Abstract
Moenkhausia cotinho is redescribed based on the examination of meristic and morphometric data and external morphological characters of specimens from the Amazon and Orinoco basins and from rivers in the Guyana. The species is distinguished from congeners based on the combination of a reticulate color pattern, the presence of dark spots on the posterior edge of scales of the longitudinal rows dorsal to the lateral line, the presence of a dark blotch on the caudal-fin base failing to reach the posterior tip of the middle caudal-fin rays; the dorsal region of the eyes red to orange in life; 5/3 longitudinal series of scales, above and below the lateral line, respectively; and the tips of the pectoral-fin rays not extending beyond the base of the pelvic fin. All specimens of M. cotinho have a completely pored literal line except for a few specimens from drainages in the Guyana. The redefinition of the taxonomic limits and geographic distribution of M. cotinho allowed us to reevaluate the taxonomic limits and geographic distribution of Hemigrammus newboldi, the identity of which has been previously questioned, but which is clearly diagnosed from M. cotinho based on the relatively longer pectoral fin, the presence of conical teeth in the posterior portion of the dentary, the absence of a reticulate color pattern of body and in the shorter longitudinal dark stripe on body. In addition, two new species of Moenkhausia are described: Moenkhausia rondoni sp n. from the rio Madeira basin, Brazil and Moenkhausia pirahan sp n. from the lower rio Madeira and rio Trombetas and rio Tapajós basins, Brazil.
References
Albornoz-Garzón, J.G., Méndez-López, A., DoNascimiento, C. & Lima, F.C.T. (2019) A new species of Hemigrammus (Characiformes: Characidae) from western Amazon River basin, Colombia. Journal of Fish Biology, 95 (3), 932–939.
https://doi.org/10.1111/jfb.14091
Benine, R.C., Mariguela, T.C. & Oliveira, C. (2009) New species of Moenkhausia Eigenmann, 1903 (Characiformes: Characidae) with comments on the Moenkhausia oligolepis species complex. Neotropical Ichthyology, 7 (2), 161–168.
https://doi.org/10.1590/s1679-62252009000200005
Bertaco, V.A., Jerep, F.C. & Carvalho, F.R. (2011) A new characid fish, Moenkhausia aurantia (Ostariophysi: Characiformes: Characidae), from the upper rio Tocantins basin in Central Brazil. Zootaxa, 2934 (1), 29–38.
https://doi.org/10.11646/zootaxa.2934.1.3
Calegari, B.B., Delapieve, M.L.S. & Sousa, L.M. (2016) Tutorial para preparação de mapas de distribuição geográfica. Boletim Sociedade Brasileira de Ictiologia, 118, 15–30.
Casatti, L., Pérez-Mayorga, M.A., Carvalho, F.R., Brejão, G.L. & Costa, I.D. (2013) The stream fish fauna from the rio Machado basin, Rondônia State, Brazil. Check List, 9 (6), 1496–1504.
https://doi.org/10.15560/9.6.1496
Castellanos, C. & Sánchez-Duarte, P. (2007) Moenkhausia cotinho Eigenmann, 1908. In: Sanabria-Ochoa, A.I., Victoria-Daza, P. & Beltrán, I.C. (Eds.), Peces de la Amazonía Colombiana com Énfasis em Especies de Interés Ornamental. Instituto Colombiano de Desarrollo Rural (INCODER), Universidad Nacional de Colombia e Instituto Amazónico de Investigaciones Científicas (SINCHI), Bogotá, pp. 132–133.
Castilho, Z.C. & Buckup, P.A. (Eds.) (2011) Ecorregião Aquática Xingú-Tapajós. CETEM, Rio de Janeiro, 246 pp.
Chernoff, B., Machado-Alison, A., Buckup, P.A. & Leon, R.R. (1994) Systematic status and neotype designation for Autanichthys giacopinii Fernández-Yépez with comments on the morphology of Bryconops melanurus (Bloch). Copeia, 1994 (1), 238–242.
https://doi.org/10.2307/1446694
Costa, W.J.E. (1994) Description of two new species of the genus Moenkhausia (Characiformes: Characidae) from the central Brazil. Zoologischer Anzeiger, 232 (1–2), 21–29.
