SERRASALMIDAE Bleeker, 1859

Toledo-Piza, Mônica, Baena, Eduardo G., Dagosta, Fernando C. P., Menezes, Naércio A., Ândrade, Marcelo, Benini, Ricardo C., Bertaco, Vinicius A., Birindelli, José Luís O., Boden, Gert, Buckup, Paulo A., Camelier, Priscila, Carvalho, Fernando R., Castro, Ricardo M. C., Chuctaya, Junior, Decru, Eva, Derijst, Eddy, Dillman, Casey B., Ferreira, Katiane M., Merxem, Dimitri G., Giovannetti, Victor, Hirschmann, Alice, Jégu, Michel, Jerep, Fernando C., Langeani, Francisco, Lima, Flávio C. T., Lucena, Carlos A. S., Lucena, Zilda Margarete S., Malabarba, Luiz R., Malabarba, Maria Cláudia S. L., Marinho, Manoela M. F., Mathubara, Kleber, Mattox, George M. T., Melo, Bruno F., Moelants, Tuur, Moreira, Cristiano R., Musschoot, Tobias, Netto-Ferreira, André L., Ota, Rafaela P., T. Oyakawa, Osvaldo, Pavanelli, Carla S., Reis, Roberto E., Santos, Osmar, Serra, Jane Piton, Silva, Gabriel S. C., Silva-Oliveira, Cárlison, Souza-Lima, Rosana, Vari, Richard P. & Zanata, Angela M., 2024, Checklist of the species of the Order Characiformes (Teleostei: Ostariophysi), Neotropical Ichthyology (e 230086) 22 (1), pp. 1-548 : 403-417

SERRASALMIDAE Bleeker, 1859

RP Ota, M Andrade, M Jégu

Serrasalmi Swainson, 1838:256. (no family-group name)

Serrasalmoniformes Bleeker, 1859: XXXI. Type genus: Serrasalmus Lacepède, 1803 . Additional comments: Bleeker (1859) misspelled Serrasalmoniformes. Fowler (1907b:468) corrected the stem to Serrasalmi- (correct spelling Serrasalmi-).

Mylesinae Eigenmann, 1903 (correct spelling Mylei-):147. Type genus: Myleus Müller, Troschel, 1844 . Additional comments: Eigenmann (1903:147) misspelled Mylesinae. Eigenmann in Eigenmann et al. (1907:154) changed to Mylinae . Eigenmann (1907:771) corrected to Myleinae.

Catoprionidi Fowler, 1958:11. Type genus: Catoprion Müller, Troschel, 1844 .

Colossomatinae Kolmann, Hughes, Hernandez, Arcila, Betancur-R, Sabaj, López-Fernández, Ortí, 2021:Corrigendum. Type genus: Colossoma Eigenmann, Kennedy, 1903 . Additional comments: Colossomatinae not made available by Kolmann et al. (2020) or Mateussi et al. (2020), but by Corrigendum published 11 February 2021. Mateussi et al. (2020b:7) provided a morphological diagnosis.

Remarks: Two different phylogenetic hypotheses for the relationships within the Serrasalmidae have been recently proposed. Kolman et al. (2020) divided it into three subfamilies: Colossomatinae, Myleinae, and Serrasalminae . Mateussi et al. (2020) divided it into two subfamilies and two tribes (Colossomatinae ( Serrasalminae (Myleini + Serrasalmini))), and provided morphological diagnoses for each higer-level taxon.

ACNODON Eigenmann, 1903

Acnodon Eigenmann, 1903:147 . Type species: Myleus oligacanthus Müller, Troschel, 1844 . Type by original designation. Gender: Masculine. Additional comments: Monophyly proposed in Kolman et al. (2020), but their analysis did not include all congeners.

Acnodon normani Gosline, 1951

Acnodon normani Gosline, 1951:43 , Pl. 3, fig. 5b. Type locality: Rio Santa Teresa , a western tributary of the upper Rio Tocantins, State of Goiaz, Brazil. Holotype: CAS 20223.

Distribution: 322, 324 (Jégu, 2003).

Remarks: See Jégu, Santos (1990:189) for species redescription, and Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

Acnodon oligacanthus (Müller, Troschel, 1844)

Myleus oligacanthus Müller, Troschel, 1844:98 . Type locality: Surinam. Holotype: ZMB 3635. [See Zarske (2012b:328, fig. 93b, c) for holotype detailed description and x-ray image].

Distribution: 311 (Jégu, 2003).

Remarks: Also appeared as new species in Müller, Troschel (1845:24).

Acnodon senai Jégu, Santos, 1990

Acnodon senai Jégu, Santos, 1990:195 , fig. 6D. Type locality: Plage à la pointe aval de l’ile de Carapatinho, rio Jari , Amapá, Brésil (0°35’45”S, 52°38’20”W). [ Brazil]. Holotype: INPA 2680 .

Distribution: 315 (Jégu, 2003).

Remarks: See Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

CATOPRION Müller, Troschel, 1844

Catoprion Müller, Troschel, 1844:96 . Type species: Serrasalmus mento Cuvier, 1819 . Type by monotypy. Gender: Masculine.

Remarks: Recovered as monophyletic and sister to Pygopristis Müller, Troschel, 1844 (Kolmann et al., 2020; Mateussi et al., 2020b).

Catoprion absconditus Mateussi, Melo, Oliveira, 2020

Catoprion absconditus Mateussi, Melo, Oliveira, 2020:12 , figs. 8–12. Type locality: Caracaraí , igarapé do Campo , rio Jufari, rio Branco, 01°03’54”S, 62°07’46”W. [ Brazil]. Holotype: MZUSP 125764 .

Distribution: 310, 314, 315, 316 (Mateussi et al., 2020a).

