The representatives of cyprinid lineage 'Poropuntiinae' with 16 recognized genera and around 100 species form a significant part of Southeast Asian ichthyofauna. Cytogenetics are valuable when studying fish evolution, especially the dynamics of repetitive DNAs, such as ribosomal DNAs (5S and 18S) and microsatellites, that can vary between species. Here, karyotypes of seven 'poropuntiin' species, namely Cosmochilus harmandi, Cyclocheilichthys apogon, Hypsibarbus malcomi, H. wetmorei, Mystacoleucus chilopterus, M. ectypus, and Puntioplties proctozysron occurring in Thailand were examined using conventional and molecular cytogenetic protocols. Variable numbers of uni- and bi-armed chromosomes indicated widespread chromosome rearrangements with a stable diploid chromosome number (2n) of 50. Examination with fluorescence in situ hybridization using major and minor ribosomal probes showed that Cosmochilus harmandi, Cyclocheilichthys apogon, and Puntioplites proctozystron all had one chromosomal pair with 5S rDNA sites. However, more than two sites were found in Hypsibarbus malcolmi, H. wetmorei, Mystacoleucus chilopterus, and M. ectypus. The number of chromosomes with 18S rDNA sites varied amongst their karyotypes from one to three; additionally, comparative genomic hybridization and microsatellite patterns varied among species. Our results reinforce the trend of chromosomal evolution in cyprinifom fishes, with major chromosomal rearrangements, while conserving their 2n.

Departamento de Genética e Evolução Universidade Federal de São Carlos Rodovia Washington Luiz Km 235 C P 676 São Carlos 13565 905 Brazil

Department of Biology Faculty of Science Khon Kaen University Muang Khon Kaen 40002 Thailand

Department of Fundamental Science Faculty of Science and Technology Surindra Rajabhat University Muang Surin 32000 Thailand

Department of Science Faculty of Science and Technology Prince of Songkla University Pattani 94000 Thailand

Division of Biology Department of Science Faculty of Science and Technology Rajamangala University of Technology Krungthep Bangkok 10120 Thailand

Faculty of Applied Science and Engineering Khon Kaen University Nong Khai Campus Muang Nong Khai 43000 Thailand

Institute of Human Genetics University Hospital Jena 07747 Jena Germany

Laboratory of Fish Genetics Institute of Animal Physiology and Genetics Czech Academy of Sciences Rumburská 89 277 21 Liběchov Czech Republic

Program in Biology Faculty of Science Buriram Rajabhat University Muang Buriram 31000 Thailand

School of Agriculture and Natural Resources University of Phayao Tumbol Maeka Muang Phayao 56000 Thailand

Zobrazit více v PubMed

Tan M., Armbruster J.W. Phylogenetic classification of extant genera of fishes of the order Cypriniformes (Teleostei: Ostariophysi) Zootaxa. 2018;4476:6–39. doi: 10.11646/zootaxa.4476.1.4. PubMed DOI

Yang L., Sado T., Hirt M.V., Pasco-Viel E., Arunachalam M., Li J., Wang X., Freyhof J., Saitoh K., Simons A.M., et al. Phylogeny and polyploidy: Resolving the classification of cyprinine fishes (Teleostei: Cypriniformes) Mol. Phylogenetics Evol. 2015;85:97–116. doi: 10.1016/j.ympev.2015.01.014. PubMed DOI

Yang L., Naylor G.J., Mayden R.L. Deciphering reticulate evolution of the largest group of polyploid vertebrates, the subfamily cyprininae (Teleostei: Cypriniformes) Mol. Phylogenetics Evol. 2021;166:107323. doi: 10.1016/j.ympev.2021.107323. PubMed DOI

Wolf U., Ritter H., Atkin N.B., Ohno S. Polyploidization in the fish family Cyprinidae, order Cypriniformes. Hum. Genet. 1969;7:240–244. doi: 10.1007/BF00273173. PubMed DOI

