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Phylogeny of the family Characidae (Teleostei: Characiformes): from characters to taxonomy
Abstract
The family Characidae is the most diverse among Neotropical fishes. Systematics of this family are mainly based on precladistic papers, and only recently a phylogenetic hypothesis for Characidae was proposed by the author. That phylogeny was based on 360 morphological characters studied for 160 species, including representatives of families related to Characidae. This paper is based on that phylogenetic analysis, with the analyzed characters described herein and documented, accompanied by comparisons of their definition and coding in previous papers. Synapomorphies of each node of the proposed phylogeny are listed, comparisons with previous classifications provided, and autapomorphies of the analyzed species listed. Taxonomic
implications of the proposed classification and the position of the incertae sedis genera within Characidae are discussed. A discussion of the phylogenetic information of the characters used in the classical systematics of the Characidae is provided.
... The phylogenetic relationships of Amazonichthys were properly addressed for the first time by B€ uhrnheim (2006) in phylogenetic study based on morphological characters, in which A. lu (¼ her A. lindeae) was hypothesized as a member of the subfamily Cheirodontinae; however, as the main objective of this study was to investigate the phylogenetic relationships of members of the subfamily Cheirodontinae, the sampling of the other representatives of Characidae was scarce. In a more comprehensive analysis, Mirande (2009Mirande ( , 2010 reported a close relationship between A. lu (¼ his A. lindeae) and Aphyocharacidium bolivianum Géry, 1973 in a phylogenetic study based on morphological characters, corroborating the hypothesis of Géry (1973), reporting the close relationship of these two genera with Microschemobrycon casiquiare Böhlke, 1953, Aphyodite grammica Eigenmann, 1912, and Parecbasis cyclolepis Eigenmann, 1914, all of them grouped in the subfamily Aphyoditeinae. Over time, the allocation of Amazonichthys in the subfamily Aphyoditeinae (sensu Mirande, 2010) was supported primarily by its dentition morphology (Géry, 1973(Géry, , 1977Mirande, 2009Mirande, , 2010, such as the presence of only one row of more than seven uni-to tricuspid teeth on the premaxillary bone. ...
... In a more comprehensive analysis, Mirande (2009Mirande ( , 2010 reported a close relationship between A. lu (¼ his A. lindeae) and Aphyocharacidium bolivianum Géry, 1973 in a phylogenetic study based on morphological characters, corroborating the hypothesis of Géry (1973), reporting the close relationship of these two genera with Microschemobrycon casiquiare Böhlke, 1953, Aphyodite grammica Eigenmann, 1912, and Parecbasis cyclolepis Eigenmann, 1914, all of them grouped in the subfamily Aphyoditeinae. Over time, the allocation of Amazonichthys in the subfamily Aphyoditeinae (sensu Mirande, 2010) was supported primarily by its dentition morphology (Géry, 1973(Géry, , 1977Mirande, 2009Mirande, , 2010, such as the presence of only one row of more than seven uni-to tricuspid teeth on the premaxillary bone. ...
... In a more comprehensive analysis, Mirande (2009Mirande ( , 2010 reported a close relationship between A. lu (¼ his A. lindeae) and Aphyocharacidium bolivianum Géry, 1973 in a phylogenetic study based on morphological characters, corroborating the hypothesis of Géry (1973), reporting the close relationship of these two genera with Microschemobrycon casiquiare Böhlke, 1953, Aphyodite grammica Eigenmann, 1912, and Parecbasis cyclolepis Eigenmann, 1914, all of them grouped in the subfamily Aphyoditeinae. Over time, the allocation of Amazonichthys in the subfamily Aphyoditeinae (sensu Mirande, 2010) was supported primarily by its dentition morphology (Géry, 1973(Géry, , 1977Mirande, 2009Mirande, , 2010, such as the presence of only one row of more than seven uni-to tricuspid teeth on the premaxillary bone. ...
... Since the taxonomic revision of the genus by Eigenmann (1917Eigenmann ( , 1921Eigenmann ( ,1927, the genus Astyanax has been diagnosed by a combination of characters that includes two rows of premaxillary teeth, five teeth in the inner premaxillary series, maxillary with a few or no teeth; lateral line complete, adipose fin present, caudal fin lacking scales at its basis, and no predorsal spine. Astyanax monophyly has been questioned in the last decades, and according to recent studies, the genus was not monophyletic according to its classical diagnosis (see Javonillo et al. 2010;Mirande 2010;Oliveira et al. 2011;2019;Terán et al. 2020). Terán et al. (2020) provided the most comprehensive phylogenetic analysis of the genus, based on a dataset combining morphological and molecular data, and concluded that some species previously described in Astyanax should be reassigned to the genera Deuterodon Eigenmann, 1907;Eretmobrycon Fink, 1976;Jupiaba Zanata, 1997;Psalidodon Eigenmann, 1911; Tetragonopterus Cuvier, 1816; and two newly described genera, Andromakhe and Makunaima (Terán et al. 2020). ...
... The analyzed dataset is composed of 520 morphological characters and 9 blocks of molecular data. The morphological dataset was initially published by Mirande (2009Mirande ( , 2010 and was successively improved and expanded through several contributions (Mirande et al. 2013;Ohara et al. 2017Ohara et al. , 2019Mirande 2019;Terán et al. 2020). The molecular data include mitochondrial ribosomal 12S and 16S, ATP synthase F0 subunit 6 (ATP6), cytochrome c oxidase subunit I (COX1), cytochrome b (CYTB) and the nuclear coding markers myosin heavy chain 6 (MYH6), patched domain containing 1 (PTCHD1), and recombination activating 1 and 2 (RAG1 and RAG2). ...