Catoprion mento (Cuvier, 1819)

Serrasalmus mento Cuvier, 1819:369 , Pl. 28 (fig. 3). Type locality: Venu de Lisbonne , paroît aussi originaire du Brésil. [Likely from upper Paraguay, Araguaia or tributaries of the Amazon basin (Mateussi et al., 2020a:678)]. Holotype: MNHN A-9869 (poor condition). [Mateussi et al., 2020a]

Mylesinus macropterus Ulrey, 1894:612 . Type locality: Brazil. Holotype: CUMV 3267. [Mateussi et al., 2020a:678]. Species description supplemented by Ulrey (1895:296). Status of species as synonym corroborated by Mateussi et al., 2020a.

Distribution: 307, 308, 316, 318, 319, 321, 324, 343 (Jégu, 2003; Ferreira et al., 2011; Ota et al., 2013; Mateussi et al., 2020a).

Remarks: Species redescribed by Mateussi et al. (2020a), including molecular data.

COLOSSOMA Eigenmann, Kennedy, 1903

Colossoma Eigenmann, Kennedy, 1903:530 . Type species: Myletes oculus Cope, 1872 . Type by original designation. Gender: Neuter. Additional comments: Also appeared in Eigenmann 1903:148.

Waiteina Fowler, 1907:473 . Type species: Myletes nigripinnis Cope, 1878 . Type by original designation. Gender: Feminine. Additional comments: Originally proposed as a subgenus of Colossoma .

Melloina Amaral Campos, 1946:219 . Type species: Melloina tambaqui Amaral Campos, 1946 . Type by original designation. Gender: Feminine.

Colossoma macropomum (Cuvier, 1816)

Myletes macropomus Cuvier, 1816:185 , Pl. 10 (fig. 1). Type locality: Brasil. Holotype: MNHN A.8626. [Stuffed specimen].

Myletes oculus Cope, 1872b:268 , Pl. 12 (fig. 2). Type locality: Ambyiacu. [ Río Ampyiacu , Peru]. Lectotype: ANSP 7991 . [Designated by Fowler (1907:472). Treated as holotype by Böhlke (1984:53)]. See Taphorn (1992:161) for synonymy with Colossoma macropomum (Cuvier, 1816) .

Myletes nigripinnis Cope, 1878:693 . Type locality: Probably from Nauta, Peruvian Amazon. Lectotype: ANSP 21124 . [Designated by Fowler (1907:473, fig. 55). Treated as holotype by Böhlke (1984:51)]. See Taphorn (1992:161) for synonymy with Colossoma macropomum (Cuvier, 1816) .

Melloina tambaqui Amaral Campos, 1946:220 , fig. 3. Type locality: Taperinha , Estado do Pará (Santarém). [ Brazil]. Holotype: MPEG.

Distribution: 203i, 206i, 207i, 208i, 209i, 210i, 301, 302, 305, 306, 307, 308, 315, 316, 317, 318, 319, 320, 321, 322, 323, 324, 325, 327, 330, 342, 344 (Jégu, 2003. For information about introduction of species see Castellani, Barrella, 2006; Barriga, 2012; Ramos et al., 2014; Langeani et al., 2007 and Ota et al., 2018).

Remarks: Also described as new in Cuvier (1818:453, Pl. 21, fig. 3). Detailed description of ontogeny, neurocranium osteology, and morphology in Machado-Allison (1982, 1986) and Géry (1986). See Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

METYNNIS Cope, 1878

Metynnis Cope, 1878:692 . Type species: Metynnis luna Cope, 1878 . Type by monotypy. Gender: Masculine.

Myleocollops Eigenmann, 1903:147 . Type species: Metynnis goeldii Eigenmann,

1903. Type by original designation. Gender: Masculine. Additional comments: Originally proposed as a subgenus of Metynnis .

Sealeina Fowler, 1907:478 . Type species: Myletes lippincottianus Cope, 1870 . Type by original designation. Gender: Feminine.

Metynnis altidorsalis Ahl, 1923

Metynnis altidorsalis Ahl, 1923b:22 . Type locality: Paramaribo, Surinam. Holotype: ZMB 19626 . [See Zarske (2012b:299, fig. 74a, b) for holotype detailed description and X-ray image].

Distribution: 307, 308, 309, 310, 311, 314, 315, 316, 317, 320 (Jégu, 2003; Jégu, Ingenito, 2007; Ota, 2015; Machado et al. (2018)).).

Remarks: See Zarske, Géry (2000:17–22) for variation in color pattern and body shape, and Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

Metynnis anisurus Ahl, 1923

Metynnis anisurus Ahl, 1923b:27 , fig. 2. Type locality: Rio Tapajoz , Borin. [Rio Tapajós, Boim, Brazil]. Holotype: ZMB 20678 . [See Zarske (2012b:300, fig. 75b, c) for holotype detailed description and X-ray image].

Distribution: 315, 320, 322 (Ota, 2015:189) and Ota (2015) based on color pattern.

Metynnis argenteus Ahl, 1923

Metynnis argenteus Ahl, 1923b:24 . Type locality: Rio Tapajoz , Borin. [Rio Tapajós, Boim, Brazil]. Holotype: ZMB 20676 . [See Zarske (2012b:302, fi. 76a, b) for holotype detailed description and X-ray image].

Metynnis eigenmanni Ahl, 1923b:25 . Type locality: Rio Tapajoz, Borin. [Rio Tapajós, Boim, Brazil]. Holotype: ZMB 20677. [See Zarske (2012b:310, fig. 81a, b) for holotype detailed description and X-ray image]. See Gosline (1951:52) for synonymy with Metynnis argenteus Ahl, 1923 . According to Ota (2015:58) Metynnis eigenmanni Ahl, 1923 is a junior synonym of Metynnis altidorsalis Ahl, 1923 .

Distribution: 320 (Jégu, 2003; Jégu, Ingenito, 2007).

Remarks: Metynnis argenteus was considered a junior synonym of Metynnis altidorsalis Ahl, 1923 by Gosline (1951:52) and Ota (2015:58), based on similar morphology and color pattern.

