Characinae 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 : 16

Characinae Latreille, 1825 , new usage

Type genus: Charax Scopoli, 1777 .

Includedgenera: Acanthocharax Eigenmann,1912 , Acestrocephalus Eigenmann, 1910 , Atopomesus , Charax , Cynopotamus Valenciennes, 1850 , Galeocharax Fowler, 1910 , Phenacogaster , and Roeboides . Not sampled: Microschemobrycon .

Definition: The least inclusive crown clade that contains Charax gibbosus (Linnaeus, 1758) , Atopomesus pachyodus Myers, 1927 , Phenacogaster pectinata (Cope, 1870) , and Acestrocephalus anomalus (Steindachner, 1880) . This is a minimum-crown-clade definition. See Figure 4 for a reference phylogeny of Characinae .

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: A group that includes Charax , the type genus of Characiformes, and small to medium-sized predators such as Acanthocharax , Acestrocephalus , Cynopotamus , Galeocharax , and Roeboides have been treated as closely related prior to the application of Hennigian phylogenetic systematics (Howes 1976, Géry 1977). Phylogenetic analysis of morphological characters led to a delimitation of Characinae that included Phenacogaster , Priocharax Weitzman and Vari, 1987 , and six genera of heterocharacins (Lucena 1998) currently classified in Acestrorhynchidae ( Oliveira et al. 2011) . Subsequent studies identified a number of morphological synapomorphies, removed the heterocharacins, and added Microschemobrycon to the Characinae (Mirande 2009, 2010, 2019, Mattox and Toledo-Piza 2012).

Phylogenomic analysis of UCE loci results in the resolution of four major lineages of Characinae delimited as tribes ( Souza et al. 2022): Phenacogasterini ( Phenacogaster ), Acanthocharacini ( Acanthocharax ), Cynopotamini ( Acestrocephalus , Cynopotamus, and Galeocharax), and Characini (Charax and Roeboides). The UCE inferred phylogeny presented here is congruent with trees presented by Souza et al. (2022) and includes Atopomesus as the sister-lineage of all other species of Characinae ( Fig. 4). Previous morphological phylogenetic studies resolved Atopomesus in the Spintherobolinae ( Mirande 2019). To investigate this novel phylogenetic hypothesis, specimens from the sequenced lot of Atopomesus (LBP 23871) were prepared for muscle and skeleton observation, revealing that Atopomesus possesses four Characinae synapomorphies, i.e. characters 3, 7, 8, and 10 of Mattox and Toledo-Piza (2012). Microschemobrycon was not sampled in the UCE inferred phylogeny but is treated here as incertae sedis in Characinae following the results from combined multilocus and morphological phylogenetics ( Mirande 2019).