Sola L., Gornung E. Classical and molecular cytogenetics of the zebrafish, Danio rerio (Cyprinidae, Cypriniformes): An overview. Genetica. 2001;111:397–412. doi: 10.1023/A:1013776323077. PubMed DOI

Sember A., Bohlen J., Šlechtová V., Altmanová M., Symonová R., Ráb P. Karyotype differentiation in 19 species of river loach fishes (Nemacheilidae, Teleostei): Extensive variability associated with rDNA and heterochromatin distribution and its phylogenetic and ecological interpretation. BMC Evol. Biol. 2015;15:251. doi: 10.1186/s12862-015-0532-9. PubMed DOI PMC

Šlechtová V., Bohlen J., Freyhof J., Ráb P. Molecular phylogeny of the Southeast Asian freshwater fish family Botiidae (Teleostei: Cobitoidea) and the origin of polyploidy in their evolution. Mol. Phylogenetics Evol. 2006;39:529–541. doi: 10.1016/j.ympev.2005.09.018. PubMed DOI

Ráb P., Hnátková E., Majtánová Z., Šlechtová V.B., Bohlen J. Karyotype Record for the Morphologically Derived, Rarely Collected, Freshwater Fish Ellopostoma mystax (Cypriniformes, Cobitoidea, Ellopostomatidae) Ichthyol. Herpetol. 2021;109:998–1001. doi: 10.1643/i2020032. DOI

Bohlen J., Völker M., Rábová M., Ráb P. Note on the banded karyotype of the enigmatic South Asian loach Vaillantella maassi (Cypriniformes) Ichthyol. Res. 2007;55:82–84. doi: 10.1007/s10228-007-0007-0. DOI

Cioffi M.B., Bertollo L.A.C. Chromosomal distribution and evolution of repetitive DNAs in fish. In: Garrido-Ramos M.A., editor. Repetitive DNA Genome Dyn. Basel. 1st ed. Volume 7. Karger Publishers; Basel, Switzerland: 2012. pp. 197–221. PubMed

Goes C.A.G., dos Santos R.Z., Aguiar W.R.C., Alves D.C.V., Silva D.M.Z.D.A., Foresti F., Oliveira C., Utsunomia R., Porto-Foresti F. Revealing the Satellite DNA History in Psalidodon and Astyanax Characid Fish by Comparative Satellitomics. Front. Genet. 2022;13:884072. doi: 10.3389/fgene.2022.884072. PubMed DOI PMC

Miura I., Shams F., Lin S.M., Cioffi M.B., Liehr T., Al-Rikabi A., Kuwana C., Srikulnath K., Higaki Y., Ezaz T. Evolution of a multiple sex-chromosome system by three-sequential translocations among potential sex-chromosomes in the Taiwanese frog Odorrana swinhoana. Cells. 2021;10:661. doi: 10.3390/cells10030661. PubMed DOI PMC

Waters P.D., Patel H.R., Ruiz-Herrera A., Álvarez-González L., Lister N.C., Simakov O., Ezaz T., Kaur P., Frere C., Grützner F., et al. Microchromosomes are building blocks of bird, reptile, and mammal chromosomes. Proc. Natl. Acad. Sci. USA. 2021;118:e2112494118. doi: 10.1073/pnas.2112494118. PubMed DOI PMC

Kretschmer R., de Souza M.S., Furo I.d.O., Romanov M.N., Gunski R.J., Garnero A.d.V., de Freitas T.R.O., de Oliveira E.H.C., O’connor R.E., Griffin D.K. Interspecies Chromosome Mapping in Caprimulgiformes, Piciformes, Suliformes, and Trogoniformes (Aves): Cytogenomic Insight into Microchromosome Organization and Karyotype Evolution in Birds. Cells. 2021;10:826. doi: 10.3390/cells10040826. PubMed DOI PMC