Metynnis cuiaba Pavanelli, Ota, Petry, 2009

Metynnis cuiaba Pavanelli, Ota, Petry, 2009:142 , figs. 1–5. Type locality: Brazil, Mato Grosso State, Município de Barão de Melgaço and Santo Antônio do Leverger, baía Sinhá Mariana , rio Cuiabá , rio Paraguai basin, 16°20’20”S, 55°54’10”W. Holotype: MZUSP 100626 .

Distribution: 318, 319, 321, 323, 324, 343 (Pavanelli et al., 2009; Ota, 2015; Machado et al. (2018)).

Remarks: See Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

Metynnis fasciatus Ahl, 1931

Metynnis fasciatus Ahl, 1931b:407 , Pl. 3a, fig. unnumbered p. 409. Type locality: Rio Capiuru. [correct spelling of Capiuru is “Capim”. According to Géry (1964d:461) the type locality is probably in the lower Amazonas River basin, Brazil]. Holotype: ZMB (not found). [In original description Ahl (1931:409) mentioned the type specimen was 102 mm SL].

Distribution: 316, 320, 322, 323, 324 (Jégu, 2003; Ota, 2015; Machado et al. (2018)).

Remarks: See Géry (1964d:461 and plate III, fig. a) for a diagnosis from Metynnis hypsauchen (Müller, Troschel, 1844) , and color pattern. See Zarske (2012b:311) for a discussion on taxonomic status of the species, and Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

Metynnis guaporensis Eigenmann, 1915

Metynnis guaporensis Eigenmann, 1915:267 , Pl. 54. Type locality: Maciél , Rio Guaporé. [ Brazil]. Holotype: FMNH 56913 .

Distribution: 307, 308, 310, 314, 315, 316, 318, 319, 321, 322, 323, 324 (Jégu, 2003; Ota et al., 2013; Ota, 2015; Machado et al. (2018)).

Remarks: See Zarske, Géry (1999b:193) for species redescription, Ota et al. (2013:19) for information about sexual dimorphism and color pattern and Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

Metynnis hypsauchen (Müller, Troschel, 1844)

Myletes hypsauchen Müller, Troschel, 1844:97 . Type locality: Guiana. [ Guyana]. Lectotype: ZMH 2865. [Designated and figured by Zarske, Géry (1999b:172, fig. 1). Zarske (2012b: 322, fig. 89b. c) for paralectotype detailed description and X-ray image].

Metynnis calichromus schreitmuelleri Ahl, in Schreitmüller, Ahl, 1922:265. Type locality: Amazonenstrom. Holotype: ZMB 20664 . [See Zarske (2012b:305, fig. 78b, c) for holotype detailed description and X-ray image]. Also appeared in Ahl (1923:19).

Metynnis calichromus Ahl, 1923b:18 , fig. 1. Type locality: Lago de Faro (Jamundá). [“Lago do Faro (Nhamundá)”, Brazil]. Holotype: ZMB 20674. [See Zarske (2012b:304, fig. 77b, c) for holotype detailed description and X-ray image]. See Zarske, Géry (1999b:197) for description of type and synonymy with Metynnis hypsauchen (Müller, Troschel, 1844) .

Metynnis ehrhardti Ahl, 1927:273 . Type locality: Mundurucú, Rio Manacapuru, Solimoes, Amazonas, Brazil. Lectotype: ZMB 23685. [Designated and figured by Zarske, Géry (1999b:2010, fig. 33). See Zarske (2012b:309, fig. 80a, b) for lectotype detailed description and X-ray image]. See Zarske, Géry (1999b:211) for types description and synonymy with Metynnis hypsauchen (Müller, Troschel, 1844) .

Distribution: 307, 308, 310, 314, 315, 316, 320 (Jégu, 2003; Ota, 2015; Ota et al. 2016; Machado et al. (2018)).

Remarks: Also appeared as new species in Müller, Troschel (1845:23). See Zarske, Géry (1999b:173) for information about sexual dimorphism, Ota et al. (2016) for a discussion of distribution, and Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

Metynnis lippincottianus (Cope, 1870)

Myletes lippincottianus Cope, 1870:561 , figure of head unnumbered. Type locality: Para, Brazil, J. C. Brevoort’s collection. Syntype: ANSP 8024 . [Bohlke (1984:49); photograph of specimen and vertebral counts presented by Zarske, Géry (1999:180, fig. 8); type status based on a label identifying the specimen as a syntype.].

Metynnis goeldii Eigenmann, 1903:147 . Type locality: South America. [Lectotype locality: Brazil ( Zarske, Géry, 2008)]. Lectotype: CAS 70686 [ex. SU 5205]. [Designated and figured by Zarske, Géry (2008:193, figs. 20–21)]. See Zarske, Géry (2008:193) for a discussion of the species taxonomic status.

Myletes (Myleus) orbicularis Steindachner, 1908b:364 . Type locality: Santarem am Amazonenstrom. [The correct is “Santarém”, Brazil]. Lectotype: NMW 56428. [Designated and figured by Zarske, Géry (1999b:189, fig. 17)]. See Zarske, Géry (1999b:189) for a discussion of the species taxonomic status.

Metynnis roosevelti Eigenmann, 1915:268 , Pl. 55. Type locality: Manaos. [ Brazil]. Holotype: FMNH 56925 . See Zarske, Géry (1999b:194) for type description and discussion of the species taxonomic status.

Metynnis heinrothi Ahl, 1923b:29 , fig. 3. Type locality: Amazonenstrom? Aquarium, Zoo, Berlin. Holotype: ZMB 20662. [See Zarske (2012b:312, fig. 83b, c) for holotype detailed description and X-ray image]. See Zarske, Géry (1999b:208) for a discussion of the species taxonomic status.

Metynnis seitzi Ahl, 1923b:28 . Type locality: Amazonas? Aquarium, Zoo, Berlin. Holotype: ZMB 20663. [See Zarske (2012b:317, fig. 86b, c) for holotype detailed description and X-ray image]. See Zarske, Géry (1999b:207) for a discussion of the species taxonomic status.

