Characidae Latreille, 1825

Melo, Bruno F, Ota, Rafaela P, Benine, Ricardo C, Carvalho, Fernando R, Lima, Flavio C T, Mattox, George M T, Souza, Camila S, Faria, Tiago C, Reia, Lais, Roxo, Fabio F, Valdez-Moreno, Martha, Near, Thomas J & Oliveira, Claudio, 2024, Phylogenomics of Characidae, a hyper-diverse Neotropical freshwater fish lineage, with a phylogenetic classification including four families (Teleostei: Characiformes), Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (1), pp. 1-37 : 11-12

Characidae Latreille, 1825 , new usage

Type genus: Charax Scopoli, 1777 .

Included subfamilies: Aphyocharacinae , Characinae , Cheirodontinae , Exodontinae, and Tetragonopterinae .

Definition: The least inclusive crown clade that contains Charax gibbosus (Linnaeus, 1758) , Aphyocharax pusillus Günter, 1868 , Cheirodon pisciculus Girard, 1855 , Exodon paradoxus Müller and Troschel, 1844 , and Tetragonopterus argenteus Cuvier 1816 . This is a minimum-crown-clade definition. See Figure 4 for a reference phylogeny of Characidae.

Etymology: From the ancient Greek ΧάΡαξ (kˈɑː͡ɹɹaeks) as a name for species of Sparidae that exhibit teeth on the oral jaws ( Thompson 1947: 284–5).

Remarks: There are no known morphological synapomorphies for Characidae ; however, most species in the clade have a pseudotympanum, a hiatus of the hypaxial muscle along the anterior portion of the gas bladder (Malabarba 1998; Mattox and Toledo-Piza 2012). The pseudotympanum exhibits different shapes, muscles, and rib delimitation among the Aphyocharacinae , Characinae , Cheirodontinae , Hyphessobrycon (cf. Carvalho 2011), and some species of Tetragonopterinae (Malabarba 1998, Mattox and Toledo-Piza 2012). A pseudotympanum is also present in members of Alestidae , Crenuchidae , Cynodontidae , and Serrasalmidae (Zanata and Vari 2005; Mattox and Toledo-Piza 2012, Zanata and Camelier 2014, 2015) and may represent a plesiomorphic condition in Characiformes as it is present in cithariniforms ( Vari 1979, Mattox and Toledo-Piza 2012), siluriforms ( Shibatta and Vari 2017, Slobodian et al. 2017, 2021), gymnotiforms (Dutra et al. 2015), and cypriniforms (e.g. Britz et al. 2021). However, additional investigations are needed to refine our understanding of this character across Characiformes and Ostariophysi .

Several authors have hypothesized a close relationship between Cheirodontinae and Aphyocharacinae based on morphological similarities (Eigenmann 1917, Géry 1977, Malabarba 1998). Alternatively, Mirande (2010) resolved Cheirodontinae as the sister-group of Aphyoditeinae and this clade as the sister-group of Aphyocharacinae . Molecular phylogenetic studies consistently resolve Cheirodontinae and Aphyocharacinae as a monophyletic group (Calcagnotto et al. 2005, Javonillo et al. 2010, Oliveira et al. 2011, Tagliacollo et al. 2012, Melo et al. 2016).

In a pre-cladistic study, Géry (1964c) hypothesized a close relationship between Characinae and Exodontinae by proposing the tribe Exodontidi , which included Exodon , Roeboexodon , and Roeboides Günther, 1864 . Molecular phylogenies support a close relationship between the Characinae and Exodontinae (Javonillo et al. 2010, Oliveira et al. 2011, Melo et al. 2016), and phylogenomic studies resolve Exodontinae as the sister-lineage of a clade containing Tetragonopterinae and Characinae ( Fig. 4; Betancur-R et al. 2019, Melo et al. 2022a, Souza et al. 2022).