided by CNPq/
COLCIENCIAS cooperative grant # 490682/2010-3 to TH. TH
and IPF were supported by a Bolsa de Pesquisa scholarship
from CNPq during the study. MDE is supported by a
fellowship from CAPES.
Literature Cited
Alvarez-León, R. 2002a. Osteoglossum bicirrhosum Pp. 168-170.
In: Mojica, J. I., C. Castellanos, J. S. Usma & R. Alvarez-León
(Eds.). Libro rojo de peces de aguas dulces de Colombia Serie
Libros Rojos de Especies Amenzadas de Colombia. Bogotá,
Colombia, Instituto de Ciencias Naturales Universidad Nacional
de Colombia, Instituto Alexander von Humboldt, Ministerio de
Medio Ambiente.
Alvarez-León, R. 2002b. Osteoglossum ferreirai. Pp. 121-122. In:
Mojica, J. I., C. Castellanos, J. S. Usma & R. Alvarez-León
(Eds.). Libro rojo de peces de aguas dulces de Colombia Serie
Libros Rojos de Especies Amenzadas de Colombia. Bogotá,
Colombia, Instituto de Ciencias Naturales Universidad Nacional
de Colombia, Instituto Alexander von Humboldt, Ministerio de
Medio Ambiente.
Cala, P. 1973. Presencia de Osteoglossum en los Llanos (Orinoquia).
Acta Zoologica Colombiana, 18: 8.
Carroll, R. L. 1988. Vertebrate Paleontology and Evolution. New
York, NY, W. H. Freeman and Company, 698p.
CCI (Corporación Colombia Internacional), 2009. Captura y
comercialización de la arawana. Sistema de Información de Pesca
y Acuicultura. Boletin mayo 17p. http://www.cci.org.co/cci/
cci_x/datos/BoletinesIncoder/Mensual/BolMay2009.pdf
(Accessed 14 February 2009).
Cook, L. G., R. D. Edwards, M. D. Crisp & N. B. Hardy. 2010.
Need morphology always be required for new species
descriptions? Invertebrate Systematics, 24: 322-326.
Davis, J. I. & K. C. Nixon. 1992. Populations, genetic variation,
and the delimitation of phylogenetic species. Systematic
Biology, 41: 421-435.
Doyle, J. J. & J. L. Doyle. 1987. A rapid DNA isolation procedure
for small quantities of fresh leaf tissue. Phytochemical Bulletin,
19: 11-15.
Gutiérrez, M., Y. López-Pinto & A. I. Sanabria. 2009. Aspectos
reproductivos de la “arawana azul”, Osteoglossum ferreirai
Kanazawa, 1966 (Osteoglossiformes: Osteoglossidae), durante
el ciclo hidrológico de mayo de 2007 a mayo 2009 en el río Bita
(Vichada), Colombia, p. 134. In: X Simposio Colombiano de
Ictiología, II Encuentro Colombo-Venezolano de Ictiólogos, I
Encuentro Suramericano de Ictiólogos. Vol. 31. Actualidades
Biológicas, Medellin, Colombia.
Hall, T. 1999. BioEdit: a user-friendly biological sequence alignment
editor and analysis program for Windows 95/98/NT. Nucleic
Acids Symposium Series, 41: 95-98.
Hasegawa, M., H. Kishino & T. A. Yano. 1985. Dating of the humanape splitting by a molecular clock of mitochondrial DNA.
Journal of Molecular Evolution, 22: 160-174.
340
Molecular diagnosis of the arowanas Osteoglossum ferreirai and O. bicirrhossum
Hrbek, T., I. P. Farias, M. Crossa, I. Sampaio, J. I. R. Porto & A.
Meyer. 2005. Population genetic analysis of Arapaima gigas,
one of the largest freshwater fishes of the Amazon basin:
implications for its conservation. Animal Conservation, 8: 297308.
Jobb, G., A. v. Haeseler & K. Strimmer. 2004. TREEFINDER: a
powerful graphical analysis environment for molecular
phylogenetics. BMC Evolutionary Biology 4: 1-9.
Kanazawa, R. H. 1966. The fishes of the genus Osteoglossum with
a description of a new species from the rio Negro. Ichthyology
and Aquarium Journal, 37: 161-172.
Kimura, M. 1980. A simple method for estimating evolutionary
rates of base substitutions through comparative studies of
nucleotide sequences. Journal of Molecular Evolution, 16: 111120.
Kumazawa, Y. & M. Nishida. 2000. Molecular phylogeny of
osteoglossoids: A new model for Gondwanian origin and plate
tectonic transportation of the Asian Arowana. Molecular
Biology and Evolution, 17: 1869-1878.
Lasso, C. A., J. I. Mojica, J. S. Usma, J. A. Maldonado-Ocampo, C.
DoNascimiento, D. C. Taphorn, F. Provenzano, O. Lasso-Alcalá,
G. Galvis, L. Vásquez, M. Lugo, A. Machado-Allison, R.
Royero, C. Suárez & A. Ortega-Lara. 2005. Peces de la cuenca
del río Orinoco. Parte I: lista de especies y distribución por
cuencas. Biota Colombiana, 5: 95-157.