Dagosta, F.C.P., Marinho, M.M.F. & Benine, R.C. (2015) A new species of Moenkhausia Eigenmann (Characiformes: Characidae) from the upper rio Juruena, Central Brazil. Zootaxa, 4032 (4), 417–425.
https://doi.org/10.11646/zootaxa.4032.4.6
Dagosta, F.C.P. & de Pinna, M.C.C. (2017) Biogeography of Amazonian fishes: deconstructing river basins as biogeographic units. Neotropical Ichthyology, 15 (3), e170034.
https://doi.org/10.1590/1982-0224-20170034
Dagosta, F.C.P. & de Pinna, M.C.C. (2019) The Fishes of the Amazon: distribution and biogeographical patterns, with a comprehensive list of species. Bulletin of the American Museum of Natural History, 431, 1–163.
https://doi.org/10.1206/0003-0090.431.1.1
DoNascimiento, C., Herrera-Collazos, E.E., Herrera-R, G.A., Ortega-Lara, A., Villa-Navarro, F.A., Oviedo, J.S.U. & Maldonado-Ocampo, J.A. (2017) Checklist of the freshwater fishes of Colombia: a Darwin Core alternative to the updating problem. Zookeys, 708, 25–138.
https://doi.org/10.3897/zookeys.708.13897
Eigenmann, C.H. (1908) Preliminary descriptions of new genera and species of tetragonopterid characins (Zoölogical Results of the Thayer Brazilian expedition). Bulletin of the Museum of Comparative Zoology, 52 (6), 91–106.
Eigenmann, C.H. (1912) The freshwater fishes of British Guiana, including a study of the ecological grouping of species, and the relation of the fauna of the plateau to that of the lowlands. Memoirs of the Carnegie Museum, 67 (5), i–xxii + 1–578, pls. 1–103.
https://doi.org/10.5962/bhl.part.14515
Eigenmann, C.H. (1914) Some results from Studies of South American Fishes. Indiana University Studies, 2, 19–31.
Eigenmann, C.H. (1917) The American Characidae—I. Memoirs of the Museum of Comparative Zoology, 43, 1–102, 16 pls.
Fernández-Yépez, A. (1949) Ramirezella newboldi nuevo género y especíe de pez Tetragonopteridae colectado em Venezuela. Evencias, 11 (6), 1–3.
Ferreira, K.M. & Lima, F.C.T. (2006) A new species of Knodus (Characiformes: Characidae) from the Rio Tiquié, upper Rio Negro basin, Brazil. Copeia, 2006 (4), 630–639.
https://doi.org/10.1643/0045-8511(2006)6[630:ANSOKC]2.0.CO;2.
Fink, W.L. (1979) A new species of Moenkhausia from the Mato Grosso region of Brazil (Pisces: Characidae). Breviora, 450, 1–12.
Fink, W.L. & Weitzman, S.H. (1974) The so-called Cheirodontin fishes of Central America with description of two new species (Pisces, Characidae). Smithsonian Contributions to Zoology, 172, 1–46.
https://doi.org/10.5479/si.00810282.172
Flores, H.G. (1995) Estado actual del conocimiento de la pesqueria en la Amazonia Peruana. Instituto de Investigaciones de la Amazonía Peruana, Iquitos, 53 pp.
Fowler, H.W. (1940) Zoological results of the second Bolivian expedition for the Academy of Natural Sciences of Philadelphia, 1936–1937. Part I: the fishes. Proceedings of the Academy of Natural Sciences of Philadelphia, 92, 43–103
Géry, J. (1964) Poissons characoides d’amazonie péruvienne. Beiträge zur Neotropischen Fauna, 4 (1), 1–44.
https://doi.org/10.1080/01650526409360375
Géry, J. (1977) Characoids of the World. T. F. H. Publications, Inc., Neptune City, New Jersey, 672 pp.