Ferreira P.H.N., Souza F.H.S., de Moraes R.L., Perez M.F., Sassi F.D.M.C., Viana P.F., Feldberg E., Ezaz T., Liehr T., Bertollo L.A.C., et al. The Genetic Differentiation of Pyrrhulina (Teleostei, Characiformes) Species is Likely Influenced by Both Geographical Distribution and Chromosomal Rearrangements. Front. Genet. 2022;13:869073. doi: 10.3389/fgene.2022.869073. PubMed DOI PMC

Phimphan S., Chaiyasan P., Suwannapoom C., Reungsing M., Juntaree S., Tanomtong A., Supiwong W. Comparative karyotype study of three Cyprinids (Cyprinidae, Cyprininae) in Thailand by classical cytogenetic and FISH techniques. Comp. Cytogenet. 2020;14:597–612. doi: 10.3897/CompCytogen.v14i4.54428. PubMed DOI PMC

Pereira C., Ráb P., Collares-Pereira M. Chromosomes of Iberian Leuciscinae (Cyprinidae) Revisited: Evidence of Genome Restructuring in Homoploid Hybrids Using Dual-Color FISH and CGH. Cytogenet. Genome Res. 2013;141:143–152. doi: 10.1159/000354582. PubMed DOI

Symonová R., Majtánová Z., Sember A., Staaks G.B., Bohlen J., Freyhof J., Rábová M., Ráb P. Genome differentiation in a species pair of coregonine fishes: An extremely rapid speciation driven by stress-activated retrotransposons mediating extensive ribosomal DNA multiplications. BMC Evol. Biol. 2013;13:42–52. doi: 10.1186/1471-2148-13-42. PubMed DOI PMC

Symonová R., Sember A., Majtánová Z., Ráb P. Characterization of fish genomes by GISH and CGH. In: Ozouf-Costaz C., Pisano E., Foresti F., de Almeida L., editors. Fish Cytogenetic Techniques. 1st ed. CRC Press; London, UK: 2015. pp. 118–131.

Doležálková M., Sember A., Marec F., Ráb P., Plötner J., Choleva L. Is premeiotic genome elimination an exclusive mechanism for hemiclonal reproduction in hybrid males of the genus Pelophylax? BMC Genet. 2016;17:100. doi: 10.1186/s12863-016-0408-z. PubMed DOI PMC

Majtánová Z., Symonová R., Arias-Rodriguez L., Sallan L., Ráb P. “Holostei versus Halecostomi” Problem: Insight from Cytogenetics of Ancient Nonteleost Actinopterygian Fish, Bowfin Amia calva: Molecular Cytogenetics of Amia calva. J. Exp. Zool. Part B Mol. Dev. Evol. 2017;328:620–628. doi: 10.1002/jez.b.22720. PubMed DOI

Majtánová Z., Unmack P.J., Prasongmaneerut T., Shams F., Srikulnath K., Ráb P., Ezaz T. Evidence of Interspecific Chromosomal Diversification in Rainbowfishes (Melanotaeniidae, Teleostei) Genes. 2020;11:818. doi: 10.3390/genes11070818. PubMed DOI PMC

Shams F., Dyer F., Thompson R., Duncan R.P., Thiem J.D., Majtánová Z., Ezaz T. Karyotypes and Sex Chromosomes in Two Australian Native Freshwater Fishes, Golden Perch (Macquaria ambigua) and Murray Cod (Maccullochella peelii) (Percichthyidae) Int. J. Mol. Sci. 2019;20:4244. doi: 10.3390/ijms20174244. PubMed DOI PMC

LeVan A., Fredga K., Sandberg A.A. Nomenclature for Centromeric Position on Chromosomes. Hereditas. 1964;52:201–220. doi: 10.1111/j.1601-5223.1964.tb01953.x. DOI

Donsakul T., Magtoon W., Rangsiruji A. Karyological studies of four Cyprinid fishes: Barbichthys nitidus, Mystacoleucus argenteus, Cychocheilichthys lagleri and Systomus sp. 1 from Thailand; Proceedings of the 44rd Kasetsart University Annual Conference: Fisheries; Bangkok, Thailand. 30 January–2 February 2006; Bangkok, Thailand: Kasetsart University; pp. 469–476.