Metynnis snethlageae Ahl, 1923b:30 . Type locality: Amazonenstrom? Aquarium, Zoo, Berlin. Holotype: ZMB 19722. [See Zarske (2012b:319, fig. 87b, c) for holotype detailed description and X-ray image]. See Zarske, Géry (1999b:210) for a discussion of the species taxonomic status.

Metynnis dungerni Ahl, 1925:108 . Type locality: Amazonenstrom bei Para. [ Brazil]. Holotype: ZMB 23683. [See Zarske (2012b:307, fig. 79b, c) for detailed description of holotype and X-ray image]. See Zarske, Géry (1999b:210) for a discussion of the species taxonomic status.

Distribution: 307, 311, 315, 316, 318, 319, 321, 323, 324, 325, 326, 327, 328, 329, 344i (Jégu, 2003; Jégu, Ingenito, 2007; Ota et al., 2013; Ota, 2015; Ota et al., 2016; Machado et al., 2018).

Remarks: See Zarske, Géry (1999b) for discussion on synonymies, Ota et al. (2013:19) for information about sexual dimorphism and color pattern and Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI). Species currently misidentified as M. maculatus (Kner, 1858b) in Paraná and São Francisco River basins (e.g., Langeani et al., 2007).

Metynnis longipinnis Zarske, Géry, 2008

Metynnis longipinnis Zarske, Géry, 2008:175 , figs. 1–2. Type locality: Venezuela, Mündung rio Casiquiare -- rio Negro. Holotype: SU 54688 :1. [According to Fricke et al. (2022) the holotype ( CAS-SU 54688 :1, 66.5 mm SL) is not identified because of similar size to the paratypes ( CAS-SU 54688 :2, 66.4 mm SL)].

Distribution: 306, 308, 314, 315, 316 ( Zarske, Géry, 2008; Ota, 2015).

Remarks: See Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

Metynnis luna Cope, 1878

Metynnis luna Cope, 1878:692 . Type locality: Near Pebas, Peruvian Amazon. Holotype: ANSP 21443. [Fowler (1907:479, fig. 58); Böhlke(1984:50)].

Distribution: 308, 310, 314, 315, 316, 317, 319, 321, 322, 324 (Jégu, 2003; Ota et al., 2013; Ota, 2015).

Remarks: See Zarske, Géry (1999b:181) for species redescription, Ota et al. (2013:21) for information about sexual dimorphism and color pattern. Hypothesis of species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI) proposed that this species is represented by three or more lineages (Machado et al., 2018).

Metynnis maculatus (Kner, 1858)

Myletes maculatus Kner, 1858b:164 . Type locality: not given. [Kner (1858b) mentioned the species co-occurs with M. hypsauchen . Kner (1860:27) later mentioned the type locality as “ Rio Guaporé ”]. Lectotype: NMW 56396 . [Designated and figured by Zarske, Géry (1999b:178, fig. 5)].

Distribution: 307, 308, 314, 315, 316, 318, 319, 321, 343 (Jégu, 2003; Ota et al., 2013; Ota, 2015).

Remarks: Described as new species in Kner (1860:26). See Ota et al. (2013:21) for information about sexual dimorphism and color pattern, and Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

Metynnis melanogrammus Ota, Rapp Py-Daniel, Jégu, 2016

Metynnis melanogrammus Ota, Rapp Py-Daniel, Jégu, 2016:804 , figs. 1–8. Type locality: rio Parauari , Monte Sinai community, Maués, Amazonas, Brazil, 4°21’28”S, 57°35’54”W. Holotype: INPA 52216 .

Distribution: 314, 315, 316, 321 (Ota et al., 2016).

Remarks: See Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

Metynnis mola Eigenmann, Kennedy, 1903

Metynnis mola Eigenmann, Kennedy, 1903:528 . Type locality: Arroyo Trementina. [ Paraguay]. Holotype: CAS 60784. [ According to Zarske (2012b:314), the lot ZMB 16978 , 71.3 mm SL, is not a paratype].

Distribution: 343 (Jégu, 2003).

Remarks: See Zarske, Géry (1999b:187, fig. 15b) for information about postpelvic spines, Zarske (2012b:314) for detailed description and X-ray image, and Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

Metynnis orinocensis (Steindachner, 1908)

Myletes (Myleus) orinocensis Steindachner, 1908b:365 . Type locality: Orinoco bei Ciudad Bolivar. [ Venezuela]. Syntypes: (3) NMW (apparently lost). [Syntypes probably lost according to Jégu (2003)].

Distribution: 308 (Jégu, 2003).

Remarks: See Zarske, Géry (1999b:190, fig. 18) for species redescription. Status of species discussed by Ota (2015:144) as a possible junior synonym of M. hypsauchen .

Metynnis otuquensis Ahl, 1923

Metynnis otuquensis Ahl, 1923b:26 . Type locality: Rio Otuquis, Bahia Negra. Holotype: ZMB 16980, and paratype ZMB 34505. [See Zarske (2012b:315, fig. 85a, b) for holotype detailed description and X-ray image. Zarske, Géry (1999b:202) separated the holotye (ZMB 16980, 95.6 mm SL), from the paratype (ZMB 34505, 87.0 mm SL)].

Distribution: 342, 343 (Jégu, 2003; Ota, 2015).

Remarks: See Zarske, Géry (1999b:203) for a comparison with Metynnis mola Eigenmann, Kennedy, 1903 .

Metynnis polystictus Zarske, Géry, 2008

Metynnis polystictus Zarske, Géry, 2008:184 , figs. 3–4. Type locality: Lagune en bordure du Ribeiro Tuatuari, en aval de Posto Leonardo, haut Xingú. [ upper rio Xingu , Brazil]. Holotype: IRSNB 833 .

Distribution: 320, 322 ( Zarske, Géry, 2008).

Remarks: Status of species discussed by Ota (2015:76) as a possible junior synonym of M. anisurus . See Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

MYLESINUS Valenciennes, 1850

Mylesinus Valenciennes , in Cuvier, Valenciennes, 1850:234 . Type species: Mylesinus schomburgkii Valenciennes, 1850 . Type by monotypy. Gender: Masculine.

