Aphpocharacinae Eigenmann, 1909

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 : 12

Aphpocharacinae Eigenmann, 1909 , new usage

Type genus: Aphyocharax Günther, 1868 .

Included genera: Aphyocharax , Cyanogaster , Leptagoniates Boulenger, 1887 , Paragoniates Steindachner, 1876 , Phenagoniates Eigenmann and Wilson in Eigenmann et al., 1914, Prionobrama Fowler, 1913 , Xenagoniate s Myers, 1942. Not sampled: Amazonichthys Esguícero and Mendonça, 2023, and Aphyocharacidium .

Definition: The least inclusive crown clade that contains Aphyocharax pusillus Günther, 1868 and Cyanogaster noctioaga Mattox et al., 2013 . This is a minimum-crown-clade definition. See Figure 4 for a reference phylogeny of Aphyocharacinae . Species of Amazonichthys and Aphyocharacidium were not sampled in this study.

Etymology: From the ancient Greek ἀϕύΗ (ɐfɪˈae), which is a name used by ancient authors for anchovies, smelts, silversides, and the goby Aphia minuta (Risso 1810) and ΧάΡαξ (kˈɑː͡ɹɹaeks) as a name for species of Sparidae that exhibit teeth on the oral jaws ( Thompson 1947: 21–2, 284–5).

Remarks: Recent taxonomic treatments of Characidae included eight genera in the Aphyocharacinae : Paragoniates , Phenagoniates , Xenagoniates , Inpaichthys , Leptagoniates (not analysed), Rachooiscus Myers, 1926 (not analysed), Aphyocharax , and Prionobrama (Mirande 2009, 2010). Phylogenetic studies using molecular data or combined morphological and molecular characters demonstrate that Rachooiscus and Inpaichthys are not closely related to Aphyocharacinae ( Oliveira et al. 2011) , resolves Aphyocharacidium (latter identified as Hemigrammus cf. geisleri Zarske and Géry, 2007 ) within Aphyocharacinae ( Tagliacollo et al. 2012) , and resulted in a delimitation of Aphyocharacinae that includes Aphyocharacidium , Aphyocharax , Prionobrama , Paragoniates , Phenagoniates , Leptagoniates , and Xenagoniate s supported by two dorsal-fin rays articulating with the first dorsal pterygiophore ( Tagliacollo et al. 2012, Vari et al. 2016). More recently, the composition of Aphyocharacinae was expanded to include Axelrodia lindeae Géry, 1973 and 12 synapomorphies were identified for the clade ( Mirande 2019); A. lindeae has been transferred to the newly described Amazonichthys (Esguícero and Mendonça 2023).

The phylogeny inferred from the UCE loci resolved Aphyocharax and Prionobrama as sister-lineages and a clade containing Paragoniates , Phenagoniates , Leptagoniates , and Xenagoniate s ( Fig. 4). A result not presented in other phylogenetic studies is the resolution of a clade in Aphyocharacinae that contains Cyanogaster noctioaga Mattox et al., 2013 , four undescribed species of Cyanogaster (Leticia, Tapajós, Negro, and Apure), and a specimen previously identified as Hemigrammus geisleri ( Fig. 4). Cyanogaster noctioaga was described as a miniature characid belonging to the Stevardiinae based on the presence of ii + 8 dorsal-fin rays and four teeth in the inner premaxillary series (Mattox et al. 2013). Phylogenetic analysis of morphological characters resolves Cyanogaster in the Stevardiinae ( Mirande 2019) . In the phylogenies inferred from the UCE loci, Cyanogaster was consistently resolved as the sister-lineage of all other species of Aphyocharacinae ( Fig. 4). Considering that the type species of Hemigrammus ( H. unilineatus Gill, 1858 ) is phylogenetically placed in the Pristellinae, we transfer Hemigrammus geisleri to the genus Cyanogaster as Cyanogaster geisleri , new combination ( Fig. 4; Table 1 ). Aphyocharacidium remains unsampled in phylogenomic studies and thus with uncertain position and here tentatively included in Aphyocharacinae . Species of Aphyocharacinae are primarily distributed in Amazon–Orinoco–Guianas; Cyanogaster geisleri and Prionobrama paraguayensis (Eigenmann 1914) extend to the Paraguay basin of La Plata, and Phenagoniates to the trans-Andean region ( Fig. 4).