Chantapan T. Master’s Thesis. Khon Kaen University; Khon Kaen, Thailand: 2015. Standardized karyotype and ideogram of cyprinid fishes (Subfamily: Cyprininae) in Thailand.

Seetapan K. Karyotypes of sex fish species of the family Cyprinidae; Proceedings of the 45th Kasetsart University Annual Conference: Fisheries; Bangkok, Thailand,. 30 January–2 February 2007; Bangkok, Thailand: Kasetsart University; pp. 749–758.

Magtoon W., Arai R. Karyotypes of five Puntius species and one Cyclocheilichtltys species (Pisces, Cyprinidae) from Thailand. Bull. Nat. Sci. Mus. Tokio. 1989;15:167–175.

Suzuki A., Taki Y., Mochizuki M., Hirata J. Chromosomal speciation in Eurasian and Japanese Cyprinidae (Pisces, Cypriniformes) Cytobios. 1995;83:171–186.

Wu G.M., Zhu X.P., Hu G., Luo J.R. The karyotype of Puntius gonionotus (Cyprinidae) Chin. J. Zool. 1991;26:20–21.

Donsakul T., Poopitayasathaporn A. Research Project. Srinakharinwirot University; Bangkok, Thailand: 2002. Karyotype of Fifteen Species of Cyprinid Fishes (Family Cyprinidae) from Thailand.

Khunda-Bukhsh A.R., Das J.K. Cytogenetic analyses in eight species of teleostean fishes (Pisces): Karyotype, multiple Ag-NORs, sex chromosomes. Res. Rev. BioScience. 2007;1:47–52.

Arai R. Fish Karyotypes: A Check List. Springer Science & Bussiness Media; Berlin/Heidelberg, Germany: 2011.

Donsakul T., Magtoon W., Rangsiruji A. Karyotypes of four cyprinid fishes: Cyclocheilichthys repasson, Cosmochilus harmandi, Poropuntius deauratus and Sikukia gudgeri from Thailand; Proceedings of the 43rd Kasetsart University Annual Conference: Fisheries, Kasetsart University; Bangkok, Thailand. 1–4 February 2005; pp. 344–351.

Chaiyasan P., Supiwong W., Saenjundaeng P., Seetapan K., Pinmongkhonkul S., Tanomtong A. A Report on Classical Cytogenetics of Hihgfin Barb Fish, Cyclocheilichthys armatus (Cypriniformes, Cyprinidae) Cytologia. 2018;83:149–154. doi: 10.1508/cytologia.83.149. DOI

Donsakul T., Rangsiruji A., Magtoon W. Karyotypes of four cyprinid fishes: Poropuntius normani, Hypsibarbus malcolmi, Scaphognathops bandanensis and Henicorhynchus caudiguttatus from Thailand; Proceedings of the 45th Kasetsart University Annual Conference: Fisheries, Kasetsart University; Bangkok, Thailand. 30 January–2 February 2007; pp. 740–748.

Magtoon W., Arai R. Karyotypes and distribution of nucleolus organizer regions in cyprinid fishes from Thailand. Jpn. J. Ichthyol. 1993;40:77–85. doi: 10.11369/jji1950.40.77. DOI

Donsakul T., Magtoon W. Karyotypes of four cyprinid fishes, Osteochilus melanopleura, Puntioplites proctozysron, Paralaubuca riveroi and Rasbora sumatrana, from Thailand; Proceedings of the 33rd Kasetsart University Annual Conference: Fisheries, Kasetsart University; Bangkok, Thailand. 30 January–1 February 1995; pp. 128–138.