Géry, J. (1992) Description de deux nouvelles espèces proches de Moenkhausia lepidura (Kner) (Poissons, Characiformes, Tetragonopterinae), avec une revue du groupe. Revue Française d’Aquariologie and Herpétologie, 19, 69–78.
Géry, J. & Mahnert, V. (1984) Poissons characoïdes (ostariophysi) récoltés par CL. Vaucher en Amazonie péruvienne. Archives des sciences et compte rendu des séances de la Société, 37 (2), 171–180.
https://doi.org/10.5169/seals-740535
Géry, J. & Zarske, A. (2004) Moenkhausia heikoi n. sp., a new tetra (Teleostei: Ostariophysi: Characiformes: Characidae) from the rio Xingú basin, Brazil, with a supplementary description of the genus type species. Journal of Ichthyology and Aquatic Biology, 9 (1), 29–43.
Hoeinghaus, D.J., Winemiller, K.O. & Taphorn, D.C. (2004) Compositional change in fish assemblages along the Andean piedmont-Llanos floodplain gradient of the Río Portuguesa, Venezuela. Neotropical Ichthyology, 2 (2), 85–92.
https://doi.org/10.1590/S1679-62252004000200005
Holly, M. (1929) Einige neue Fischformen aus Brasilien. Anzeiger der Wissenschaften Wien, 66, 1–4.
International Commission on Zoological Nomenclature (1999) International Code of Zoological Nomenclature. 4th Edition. International Trust for Zoological Nomenclature, London, xxix + 306 pp.
Jégu, M., Queiroz, L.J., Camacho-Terrazas, J., Teroente-Vilara, G., Carvajal-Vallejos, F.M., Pouilly, M., Yunoki, T. & Zuanon, J.A.S. (2014) Catálogo de los peces de la cuenca Iténez (Bolivia Y Brasil). In: Van Damme, P.A., Maldonado, M., Pouilly, M., Doria, C.R.C. (Eds.), Aguas del Iténez o Guaporé. Recursos Hidrobiológicos de un patrimonio binacional (Bolivia y Brasil). Editorial INIA, Cochabamba, pp. 111–156.
https://doi.org/10.4000/books.irdeditions.18741
Jerep, F.C., Carvalho, F.R. & Bertaco, V.A. (2011) Geographic distribution of Hemigrammus ora (Ostariophysi: Characiformes: Characidae) in the Amazon basin, Brazil. Zoologia, 28 (4), 545–550.
https://doi.org/10.1590/S1984-46702011000400019
Lasso, C.A., Sánchez-Duarte, P., Lasso-Alcalá, O. M., Martín, R., Samudio,H., González-Oropeza, K., Hernández-Acevedo, J. & Mesa, L. (2009) Lista de los peces del delta del río Orinoco, Venezuela. Biota Colombiana, 10 (1–2), 123–148.
Lasso, C.A., Mojica, J.I., Usma, J.S., Maldonado, J.A., do Nascimiento, C., Taphorn, D.C., Provenzano, F., Alcalá, O.M.L., Galvis, G., Vásquez, L., Lugo, M., Machado-Allison, A., Royero, R., Suárez, C. & Lara, A.O. (2014) Peces de la cuenca del Río Orinoco. Parte I: lista de especies y distribución por subcuencas. Biota Colombiana, 5 (2), 95–158.