Remarks: Recovered as paraphyletic in several molecular phylogenies (e.g., Machado et al., 2018; Mateussi et al., 2020b). Morphological diagnosis based on teeth arrangement is homoplastic and more comprehensive studies are required to understand its composition and relationships with remaining serrasalmid genera.

Mylesinus paraschomburgkii Jégu, Santos, Ferreira, 1989

Mylesinus paraschomburgkii Jégu, Santos, Ferreira, 1989:53 , Pl. 1. Type locality: Rio Trombetas , en aval de Cachoeira Vira-Mundo, PA. [ Brazil]. Holotype: INPA 1226 .

Distribution: 311, 315, 323* (Jégu, 2003; Vari, Ferraris, 2009 ; Andrade et al., 2013).

Remarks: See Andrade et al. (2013:304) for identification key. Hypothesis of species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI) proposed that this species is represented by three or more lineages (Machado et al., 2018). A taxonomic review is necessary to understand its intra- and interspecific variation.

Mylesinus paucisquamatus Jégu, Santos, 1988

Mylesinus paucisquamatus Jégu, Santos, 1988:332 , Pl. 1. Type locality: Jatobal , rio Tocantins. [ Brazil]. Holotype: INPA 1808 .

Distribution: 324 (Jégu, 2003; Andrade et al., 2013).

Remarks: See Andrade et al. (2013:304) for identification key, and Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

Mylesinus schomburgkii Valenciennes, 1850

Mylesinus schomburgkii Valenciennes , in Cuvier, Valenciennes, 1850:235 , Pl. 644. Type locality: not given. [accoding to Bertin (1948:27–28) the type locality is Essequibo River, Guyana]. Holotype: MNHN A.9855. [Incomplete specimen, head only, used for original description along with the drawing depicted in Pl.

644 in Cuvier, Valenciennes (1850) ]. Distribution: 308, 310 (Jégu, 2003; Andrade et al., 2013). Remarks: See Andrade et al. (2013:304) for identification key.

MYLEUS Müller, Troschel, 1844

Myleus Müller, Troschel, 1844:98 . Type species: Myleus setiger Müller, Troschel, 1844 . Type by subsequent designation by Eigenmann (1910:443). Gender: Masculine.

Remarks: Recovered as monophyletic in Kolmann et al. (2020), but paraphyletic in Mateussi et al. (2020b). Morphological diagnosis based on teeth arrangement is homoplastic and more comprehensive studies are required to understand its composition and relationship with remaining serrasalmid genera.

Myleus altipinnis ( Valenciennes, 1850)

Tometes altipinnis Valenciennes , in Cuvier, Valenciennes, 1850:230 , Pl. 643. Type locality: Rio San-Francisco. [rio São Francisco, Brazil]. Holotype: MNHN A-8652. [stuffed specimen, lacking axial skeleton].

Distribution: 327 (Jégu, 2003; Jégu, Ingenito, 2007).

Remarks: Species only known from the stuffed holotype. See Andrade et al. (2018) for description of teeth arrangement.

Myleus knerii (Steindachner, 1881)

Myletes knerii Steindachner, 1881a:127 [27], Pl. 7 (fig. 2). Type locality: Maroni- Fluss in Guiana [French Guiana]. Holotype: MNW 56394. [See Jégu, Santos (2002:55, fig 13a, b, c) for holotype images].

Distribution: 311 (Jégu, 2003).

Remarks: See Jégu, Santos (2002:54) for detailed redescription, Andrade et al. (2013:304) for identification key, and Andrade et al. (2018) for description of teeth arrangement.

Myleus micans (Lütken, 1875)

Myletes (Tometes) micans Rhdt. Lütken, 1875c:137 [11]. Type locality: Rio das Velhas og dens niilod Rio Taquoaruru , der falder i forst naevnte lidt nord for Lagoa Santa. [ Brazil]. ZMUC 260 (1) and ZMUC 261 (1).

Distribution: 327 (Jégu, 2003).

Remarks:Also described in Lütken (1875:241). See Andrade et al. (2018) for description of teeth arrangement, and Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

Myleus pachyodus Andrade, Jégu, Buckup, Netto-Ferreira, 2018

Myleus pachyodus Andrade, Jégu, Buckup, Netto-Ferreira, 2018:1903 , figs. 1–7. Type locality: Brazil, Mato Grosso, Alta Floresta, Tapajós River basin, Rio Teles Pires , 9°38’01”S, 55°56’01”W. Holotype: MNRJ 35023 .

Distribution: 320 (Andrade, Jégu, Buckup, Netto-Ferreira, 2018).

Myleus planquettei (Jégu, Keith, Le Bail, 2003)

Myloplus planquettei Jégu, Keith, Le Bail, 2003:835 , figs. 2–5. Type locality: fleuve Maroni, Twenk, Guyane française, coll. De Mérona [ French Guiana]. Holotype: MNHN 1997-730 .

Distribution: 310, 311 (Jégu et al., 2003; Vari, Ferraris, 2009 ; Mol et al., 2012).

Remarks: Assignment to Myelus based on Kolmann et al., 2020:11, however the species does not share the morphological diagnosis proposed by Jégu, Santos (2002).

Myleus setiger Müller, Troschel, 1844

Myleus setiger Müller, Troschel, 1844:98 . Type locality: Guiana. Lectotype: ZMB 3634 . [Designated and figured by Jégu, Santos (2002:36), fig 1. See Zarske (2012b:330, fig. 94b) for lectotype and paralectotype photos, and X-ray image].

Myletes divaricatus Valenciennes , in Cuvier, Valenciennes, 1850:215 . Type locality: Esséquibo. [ Guyana]. Holotype: MNHN A-8629. [See Jégu, Santos (2002:51, fig. 10a) for holotype image]. See Jégu, Santos (2002:37) for the species taxonomic status discussion.

Myletes doidyxodon Valenciennes , in Cuvier, Valenciennes, 1850:222 . Type locality: Amazone. Holotype: MNHN A-9868. [See Jégu, Santos (2002:51, fig. 10b) for holotype image]. See Jégu, Santos (2002:37) for the species taxonomic status discussion.