Donsakul T., Magtoon W. Karyotypes of two cyprinid fishes, Hypsibarbus wetmorei and Morulius chrysophekadion, from Thailand; Proceedings of the 28th Congress on Science and Technology of Thailand, Queen Sirikit National Convention Centre; Bangkok, Thailand. 24–26 October 2002; p. 92.

Piyapong C. Master’s Thesis. Chulalongkorn University; Bangkok, Thailand: 1999. Karyotypes and Distribution of Nucleolus Organizer Regions in Four Cyprinid Species from Thailand.

Arai R., Magtoon W. Karyotypes of four cyprinid fishes from Thailand. Bull. Natl. Mus. Nat. Sci. 1991;17:183–188.

Zan R.G., Song Z., Liu W.G. Studies on karyotypes and nuclear DNA contents of some cyprinoid fishes, with notes on fish polyploids in China; Proceedings of the 2nd International Conference Indo-Pacific Fishes; Tokyo, Japan. 1 October 1986; pp. 877–885.

Donsakul T., Magtoon W. Karyotypes of seven cyprinid fishes: Systomos binotatus, Puntius brevis, Poropuntius laoensis, Labiobarbus siamensis, Catlocarpiosiamensis, Tor tambroides and Probarbus jullieni from Thailand. Srinakharinwirot Sci. J. 2008;24:80–92.

Zan R.G., Song Z., Liu W.G. Studies of karyotypes of seven species of fish in Barbinae, with a discussion on identification of fish polyploids. Zool. Res. 1984;5:82–90.

Supiwong W., Tanomtong A., Supanuam P., Jantarat S., Khakhong S., Sanoamuang L. A discovery of nucleolar organizer regions (NORs) polymorphism and karyological analysis of Smith’ s Barb, Puntioplites proctozysron (Cypriniformes, Cyprinidae) in Thailand. Cytologia. 2012;77:35–42. doi: 10.1508/cytologia.77.35. DOI

Rainboth W.J., Vidthayanon C., Yen M.D. Fishes of the Greater Mekong Ecosystem with Species List and Photographic Atlas, Miscellaneous Publications Museum of Zoology. University of Michigan; Ann Harbor, MI, USA: 2012. p. 314.

Bertollo L.A.C., Cioffi M.B., Moreira-Filho O. Direct chromosome preparation from freshwater teleost fishes. In: Ozouf-Costaz C., Pisano E., Foresti F., Toledo L.F.A., editors. Fish Cytogenetic Techniques: Ray-Fin Fishes and Chondrichthyans. 1st ed. RC Press; Boca Raton, FL, USA: 2015. pp. 21–26.

Howell W.M., Black D.A. Controlled silver-staining of nucleolus organizer regions with a protective colloidal developer: A 1-step method. Experientia. 1980;36:1014–1015. doi: 10.1007/BF01953855. PubMed DOI

Yano C.F., Bertollo L.A.C., Cioffi M.B. Fish-FISH: Molecular cytogenetics in fish species. In: Liehr T., editor. Fluorescence In Situ Hybridization (FISH) Springer; Berlin/Heidelberg, Germany: 2017. pp. 429–443. DOI

Martins C., Ferreira I.A., Oliveira C., Foresti F., Galetti P.M. A tandemly repetitive centromeric DNA sequence of the fish Hoplias malabaricus (Characiformes: Erythrinidae) is derived from 5S rDNA. Genetica. 2006;127:133–141. doi: 10.1007/s10709-005-2674-y. PubMed DOI

Cioffi M., Martins C., Centofante L., Jacobina U., Bertollo L. Chromosomal Variability among Allopatric Populations of Erythrinidae Fish Hoplias malabaricus: Mapping of Three Classes of Repetitive DNAs. Cytogenet. Genome Res. 2009;125:132–141. doi: 10.1159/000227838. PubMed DOI

Kubat Z., Hobza R., Vyskot B., Kejnovsky E. Microsatellite accumulation on the Y chromosome in Silene latifolia. Genome. 2008;51:350–356. doi: 10.1139/G08-024. PubMed DOI