Lasso, C. A., J. S. Usma, F. Villa, M. T. Sierra-Quintero, A. OrtegaLara, L. M. Mesa, M. A. Patiño, O. Lasso-Alcalá, M. A.
Morales-Betancourt, K. González-Oropesa, M. P. Quiceno, A.
Ferrer & C. F. Suárez. 2009. Peces de la estrella fluvial Inírida:
ríos Guaviare, Inírida, Atabapo y Orinoco (Orinoquia
colombiana). Biota Colombiana, 10: 89-122.
Librado, P. & J. Rozas. 2009. DnaSP v5: A software for
comprehensive analysis of DNA polymorphism data.
Bioinformatics, 2: 1451-1452.
Maldonado-Ocampo, J. A., M. Lugo, J. D. Bogotá-Gregory, C. A.
Lasso, L. Vásquez, J. S. Usma, D. C. Taphorn & F. Provenzano
R. 2006. Peces del río Tomo, cuenca del Orinoco, Colombia.
Biota Colombiana, 7: 113-128.
Maldonado-Ocampo, J. A., R. P. Vari & J. S. Usma. 2008. Checklist
of the freshwater fishes of Colombia. Biota Colombiana, 9:
143-237.
Miller-Hurtado, H., D. C. Taphorn B. & J. S. Usma. 2009. Lista
preliminar de los peces del río Papunahua, cuenca del río Inírida
– departamento del Vaupés, Colombia. Biota Colombiana, 10:
163-169.
Mojica, J. I. 1999. Lista preliminar de las especies de peces
dulceacícolas de Colombia. Revista de la Academia Colombiana
de Ciencias, 23: 547-566.
View publication stats
Nelson, J. S. 2006. Fishes of the World, 4th Edition. New York, NY,
John Wiley & Sons, 624 p.
Posada, D. & K. A. Crandall. 1998. MODELTEST: Testing the
model of DNA substitution. Bioinformatics, 14: 817-818.
Rabello-Neto, J. G. 1999. Biologia reprodutiva e alimentação natural
do Arauanã preto Osteoglossum ferreirari (Kanazawa, 1966),
no municipio de Barcelos, Médio rio Negro, Amazonas, Brasil.
Unpublished Dissertation, Universidade Federal do Amazonas
(UFAM), Manaus, AM, Brazil. 32p.
Rabello-Neto, J. G. 2002. Variabilidad morfológica do aruanã preto,
Osteoglossum ferreirai (Kanazawa, 1966) em seis área de pesca
do médio Rio Negro, município de Barcelos, AM, Brasil:
Implicações para a identificação de estoques. Unpublished
Dissertation, Instituto Nacional de Pesquisas da Amazônia
(INPA) and Universidade Federal do Amazonas (UFAM),
Manaus, AM, Brazil. 45p.
Reis, R. E., S. O. Kullander & C. J. Ferraris (Eds.). 2003. Check
List of the Freshwater Fishes of South and Central America.
Porto Alegre, Brazil, EDIPUCRS, 734p.
Tamura, K., D. Peterson, N. Peterson, G. Stecher, M. Nei & S.
Kumar. 2011. MEGA5: Molecular Evolutionary Genetics
Analysis using maximum likelihood, evolutionary distance, and
maximum parsimony methods. Molecular Biology and
Evolution, 28: 2731-2739.
Thompson, J. D., D. G. Higgins & T. J. Gibson. 1996. CLUSTAL
W: improving the sensitivity of progressive multiple sequence
alignment through sequence weighting, position specific gap
penalties and weight matrix choice. Nucleic Acids Research, 22:
4673-4680.
Watkins, G., W. Saul, E. Holm, C. Watson, D. Arjoon & J. Rjoon.
2004. The fish fauna of the Iwokrama forest. Proceedings of the
Academy of Natural Sciences of Philadelphia, 154: 39-53.
Winemiller, K., López-Fernández, H., Taphorn D., Nico L. G & B.
Duque. Fish assemblages of the Casiquiare river, a corridor and
zoogeographical filter for dispersal between the Orinoco and
Amazon basins. Journal of Biogeography, 35: 1551-1563
Winemiller K & S Willis. 2011. The Vaupes Arch and Casiquiare
Canal barriers and passages. Pp 225 – 242 In James S Albert J
& Roberto E. Reis (Eds). Historical Biogeography of Neotropical
Freshwater Fishers. University of California Press, 369 p.
Yue, G. H., D. Ong, C. C. Wong, L. C. Lim & L. Orban. 2003. A
strain-specific and a sex-associated STS marker for Asian
arowana (Scleropages formosus, Osteoglossidae). Aquaculture
Research, 34: 951-957.
Submitted January 23, 2012
Accepted February 26, 2013 by Claudio Oliveira
Published June 28, 2013