Hpphessobrpconinae Lima, Carvalho & Faria

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 : 25-26

Hpphessobrpconinae Lima, Carvalho & Faria , new subfamilp

ZooBank: urn:lsid:zoobank.org:act:B7A1E353-CC55-4480-842E-CFA88155FA94 .

Type genus: Hyphessobrycon Durbin in Eigenmann, 1908.

Included genera: Dinotopterygium Frainer et al., 2021 , Erythrocharax Netto-Ferreira et al., 2013 , Hemigrammus (in part), Hyphessobrycon (in part), Jupiaba (in part), Macropsobrycon (in part), Moenkhausia (in part), Parecbasis , and Phycocharax Ohara et al., 2017 .

Definition: The least inclusive crown clade that contains Hyphessobrycon agulha Fowler, 1913 and Jupiaba polylepis (Günther, 1864) . This is a minimum-crown-clade definition. See Figure 6 for a reference phylogeny of Hyphessobryconinae .

Etymology: From the ancient Greek ὑϕήσσων (hˈuːfɑːsˌo͡ʊn) meaning of lesser stature and ΒΡύκω (bɹˈʊka͡ʊ) meaning to bite.

Remarks: The new subfamily Hyphessobryconinae is delimited as containing many species and genera that have long been challenged in the systematics and taxonomy of Characiformes ( Fig. 6). A clade comprising eight species currently classified in Jupiaba and Hyphessobrycon moniliger is the sister-lineage of all other species of Hyphessobryconinae ( Fig. 6). Within Hyphessobryconinae there is a clade that includes several species with large bony hooks on the anal fin of males: Parecbasis cyclolepis Eigenmann, 1914 , eight species of Hyphessobrycon , one species of Macropsobrycon , three species of Moenkhausia , and three species of Hemigrammus ( Fig. 6). A close phylogenetic affinity of species sharing large bony hooks on the anal fins of males had been suggested for the Hyphessobrycon panamensis group (Ota et al. 2020), H. bayleyi Lima et al., 2022, H. diancistrus , and H. otrynus (Lima et al. 2022).

Hyphessobrycon compressus is the type species of the genus and in the UCE phylogeny resolves in a monophyletic group that also contains H. columbianus Zarske and Géry, 2002 , H. ecuadorensis (Eigenmann, 1915) , H. panamensis Durbin, 1908 , and Hyphessobrycon sp. Dagua ( Fig. 6). This monophyletic trans-Andean lineage was supported in other phylogenetic studies that included H. bussingi Ota et al., 2020 , H. columbianus , H. compressus , H. condotensis Regan, 1913 , H. ecuadoriensis Eigenmann and Henn, 1914 , H. panamensis , H. saoagei Bussing, 1967 , and H. tortuguerae Böhlke, 1958 (Melo et al. 2022a, Elías et al. 2023), which differs from the hypothesis based on morphology of a close relationship with cis-Andean species of the rosy tetra clade (Weitzman and Palmer 1997, Carvalho and Malabarba 2015). Pending additional taxonomic and phylogenetic research, we refrain from proposing changes in the generic classification and nomenclature for Moenkhausia gracilima , Moenkhausia mikia , Hemigrammus leois , Macropsobrycon xinguensis , Moenkhausia ceros , and Hemigrammus hyanuary .

The specimen identified as Hyphessobrycon sp. Dagua (15715) from Río Dagua in south-western Colombia in the UCE phylogeny has the morphological features of Astyanax daguae Eigenmann, 1913 , such as complete lateral line and absence of scales over caudal-fin lobes (Eigenmann 1913). The species was recently analysed based on a single mitochondrial gene and transferred to Tetragonopterus ( Terán et al., 2020) . Due to the phylogenetic position of the specimen relatively close to Hyphessobrycon compressus ( Fig. 6), and the presence of a second species Hyphessobrycon daguae (Eigenmann, 1922) in that river, we use the provisional name Hyphessobrycon sp. Dagua, with an indication that future research will evaluate the status of both species. This also reveals that the species should not be placed in the genus Tetragonopterus ( Tetragonopterinae ) within Characidae ( Fig. 3). The genus Macropsobrycon requires additional taxonomic research because Macropsobrycon xinguensis phylogenetically resolved as a lineage of Hyphessobryconinae , but the type species of the genus Macropsobrycon uruguayanae is placed in Cheirodontinae ( Characidae ) in the UCE phylogeny ( Figs 3, 6).

Erythrocharax , Dinotopterygium , and Phycocharax are speciesdepauperate genera described over the past 10 years that are nested in a clade of Hyphessobryconinae that includes several species of Hyphessobrycon ( Fig. 6; Netto-Ferreira et al. 2013, Ohara et al. 2017, Frainer et al. 2021). Species in this clade are predominantly from the Brazilian Shield and possess up to

seven cusps, with Hyphessobrycon psitacus Dagosta et al., 2016 being the exception with five cusps. Some species have a higher number of cusps, such as Erythrocharax altipinnis with up to eight cusps, and H. juruna Faria et al., 2018 , P. rasbora Ohara et al., 2017 , and Dinotopterygium diodon Frainer et al., 2021 with up to nine cusps.