Zwick M.S., Hanson R.E., Islam-Faridi M.N., Stelly D.M., Wing R.A., Price H.J., McKnight T.D. A rapid procedure for the isolation of C0t-1 DNA from plants. Genome. 1997;40:138–142. doi: 10.1139/g97-020. PubMed DOI

Saenjundaeng P., Supiwong W., Sassi F.M.C., Bertollo L.A.C., Rab P., Kretschmer R., Tanomtong A., Suwannapoom C., Reungsing M., Cioffi M.D.B. Chromosomes of Asian cyprinid fishes: Variable karyotype patterns and evolutionary trends in the genus Osteochilus (Cyprinidae, Labeoninae, “Osteochilini”) Genet. Mol. Biol. 2020;43:e20200195. doi: 10.1590/1678-4685-gmb-2020-0195. PubMed DOI PMC

Pereira R., Pereira V., Gomes I., Tomas C., Morling N., Amorim A., Prata M.J., Carracedo Á., Gusmão L. A method for the analysis of 32 X chromosome insertion deletion polymorphisms in a single PCR. Int. J. Leg. Med. 2011;126:97–105. doi: 10.1007/s00414-011-0593-2. PubMed DOI

Derenzini M. The AgNORs. Micron. 2000;31:117–120. doi: 10.1016/S0968-4328(99)00067-0. PubMed DOI

Gornung E. Twenty Years of Physical Mapping of Major Ribosomal RNA Genes across the Teleosts: A Review of Research. Cytogenet. Genome Res. 2013;141:90–102. doi: 10.1159/000354832. PubMed DOI

Rebordinos L., Cross I., Merlo A. High Evolutionary Dynamism in 5S rDNA of Fish: State of the Art. Cytogenet. Genome Res. 2013;141:103–113. doi: 10.1159/000354871. PubMed DOI

Cioffi M., Bertollo L.A.C., Villa M.A., de Oliveira E.A., Tanomtong A., Yano C.F., Supiwong W., Chaveerach A. Genomic Organization of Repetitive DNA Elements and Its Implications for the Chromosomal Evolution of Channid Fishes (Actinopterygii, Perciformes) PLoS ONE. 2015;10:e0130199. doi: 10.1371/journal.pone.0130199. PubMed DOI PMC

Symonová R., Howell W.M. Vertebrate Genome Evolution in the Light of Fish Cytogenomics and rDNAomics. Genes. 2018;9:96. doi: 10.3390/genes9020096. PubMed DOI PMC

Salim D., Gerton J.L. Ribosomal DNA instability and genome adaptability. Chromosom. Res. 2019;27:73–87. doi: 10.1007/s10577-018-9599-7. PubMed DOI

Roa F., Guerra M. Non-Random Distribution of 5S rDNA Sites and Its Association with 45S rDNA in Plant Chromosomes. Cytogenet. Genome Res. 2015;146:243–249. doi: 10.1159/000440930. PubMed DOI

Sochorová J., Garcia S., Gálvez F., Symonová R., Kovařík A. Evolutionary trends in animal ribosomal DNA loci: Introduction to a new online database. Chromosoma. 2017;127:141–150. doi: 10.1007/s00412-017-0651-8. PubMed DOI PMC

Fujiwara A., Abe S., Yamaha E., Yamazaki F., Yoshida M.C. Uniparental chromosome elimination in the early embryogenesis of the inviable salmonid hybrids between masu salmon female and rainbow trout male. Chromosoma. 1997;106:44–52. doi: 10.1007/s004120050223. PubMed DOI

Zhu H.-P., Gui J.-F. Identification of genome organization in the unusual allotetraploid form of Carassius auratus gibelio. Aquaculture. 2007;265:109–117. doi: 10.1016/j.aquaculture.2006.10.026. DOI

Valente G.T., Schneider C.H., Gross M.C., Feldberg E., Martins C. Comparative cytogenetics of cichlid fishes through genomic in-situ hybridization (GISH) with emphasis on Oreochromis niloticus. Chromosom. Res. 2009;17:791–799. doi: 10.1007/s10577-009-9067-5. PubMed DOI