Lima, F.C.T. & Ribeiro, A.C. (2011) Continental-scale tectonic controls of biogeography and ecology. In: Albert, J. & Reis, R.E. (Eds.), Historical biogeography of Neotropical freshwater fishes, University of California Press, Berkeley, California, pp. 145–164.
https://doi.org/10.1525/california/9780520268685.003.0009
Lima, F.C.T. & Toledo-Piza, M. (2001) New species of Moenkhausia (Characiformes: Characidae) from the rio Negro of Brazil. Copeia, 2001 (4), 1058–1063.
https://doi.org/10.1643/0045-8511(2001)001[1058:NSOMCC]2.0.CO;2
Lima, F.C.T., Malabarba, L.R., Buckup, P.A., Silva, J.F.P, Vari, R.P., Harold, A., Benine, R.C., Oyakawa, O.T., Pavanelli, C.S., Menezes, N.A., Lucena, C.A.S., Reis, R.E., Langeani, F., Casatti, L., Bertaco, V.A., Moreira, C.R. & Lucinda, P.H.F. (2003) Genera Incertae Sedis in Characidae. In: Reis, R.E., Kullander, S.O. & Ferraris Jr., C.J. (Eds.), Check List of the Freshwater fishes of South and Central America. Edipucrs, Porto Alegre, pp. 106–169.
Lima, F.C.T., Pires, T.H.S., Ohara, W.M., Jerep, F.C., Carvalho, F.R., Marinho, M.M.F. & Zuanon, J. (2013) Characidae. In: Queiroz, L.J., Torrente-Vilara, G., Ohara, W.M., Pires, T.H.S., Zuanon, J. & Doria, C.R.C. (Eds.), Peixes do rio Madeira. Vol.1. Dialeto Latin American Documentary, São Paulo, pp. 212–394.
Machado-Allison A., Chernoff, B., Royeron-León, R., Mago-Leccia, F., Velázquez, J., Lasso, C., López-Rojas, H., Bonilla-Rivero, A., Provenzano, F. & Silvera, C. (2000) Ictiofauna de la cuenca del Río Cuyuní em Venezuela. Interciencia, 25 (1), 1–21.
Machado-Alison, A. (2006) Contributions to the Knowledge of Freshwater Icthyology in Venezuela. Acta Biologica Venezuelica, 26 (1), 13 –52.
Maldonado-Ocampo, J.A., Vari, R. & Usma, J.S. (2008) Checklist of Freshwater Fishes of Colombia. Biota Colombiana, 9 (2), 143–237.
Mariguela, T.C., Benine, R.C., Abe, K.T., Avelino, G.S. & Oliveira, C. (2013) Molecular phylogeny of Moenkhausia (Characidae) inferred from mitochondrial and nuclear DNA evidence. Journal of Zoological Systematics and Evolutionary Research, 51 (4), 327–332.
https://doi.org/10.1111/jzs.12025
Marinho, M.M.F. & Langeani, F. (2010) Moenkhausia celibela: a new species from the Amazon basin, Brazil (Characiformes: Characidae). Journal of Fish Biology, 77 (4), 879–889.
https://doi.org/10.1111/j.1095-8649.2010.02719.x
Marinho, M.M.F. & Langeani, F. (2016) Reconciling more than 150 years of taxonomic confusion: the true identity of Moenkhausia lepidura, with a key to the species of the M. lepidura group (Characiformes: Characidae). Zootaxa, 4107 (3), 338–352.
https://doi.org/10.11646/zootaxa.4107.3.3
Mirande, J.M. (2019) Morphology, molecules and the phylogeny of Characidae (Teleostei, Characidae). Cladistics, 35 (3), 282–300.
https://doi.org/10.1111/cla.12345
Moreira, C.R. & Lima, F.C.T. (2017) Two new Hyphessobrycon species from Central Amazon basin, Brazil. Zootaxa, 4318 (1), 123–134.
https://doi.org/10.11646/zootaxa.4318.1.5
Ohara, W.M., Lima, F.C.T., Salvador, G.N. & Andrade, M.C. (2017) Peixes do Rio Teles Pires: diversidade e guia de identificação. Gráfica e editora Amazonas, Goiânia, 408 pp.