Distribution: 307, 308, 310, 311, 314, 315, 319, 320, 321, 322, 324 (Jégu, 2003; Andrade et al., 2013; Vari, Ferraris, 2009 ).

Remarks: Also appeared as new in Müller, Troschel (1845:24). Detailed redescription in Jégu, Santos (2002:37). Identification key in Andrade et al. (2013). See Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

Species inquirenda ( Myleus )

Myletes pacu Jardine , in Schomburgk, 1841:236, Pls. 20–21. Type locality: Falls of the Essequibo and Mazaruni, at the cataracts of the Paramu, a tributary of the Orinoco which joins that the river above Esmeralda. [ Guyana]. No types known. Remarks: Jégu (2003) considered Myleus pacu as species inquirenda based on Jégu, Santos (2002) who considered the types of Myleus pacu as unknown. See Andrade et al., 2019 for taxonomic history.

MYLOPLUS Gill, 1896

Myloplus Gill, 1896c:214 . Type species: Myletes asterias Müller, Troschel, 1844 . Type by subsequent designation by Jordan (1920:467). Gender: Masculine. Additional comments: Originally proposed as subgenus of Myleus .

Orthomyleus Eigenmann, 1903:148 . Type species: Myletes ellipticus Günther, 1864 . Type by original designation. Gender: Masculine. Additional comments: Originally proposed as a subgenus of Myleus .

Remarks: Recovered as paraphyletic in recent molecular phylogenies (Kolmann et al., 2020; Mateussi et al., 2020b). Morphological diagnosis based on teeth arrangement is homoplastic and more comprehensive studies are required to understand its composition and relationship with remaining serrasalmid genera.

Myloplus arnoldi Ahl, 1936

Myloplus arnoldi Ahl, 1936:25 . Type locality: Amazonenstrom. Syntype: 20812. [ Zarske (2012b) revealed that just one specimen is available as syntype rather than two. See Zarske (2012b:331, fig. 95b, c) for detailed description and X-ray image of syntype].

Distribution: 315*, 316, 322, 323*, 324 (Jégu, 2003; Jégu, Ingenito, 2007; M. Ândrade, pers. obs.).

Remarks: See Zarske (2012b:329) for new combination and Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

Myloplus asterias (Müller, Troschel, 1844)

Myletes asterias Müller, Troschel, 1844:98 . Type locality: Guiana. [ Guyana]. Lectotype: ZMB 3646 . [Designated by Jégu et al. (2004:124). See Zarske (2012b:321, fig. 88b, c) for lectotype detailed description and X-ray image.

Myletes ellipticus Günther, 1864:375 . Type locality: Essequibo. [ Guyana]. Syntypes: BMNH 1864.1 .21.33 (1), 1971.5.10.63 (1). [See Jégu et al. (2004:125–126, fig. 4a) for syntype image and detalied description]. See Jégu et al. (2004:127) for discussion on taxonomic status.

Myleus gurupyensis Steindachner, 1911b:342 . Type locality: Rio Gurupy bei Chatão. [ Brazil]. Syntypes: (2) NMW 10589 (1). [Jégu (2003:185) mentioned that only one specimen was found rather than two]. See Jégu (2003:185) for synonymy with Myloplus asterias (Muller, Troschel) based on personal observation and examination of type specimens.

Myloplus schulzei Ahl, 1938:191 . Type locality: South America. Holotype: ZMB 20814 . [See Jégu (2001:385, fig. 1), and Zarske (2012b:332, fig. 96a, b) for holotype detailed description and X-ray image]. See Jégu (2001:383) for discussion on taxonomic status.

Tomète maculatus Amaral Campos, 1944b:211 , figure. Type locality: Rio Amazonas. Holotype: MZUSP 3356. Genus should have been spelled Tometes . See Jégu (2003:185) for synonymy with Myloplus asterias (Müller, Troschel, 1844) .

Distribution: 310, 311, 314, 315, 316, 319, 321, 322, 323, 324 (Jégu, 2003; Jégu et al., 2004; Ota et al., 2013).

Remarks: Also appeared as new in Müller, Troschel (1845:24). See Jégu et al. (2004:123) for species redescription. Multiple lineages recovered in the hypothesis of species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI) proposed in Machado et al. (2018), and Ota et al. (2020). A broad taxonomic review is necessary to understand intra- and interspecific vatiarion, and consequently the limits of its geographic distribution. See Ota et al. (2020) for confirmation of synonymies.

Myloplus levis (Eigenmann, McAtee, 1907)

Myleus levis Eigenmann, McAtee , in Eigenmann , McAtee , Ward, 1907:142, Pl. 42 (fig. 2). Type locality: Bahia Negra [ Paraguay]. Holotype: CAS 62107.

Distribution: 342, 343 (Britski et al., 1999; Gomes, 2015).

Remarks: See Britski et al. (1999:58) for a short description, Andrade et al. (2016b:154) for new combination, and Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

Myloplus lobatus ( Valenciennes, 1850)

Myletes lobatus Valenciennes , in Cuvier, Valenciennes, 1850:212 . Type locality: Amazone. Syntype: MNHN 0000-5244. [unique, in poor condition].

Myletes discoideus Kner, 1860:30 [22]. Type locality: Bananeira , Rio branco, Matogrosso. [Mato Grosso, Brazil]. Syntypes: NMW 16407 (1, dry). The name first appeared in Kner (1858b:165), but was described in more detail by Kner (1860:30) who mentioned it was already being described by Heckel (in preparation).

Myletes parma Günther, 1864:374 . Type locality: River Capin , Para. [ Brazil]. Syntypes: BMNH 1849.11 .8.32 (1) and BMNH 1849.11 .8.33 (1). See Jégu (2003:185) for discussion on the species taxonomic status.

Distribution: 307, 308, 309, 315, 316, 318, 319, 321 (Jégu, 2003).