Dinotopterygium has a unique combination of 10 synapomorphies, including modifications in anal-fin morphology leading to the anal-fin base strongly convex in males, and suggesting a close relationship with Erythrocharax and Phycocharax (Frainer et al. 2021) . Phycocharax rasbora has a unique combination of characters: presence of a single row of relatively compressed premaxillary teeth, large teeth with four to nine cusps on premaxilla and dentary, absence of pseudotympanum, incomplete lateral line, sexually dimorphic males with distal margin of anal fin approximately straight, and presence of a nearly triangular and horizontally elongated blotch from the posterior half of the body to caudal peduncle ( Ohara et al. 2017). Phylogenetic analysis of morphological characters resolved a clade containing Phycocharax , Paracheirodon axelrodi (Schultz, 1956) , Hyphessobrycon elachys Weitzman, 1985 , H. loweae Costa and Géry, 1994, and H. oanzolinii Lima and Flausino Junior, 2016 ( Ohara et al. 2017). The relationships resolved in the UCE phylogeny corroborate the phylogenetic affinities of relationship between Phycocharax and H. loweae , but are not consistent with a close phylogenetic relationship with P. axelrodi . Hyphessobrycon elachys and H. oanzolinii were not included in phylogenetic analyses of the UCE loci.

The phylogeny resolves a clade containing 14 species of Hyphessobrycon , Hemigrammus cf. bellotii (Steindachner, 1882) , Hemigrammus rubrostriatus Zarske, 2015 , and Hemigrammus sp. Leticia ( Fig. 6). Species in this clade have teeth with up to five cusps; the presence of a longitudinal black stripe typically starting posteriorly to the humeral blotch (when humeral blotch is present) and typically darker on the caudal peduncle region. The black stripe presents interspecific variation, ranging from weakly marked and diffuse (e.g. Hyphessobrycon amapaensis Zarske and Géry, 1998 and Hemigrammus bellotii ) to very dark and broad stripe (e.g. Hyphessobrycon peruoianus Ladiges, 1938 ), and in some cases more than one pattern may occur on the same species (e.g. Hyphessobrycon agulha ). The known species of the group share a thin iridescent stripe above the black stripe and humeral blotch. Our phylogeny indicates that live coloration patterns might support monophyletic assemblages within the Hyphessobryconinae , opening new avenues for studies within these clades.

Many of the species in this clade were previously classified in the Hyphessobrycon agulha and Hyphessobrycon heterorhabdus (Ulrey 1894) species-groups (Géry 1977, Lima et al. 2014, Faria et al. 2020a, b). The Hyphessobrycon agulha species group is defined by the presence of ‘a broad, relatively diffuse lateral stripe (typically more discernible ventrally, posterior to the midbody), and a humeral blotch that may or may not coalesce with the stripe (although a humeral blotch is absent in H. loretoensis Ladiges, 1938 and H. mutabilis Costa and Géry, 1994)’ ( Ohara and Lima 2015) and in its most recent proposition is composed of Hyphessobrycon agulha , Hyphessobrycon claoatus Zarske , 2015, Hyphessobrycon eschwartzae García-Alzate et al., 2013,

Hyphessobrycon herbertaxelrodi Géry, 1961, Hyphessobrycon klausanni García-Alzate et al., 2017, Hyphessobrycon loretoensis , Hyphessobrycon lucenorum Ohara and Lima, 2015 , Hyphessobrycon margitae Zarske, 2016 , Hyphessobrycon metae Eigenmann and Henn, 1914 , Hyphessobrycon mutabilis , Hyphessobrycon peruoianus , Hyphessobrycon wadai Marinho et al., 2016 , and Hyphessobrycon zoe Faria et al., 2020 (Faria et al. 2020a). The Hyphessobrycon heterorhabdus species group is currently defined by a tricolor longitudinal pattern along midbody, i.e. dorsal red stripe, middle iridescent stripe and ventral longitudinal black pattern composed by a single humeral blotch, a midlateral black stripe continuous with humeral blotch and increasingly blurred towards caudal peduncle (Faria et al. 2020b); the group includes Hyphessobrycon amapaensis , Hyphessobrycon cantoi Faria et al., 2021 , Hyphessobrycon heterorhabdus , Hyphessobrycon ericae Moreira and Lima, 2017 , Hyphessobrycon montagi Lima et al., 2014, Hyphessobrycon sateremawe Faria et al., 2020 , and Hyphessobrycon wosiackii Moreira and Lima, 2017 (Faria et al. 2021). The phylogeny inferred from the UCE loci resolves both the Hyphessobrycon agulha and H. heterorhabdus species-groups as non-monophyletic ( Fig. 6).