Ohara, W.M. & Marinho, M.M.F. (2016) A new species of Moenkhausia Eigenmann (Characiformes: Characidae) from the upper rio Machado at Chapada dos Parecis, rio Madeira basin, Brazil. Neotropical Ichthyology, 14 (1), e150041.
https://doi.org/10.1590/1982-0224-20150041
Ohara, W.M., Mirande, J.M. & Lima, F.C.T. (2017) Phycocharax rasbora, a new genus and species of Brazilian tetra (Characiformes: Characidae) from Serra do Cachimbo, rio Tapajós basin. PLoS ONE, 12 (2), e0170648.
https://doi.org/10.1371/journal.pone.0170648
Ortega, H. & Vari, R. (1986) Annotated checklist of the freshwater fishes of Peru. Smithsonian Contributions to Zoology, 437, 1–25.
https://doi.org/10.5479/si.00810282.437
Pastana, M.N.L. & Dagosta, F.C.P. (2014) Moenkhausia rubra, a new species from rio Juruena, upper rio Tapajós, basin, Brazil (Characiformes: Characidae). Neotropical Ichthyology, 12 (2), 389–396.
https://doi.org/10.1590/1982-0224-20130183
Patterson, C. (1992) Supernumerary median fin-rays in teleostean fishes. Zoological Journal of the Linnean Society, 106 (2), 147–161.
https://doi.org/10.1111/j.1096-3642.1992.tb01244.x
Pearson, N.E. (1924) The fishes of the rio Beni Basin, Bolivia, collected by Mulford Expedition. Indiana University Studies, 11, 1–83.
Pedroza, W.S., Ribeiro, F.R.V., Teixeira, T.F., Ohara, W.M. & Py-Daniel, L.H.P. (2012) Ichthyofaunal survey of stretches of the Guariba and Rooselvelt Rivers, in Guariba State Park and Guariba Extractive Reserve, Madeira River basin, Amazonas, Brazil. Check List, 8 (1), 8–15.
https://doi.org/10.15560/8.1.008
Pouilly, M., Jégu, M., Terrazas, J.C., Palacios, M.Q., Chumacero, G.M., Zubieta, J.P.Z. & Yunoki, T. (2010) Updated checklist and distribution of fish in the bolivian Amazon lowland. Revista Boliviana de Ecología y Conservacíon Ambiental, 28, 73–97.
Queiroz, L.J., Torrente-Vilara, G., Vieira, F.G., Ohara, W.M., Zuanon, J. & Doria, C.R.C. (2013) Fishes of Lago Cuniã, Madeira River Basin, Brazil. Check List, 9 (3), 540–548.
https://doi.org/10.15560/9.3.540
Ramos, T.P.A., Ramos, R.T.C. & Ramos, S.A.Q.A. (2014) Ichthyofauna of the Parnaíba river Basin, Northeastern Brazil. Biota Neotropica, 14 (1), 1–8.
https://doi.org/10.1590/S1676-06020140039
Reia, L., Vicensotto, A.M.P.F., Oliveira, C. & Benine, R.C. (2019) Taxonomy of Moenkhausia australis Eigenmann, 1908 (Characiformes, Characidae) with a discussion on its phylogenetic relationships. Zootaxa, 4688 (2), 213–231.
https://doi.org/10.11646/zootaxa.4688.2.3
Rivero, A.B. & Rojas, H.L. (2013) Biogeographic analysis of the Characiformes fishes of Venezuela. Acta biologica Colombiana, 18 (1), 69–86.
Roberts, R.T. (1967) Virilia, a new genus of sexually dimorphic characid fishes from west Africa, with remarks on characoids having an incomplete latera line. Stanford Ichthyological Bulletin, 18 (4), 251–259.