Remarks: See Jégu, Ingenito (2007) for new combination, Ota et al. (2013:25) for information about diagnostic features and color pattern, and Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

Myloplus lucienae Andrade, Ota, Bastos, Jégu, 2016

Myloplus lucienae Andrade, Ota, Bastos, Jégu, 2016:572 , figs. 1–4. Type locality: Rio Carabinani , Novo Airão, Amazonas, Brazil, 2°01’25”S, 61°32’35”W. Holotype: INPA 30717 .

Distribution: 314 (Andrade et al., 2016c).

Remarks: See Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

Myloplus nigrolineatus Ota, Machado, Andrade, Collins, Farias, Hrbek, 2020

Myloplus nigrolineatus Ota, Machado, Andrade, Collins, Farias, Hrbek, 2020 : figs. 1–4. Type locality: Brazil, Amazonas State , ca. 60 km south of Manaus , lago Tracajá , tributary of rio Juma, rio Amazonas basin, 3°41’15”S, 59°57’01”W. Holotype: INPA 54773 .

Distribution: 314, 316, 320, 324 (Ota et al., 2020).

Remarks: See Machado et al. (2018) for previous results of species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

Myloplus rubripinnis (Müller, Troschel, 1844)

Myletes rubripinnis Müller, Troschel, 1844:97 . Type locality: Guiana. Lectotype: ZMB 23686 . [Designated by Jégu et al. (2004:125). See Zarkse (2012:325, fig. 91b, c) for paralectotype detailed description and X-ray image. Paralectotype also illustrated in Andrade et al. 2018:118, fig. 4].

Myletes luna Valenciennes , in Cuvier, Valenciennes, 1850:221 . Type locality: Cayenne. [ French Guiana]. Lectotype: MNHN A-9870 (290 mm SL). [Designated by Géry (1972a:163). See Jégu et al. (2004:125–126, fig. 4b) for discussion about species taxonomic status].

Distribution: 307, 308, 310, 311, 314, 315, 316, 319, 320*, 321, 322, 323, 324 (Jégu, 2003; Jégu et al., 2004; Ota et al., 2013; M. Andrade, pers. obs.).

Remarks: Also appeared as new in Müller, Troschel (1845:23). See Jégu et al. (2004:140) for detailed redescription, information about color pattern. Multiple lineages recovered in the species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI) in Machado et al. (2018). Andrade et al., 2018 presented comments on distribution of the species in the Guiana Shield, but highlighted the necessity of a broad taxonomic review to delimit its geographical distribution.

Myloplus schomburgkii (Jardine, 1841)

Tetragonopterus schomburgkii Jardine , in Schomburgk, 1841:243, Pl. 22. Type locality: Rio Negro. No types known.

Myletes palometa Valenciennes , in Cuvier, Valenciennes, 1850:214 . Type locality: Haut Orénoque. [ Venezuela]. No types known. In the original description the author mentioned this species as strictly related to Myloplus schomburgkii (Jardine 1841) , but because it came from a different river basin, it was described as a new species.

Myletes schomburgkii Valenciennes , in Cuvier, Valenciennes, 1850:212 . Type locality: Surinam. [ Suriname]. Probable Syntypes: ZMB 3638 (1) and ZMB 3639 (1). [No types known according to Jégu (2003). See Zarske (2012b:327, fig. 92a, b) for the probable syntypes description].

Distribution: 307, 308, 313, 314, 315, 316, 320*, 321, 322, 324 (Jégu, 2003; Jégu, Ingenito, 2007; Machado et al., 2018; M. Andrade, pers. obs.).

Remarks: See Andrade et al. (2016b:154) for new combination. Multiple lineages recovered in the hypothesis of species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI) in Machado et al. (2018).

Myloplus tiete (Eigenmann, Norris, 1900)

Myletes tieté Eigenmann, Norris, 1900:359 . Type locality: Piracicaba. [ Brazil]. Holotype: not found.

Distribution: 343, 344, 345 (Jégu, 2003; Jégu, Ingenito, 2007).

Remarks: See Gomes (2015:104) for color pattern and information about sexual dimorphism, and Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

Myloplus torquatus (Kner, 1858)

Myletes torquatus Kner, 1858b:164 . Type locality: not given. [Kner (1860:24) later mentioned type locality as “ Rio branco ( Marabitanos )”]. Syntypes: NMW 56449 (1) and NMW 56450 (1). [According to Jégu (2003) only two specimens are available as syntypes rather than three. See Gomes (2015:107, fig. 54) for syntype illustration].

Distribution: 308, 314 (Jégu, 2003; Jégu, Ingenito, 2007).

Remarks: Described as new species in Kner (1860:24, Taf I fig. 4). See Andrade et al. (2016b:154) for new combination.

Myloplus tumukumak Andrade, Jégu, Gama, 2018

Myloplus tumukumak Andrade, Jégu, Gama, 2018:112 , figs. 1–2, 4B. Type locality: Brazil, State of Amapá, Motoura River, a tributary of Oyapock River , 02°35’00”N, 52°33’00”W. Holotype: ZMA 107.638 .

Distribution: 311, 315 (Andrade, Jégu, Gama, 2018).

Myloplus zorroi Andrade, Jégu, Giarrizzo, 2016

Myloplus zorroi Andrade, Jégu, Giarrizzo, 2016b:155 , figs. 1, 2 and 4. Type locality: Rio Aripuanã , Corredeira dos Periquitos, Apuí, Amazonas, Brazil, 07°17’19.8”S, 60°38’10.0”W. Holotype: INPA 50880 .

Distribution: 321 (Andrade et al., 2016b).

Remarks: See Machado et al. (2018) for species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI).

MYLOSSOMA Eigenmann, Kennedy, 1903

Mylossoma Eigenmann, Kennedy, 1903:530 . Type species: Myletes albiscopus Cope, 1872 . Type by monotypy. Gender: Neuter. Additional comments: Also appeared in Eigenmann (1903:148).

Starksina Fowler, 1907:476 . Type species: Myletes herniarius Cope, 1872 . Type by original designation (also monotypic). Gender: Feminine.