Ditcharoen S., Bertollo L.A.C., Ráb P., Hnátková E., Molina W.F., Liehr T., Tanomong A., Triantaphyllidis C., Ozouf-Costaz C., Tongnunui S., et al. Genomic organization of repetitive DNA elements and extensive karyotype diversity of silurid catfishes (Teleostei: Siluriformes): A comparative cytogenetic approach. Int. J. Mol. Sci. 2019;20:3545. doi: 10.3390/ijms20143545. PubMed DOI PMC

Rampin M., Bi K., Bogart J.P., Collares-Pereira M.J. Identifying parental chromosomes and genomic rearrangements in animal hybrid complexes of species with small genome size using genomic in situ hybridization (GISH) Comp. Cytogenet. 2012;6:287–300. doi: 10.3897/compcytogen.v6i3.3543. PubMed DOI PMC

Knytl M., Kalous L., Symonová R., Rylková K., Ráb P. Chromosome studies of European cyprinid fishes: Cross-species painting reveals natural allotetraploid origin of a Carassius female with 206 chromosomes. Cytogenet. Genome Res. 2013;139:276–283. doi: 10.1159/000350689. PubMed DOI

Ráb P., Collares-Pereira M.J. Chromosomes of European cyprinid fishes (Cyprinidae, Cypriniformes). A review. Folia Zool. 1995;44:193–214.

Boroń A., Spoz A., Porycka K., Karolewska M., Ito D., Abe S., Kirtiklis L., Juchno D. Molecular cytogenetic analysis of the crucian carp, Carassius carassius (Linnaeus, 1758) (Teleostei, Cyprinidae), using chromosome staining and fluorescence in situ hybridisation with rDNA probes. Comp. Cytogenet. 2014;8:233–248. doi: 10.3897/compcytogen.v8i3.7718. PubMed DOI PMC

Knytl M., Kalous L., Rylková K., Choleva L., Merilä J., Ráb P. Morphologically indistinguishable hybrid Carassius female with 156 chromosomes: A threat for the threatened crucian carp, C. carassius, L. PLoS ONE. 2018;13:e0190924. doi: 10.1371/journal.pone.0190924. PubMed DOI PMC

Saenjundaeng P., Cioffi M.B., de Oliveira E.A., Tanomtong A., Supiwong W., Phimphan S., Collares-Pereira M.J., Sember A., Bertollo L.A.C., Liehr T., et al. Chromosomes of Asian cyprinid fishes: Cytogenetic analysis of two representatives of small paleotetraploid tribe Probarbini. Mol. Cytogenet. 2018;11:1–9. doi: 10.1186/s13039-018-0399-8. PubMed DOI PMC

Nirchio M., Rossi A.R., Foresti F., Oliveira C. Chromosome evolution in fishes: A new challenging proposal from Neotropical species. Neotrop. Ichtchyol. 2014;12:761–770. doi: 10.1590/1982-0224-20130008. DOI

Jacobina U.P., Martinez P.A., Cioffi M.B., Garcia J., Bertollo L.A.C., Molina W.F. Morphological and karyotypic differentiation in Caranx lugubris (Perciformes: Carangidae) in the St. Peter and St. Paul Archipelago, mid-Atlantic Ridge. Helgol. Mar. Res. 2014;68:17–25. doi: 10.1007/s10152-013-0365-0. DOI

Betancur-R R., Wiley E.O., Arratia G., Acero A., Bailly N., Miya M., Lecointre G., Ortí G. Phylogenetic classification of bony fishes. BMC Evol. Biol. 2017;17:162. doi: 10.1186/s12862-017-0958-3. PubMed DOI PMC

Chromosomes of Asian cyprinid fishes: Novel insight into the chromosomal evolution of Labeoninae (Teleostei, Cyprinidae)