Rodrigues-Filho, C.A.S., Gurgel-Lorenço, R.C., Bezerra, L.A.V., Sousa, W.A., Garcez, D.S., Lima, S.M.Q., Ramos, T.P.A. & Sánchez-Botero, J.I. (2016) Ichthyofauna of the humid forest enclaves in the tablelands of Ibiapaba and Araripe, Northeastern Brazil. Biota Neotropica, 16 (4), e20160273.
https://doi.org/10.1590/1676-0611-BN-2016-0273
Rojas, V.F. & Rumiz, D.I. (2008) Estudio preliminar de la ictiofauna y los hábitats acuáticos del Río Bajo Paraguá, Santa Cruz, Bolivia. Biota Neotropica, 8 (1), 73–80.
https://doi.org/10.1590/S1676-06032008000100010
Sabaj, M.H. (2019) Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an online reference. Version 7.1. 21 March 2019. American Society of Ichthyologists and Herpetologists, Washington, D.C. Electronically accessible. Available from: http://www.asih.org (accessed 3 August 2020)
Sarmiento, J., Moraes, M.R., Aguirre, L.F. & Specht, R. (2016) Vertebrados de Espíritu, Llanos de Moxos: un Palmar estacionalmente inundable de Bolivia. In: Lasso, C.A., Colonnello, G. & Moraes R., M. (Eds.), XIV. Morichales, cananguchales y otros palmares inundables de Suramérica. Parte II: Colombia, Venezuela, Brasil, Perú, Bolivia, Paraguay, Uruguay y Argentina. Serie Editorial Recursos Hidrobiológicos y Pesqueros Continentales de Colombia. Instituto de Investigación de Recursos Biológicos Alexander von Humboldt (IAvH), Bogotá, D.C., pp. 346–371.
Silva, M.J., Ramos, T.P.A., Costa, B.G., Auricchio, P. & Lima, S.M.Q. (2015) Ichthyofauna of the Gurgueia River, Parnaíba River basin, northeastern Brazil. Check List, 11 (5), 1–8.
https://doi.org/10.15560/11.5.1765
Soares, I.M. & Bührnheim, C.M. (2016) A new species of Moenkhausia Eigenmann, 1903 (Characiformes: Characidae) from Amazon basin, Brazil. Zootaxa, 4208 (4), 392–400.
https://doi.org/10.11646/zootaxa.4208.4.6
Sousa, L., Netto-Ferreira, A.L. & Birindelli, J.L.O. (2010) Two new species of Moenkhausia Eigenmann (Characiformes: Characidae) from Serra do Cachimbo, Pará, Northern Brazil. Neotropical Ichtyology, 8 (2), 255–264.
https://doi.org/10.1590/S1679-62252010000200003
Taphorn, D.C. (1992) The characiform fishes of the Apure river drainage, Venezuela. Biollania edición especial, Monografias Cientificas del Museo de Ciencias Naturales, 4, 1–221.
Taylor, W.R. & Van Dyke, G.C. (1985) Revised procedures for attaining and clearing small fishes and other vertebrates for bone and cartilage study. Cybium, 9, 107–119.
Toledo-Piza, M. (Org.) (2002) Peixes do Rio Negro: Alfred Russel Wallace. Editora da Universidade de São Paulo, Imprensa Oficial do Estado, São Paulo, 517 pp.
Trujillo, F., Antelo, R. & Usma, S. (Eds.) (2016) Biodiversidad de la cuenca baja y media del Río Meta. Fundación Omacha, Fundación Palmarito, WWF, Bogotá, 336 pp.
Vari, R.P. (1989) Systematics of the neotropical characiform genus Pseudocurimata Fernández-Yépez (Pisces: Ostariophysi). Smithsonian Contributions to Zoology, 490, 1–28.
https://doi.org/10.5479/si.00810282.490
Vari, R.P. & Harold, A.S. (2001) Phylogenetic study of the neotropical fish genera Creagrutus Günther and Piabina Reinhardt (Teleostei: Ostariophysi: Characiformes), with a revision of the Cis-Andean species. Smithsonian Contributions to Zoology, 613, 1–239.
https://doi.org/10.5479/si.00810282.613
Zarske, A. & Géry, J. (2006) Zum Status einiger von Steindachner und Holly bescheriebene Fisch-Taxa (Teleostei: Characiformes: Characidae). Zoologische Abhandlungen, 56, 3–14.