Remarks: Revision of Cis-Andean species by Mateussi et al. (2018).

Mylossoma acanthogaster ( Valenciennes, 1850)

Myletes acanthogaster Valenciennes , in Cuvier, Valenciennes, 1850:209 . Type locality: Lagune de Maracaïbo. [ Venezuela]. Holotype: MNHN A-1065.

Distribution: 303 (Jégu, 2003).

Remarks: See Machado-Allison, Castillo (1992) for taxonomic review.

Mylossoma albiscopum (Cope, 1872)

Myletes albiscopus Cope, 1872b:267 , Pl. 12 (fig. 1). Type locality: Ambyiacu. [ Río Ampyiacu , Peru]. Lectotype: ANSP 8021 . [Designated by Fowler (1907:475). Treated as holotype by Böhlke (1984:42). See Mateussi et al., 2018:282, fig.4 for lectotype image].

Mylossoma argenteum Ahl, 1928c:192 , figure. Type locality: Amazonenstrom. Syntypes: ZMB 20854 (2). [See Zarske (2012b:334, fig. 97b, c) for holotype detailed description and X-ray image. Also appeared in Ahl (1929:273). Synonymy proposed by Mateussi et al., 2018.

Distribution: 307, 314, 315, 316, 317, 321 (Mateussi et al., 2018).

Remarks: Revalidated by Mateussi et al., 2018.

Mylossoma aureum (Agassiz, 1829)

Myletes aureus Agassiz , in Spix , Agassiz, 1829:74, Pl. 31. Type locality: Brasiliae aequinoctialis flaviis. [ Brazil]. Lectotype: MHNN 788 -A. [Designated by Mateussi et al., 2018:291, fig. 10(p.289)].

Myletes herniarius Cope, 1872b:268 , Pl. 12 (fig. 3). Type locality: Ambyiacu. [ Río Ampyiacu , Peru]. Lectotype: ANSP 8025 . [Designated by Fowler (1907:476, fig. 56); Böhlke (1984:48)]. See Taphorn (1992:284) for discussion on species taxonomic status.

Mylossoma ventriosa Norman, 1929:815 , fig. 20. Type locality: Solimões, R. Amazon. Holotype: BMNH 1923.10 .28.263. See Machado-Allison, Castillo (1992:17) for discussion on species taxonomic status.

Distribution: 306, 307, 308, 309, 313, 316, 318, 319, 321 (Jégu, 2003; Ota et al., 2013; Mateussi et al., 2018).

Remarks: Author of species name is Agassiz according to Kottelat (1988:79) followed by Jégu (2003) and Mateussi et al. (2018). Species redescribed by Mateussi et al. (2018:286).

Mylossoma duriventre (Cuvier, 1818)

Myletes duriventris Cuvier, 1818:451 , Pl. 22 (fig. 2). Type locality: Brésil. [ Brazil]. MNHN A.9891. [stuffed specimen. See Mateussi et al. (2018:295, fig. 16) for holotype image].

Myletes orbignyanus Valenciennes , in Cuvier, Valenciennes, 1850:220 . Type locality: Corrientes, dans le Parana . Lectotype: MNHN A-9863 (235 mm SL specimen). [Designated by Géry et al. (1987:444). Stuffed specimen]. See Machado-Allison, Castillo (1992:19) for discussion of species taxonomic status.

Mylosoma ocellatum Eigenmann, 1915:265 . Type locality: Villa Hays, Paraguay. Holotype: FMNH 56816. Originally appeared as a doubtful new species, probably a juvenile of Myletes albiscopus Cope, 1872 . Original genus should be Mylossoma .

Mylossoma paraguayensis Norman, 1929:814 , fig. 19. Type locality: Rio Paraguay. Syntypes: BMNH 1895.5 .17.254 (1), BMNH 1895.5 .17.255 (1) Río Paraguay, BMNH 1910.5 .26.46 (1) San Salvador, BMNH 1910.5 .26.47 (1), BMNH 1910.5 .26.48 (1) Puerto Asir.

Distribution: 343, 344, 345 (Mateussi et al., 2018).

Remarks: Distribution of species limited to the Rio Paraguai basin, lower Paraná and lower Uruguai rivers basins (Mateussi et al., 2018); records for the Amazon River basin are misidentifications mainly of M. albiscopum.

Mylossoma unimaculatum (Steindachner, 1908)

Metynnis unimaculatus Steindachner, 1908d:326 . Type locality: Lagune am Rio Medonho, einem Nebenfluss des Pernahyba nördlich von Sa. Filomena. [Type locality in the original descrption most probably incorrect (Mateussi et al. (2018:303))]. Syntypes: NMW 56451 (3). [Illustrated in Mateussi et al. (2018:303, fig. 20)].

Distribution: 324 ([Mateussi et al. (2018:303)]).

Remarks: See Mateussi (2015: 51]) for new combination.

OSSUBTUS Jégu, 1992

Ossubtus Jégu, 1992:240 . Type species: Ossubtus xinguense Jégu, 1992 . Type by original designation. Gender: Neuter.

Remarks: Recovered as closely related to Myloplus schomburgkii in Kolmann et al. (2020) and Mateussi et al. (2020b).

Ossubtus xinguense Jégu, 1992

Ossubtus xinguense Jégu, 1992:240 , figs. 4, 11. Type locality: Rio Xingu , Altamira, Pará, Brésil. [ Brazil]. Holotype: INPA 6535 . [Holotype INPA 6535 wrongly illustrated by Jégu (1992). See Andrade et al. (2016d: fig. 3) for illustration of the correct holotype].

Distribution: 322 (Jégu, 2003; Andrade et al., 2016d).

Remarks: See Andrade et al. (2016d) for species redescription, illustration of the correct holotype (female instead of male in original description), and conservation status. Species delimitation using mitochondrial gene cytochrome c oxidase subunit I (COI) in Machado et al. (2018).

PARAMYLOPLUS Norman, 1929