Grundulinae Fowler, 1958

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 : 28-29

Grundulinae Fowler, 1958 , new usage

Type genus: Grundulus Valenciennes, 1846 .

Included genera: Astyanacinus Eigenmann, 1907 and Grundulus .

Definition: The least inclusive crown clade that contains Grundulus bogotensis and Astyanacinus moorii (Boulenger, 1892) . This is a minimum-crown-clade definition. See Figure 7 for a reference phylogeny of Grundulinae .

Etymology: From the Middle English grundel in reference to several species of fishes.

Remarks: Grundulus was described for Poecilia bogotensis Humboldt, 1821 (Valenciennes in Cuvier and Valenciennes, 1846) and includes three species endemic to lakes in the northern Andes of South America ( Román-Valencia et al. 2005). Morphological phylogenetic analysis resolved Grundulus as monophyletic, supported with 11 synapomorphies ( Román-Valencia et al. 2010), and G. quitoensis Román-Valencia et al., 2005 is the sister-species of a clade including G. cochae Román-Valencia et al, 2003 and G. bogotensis . Alternative phylogenetic studies using morphology resolved Grundulus and Coptobrycon from eastern Brazil as sister-lineages classified in the subfamily Gymnocharacinae (Mirande 2009, 2010). Phylogenetic analysis of combined morphological and molecular data resolved Grundulus as closely related to Coptobrycon , Stygichthys , some species of Hyphessobrycon , Phycocharax , Myxiops , Probolodus , Deuterodon , and a species of Astyanax , which were classified in the tribe Grundulini Fowler, 1958 ( Mirande 2019). This delimitation of Grundulini has not been supported in molecular phylogenetic analyses ( Figs 3–7; Oliveira et al. 2011, Melo et al. 2022a).

Astyanacinus was described by Eigenmann (1907: 769) and differentiated from Astyanax by possessing a ‘lengthened upper jaw’ (Géry 1977: 415). Astyanacinus was synonymized with Astyanax based on the phylogenetic resolution of Astyanacinus moorii (Boulenger, 1892) as nested within Astyanax , a result supported by two morphological synapomorphies (Terán et al. 2020). A phylogenomic study resolved Grundulus and Rhoadsia

as sister-lineages and Astyanacinus as the sister-lineage of a clade containing Psellogrammus Eigenmann, 1908 , Ctenobrycon Eigenmann, 1908 , Oligosarcus , and Astyanax (Betancur et al. 2019) . An earlier phylogenomic study using UCE loci reflects results presented in Figure 7, with the resolution of a clade containing Grundulus and Astyanacinus that is the sister-lineage of a clade containing Rhoadsiinae , Psellogrammus , Ctenobrycon , Oligosarcus , and Astyanax (Melo et al. 2022a) .

The phylogenetic analysis of UCE loci supports the monophyly of Grundulinae , which are the sister-lineage of a species-rich clade that we delimit as the subfamily Acestrorhamphinae ( Fig. 7). Based on the UCE phylogeny, we recognize the subfamily Grundulinae and revalidate Astyanacinus as a monotypic genus ( Fig. 7; Table 1 ). A taxonomic revision delimited the Astyanax orthodus species group that included Astyanacinus moori (Ruiz-C. et al. 2018), indicating the presence of additional species currently classified as Astyanax that may be more closely related to Astyanacinus . Grundulinae exhibit a geographically disjunct distribution with Grundulus in the Andes and Astyanacinus in the La Plata basin ( Fig. 7).