A new species of Astyanax Baird & Girard, 1854 is described from the rio Apiacás, a tributary of the rio Teles Pires, rio Tapajós basin, Mato Grosso state, Brazil. The new taxon can be distinguished from all congeners, except those belonging to the Astyanax bimaculatus species group and to the Astyanax orthodus species group, by the presence of a horizontally elongated to rounded humeral blotch. The new taxon can be readily distinguished from all species belonging to the A. bimaculatus species group and to the A. orthodus species group by presenting a distinct morphology in premaxillary and dentary teeth with conspicuous diastema (a teeth gap) between them. We also present a hypothesis about the phylogenetic relationships of the new taxon within Astyanax.
... P. melanotus (MZUSP 26068) diagnosable by three autapomorphies related to jaw and teeth morphology and one related to the color pattern. The genus then appeared in a basal polytomy in the subfamily along with the genera Odontostilbe Cope, 1870, Aphyocheirodon Eigenmann, 1915, Pseudocheirodon Meek & Hildebrand, 1916, and Cheirodontops Schultz, 1944. Mirande (2010 further recovered P. melanotus as sister group to remaining cheirodontines analyzed therein (Odontostilbe, Serrapinnus Malabarba, 1998, andCheirodon Girard, 1855) and described nine autapomorphies for the type-species of the genus, but none exclusive to Prodontocharax. ...
Prodontocharax species are revised and the genus Amblystilbe is revalidated based on analysis of type-material and additional specimens. Both genera are diagnosed based on unique synapomorphies among members of the Cheirodontinae related to shape, size, number and arrangement of teeth in the jaw bones. Prodontocharax melanotus, from the upper rio Madeira basin, Brazil and Bolivia, is redescribed, and a new species is described from the rio Huallaga basin, Peru. The genus Amblystilbe and its type-species, A. howesi, are redescribed from the Amazonas River basin, Bolivia, Brazil, and Peru, and Prodontocharax alleni is considered a junior synonym of A. howesi. The two species of Prodontocharax are distinguished by the color pattern, number of lamellae of the olfactory rosette in male and female and number of gill rakers, and differ from A. howesi by the number and shape of teeth, color pattern and number of branched anal-fin rays. According to recent studies, the species of Prodontocharax and A. howesi cluster into two distinct clades among the cheirodontines.
... None of the abovementioned characteristics is unique for Moenkhausia, and all of them have been demonstrated to be highly homoplastic within the Characidae. This not only indicates the artificially proposed delimitation, but also strongly suggests the non-monophyly of this genus (Mirande, 2010(Mirande, , 2019Dagosta & Marinho, 2016;Ohara et al., 2019). ...
A new species of Moenkhausia is described from the rio Braço Norte, a tributary of Rio Teles Pires draining the Serra do
Cachimbo, rio Tapajós basin, Pará, Brazil. The new species is diagnosed from all congeners, except M. moisae and M.
pirauba, by having a high number of scales in the longitudinal series (43–46 vs. 23–41 in other Moenkhausia species). It
can also be distinguished from the aforementioned species based on the combination of the following characters: a single
humeral blotch, 21–25 branched anal-fin rays, and a round and symmetrical caudal blotch not continuous anteriorly with
the dark midlateral stripe. The new tetra herein described represents an additional, possibly endemic, taxon from the
headwaters draining from Serra do Cachimbo, in the Brazilian Shield.
... Garcia-Ayala, Benine (2020) referred to as "Stethaprion Clade" the clade formed by the genera Stethaprion Cope, 1870, Brachychalcinus Boulenger, 1892, Poptella Eigenmann, 1908, and Orthospinus Reis, 1989, as hypothesized by Mirande (2019). These taxa are diagnosed by the presence of three unbranched dorsal-fin rays, of which the first one is modified into an anteriorly oriented spine-like element (Mirande 2010(Mirande , 2019, traditionally called "predorsal spine" (Eigenmann, 1907;Géry, 1977;Reis, 1989). ...
A new species of Poptella is described from the rio Xingu basin, Pará and Mato Grosso states, Brazil. The new species is distinguished from all congeners by having a greater number of scales around the caudal peduncle (15–18 vs. 13–14). Furthermore, this species can be easily distinguished from P. brevispina, P. longipinnis, and P. fuscata by the lower number of branched rays in the dorsal fin (9 vs. 10–11). The description of new taxon represents the third species of the genus Poptella described in the last five years.
... Digestive tract in cardinal tetra (Fricke et al., 2022). The great morphological and behavioral diversity observed in this family is related to the variety of environments that they inhabit (Mirande, 2010). Paracheirodon axelrodi is a non-migratory freshwater fish, which generally lives in shoals in the middle water layers with temperatures between 23°C and 27°C (Anjos, Anjos, 2006;Brito, Bazzoli, 2009). ...
The cardinal tetra Paracheirodon axelrodi is a species of the family Characidae of great interest as an ornamental fish. Many aspects of the biology of this species are still unknown. The present work presents a complete description of the different components of the digestive tract of P. axelrodi, analyzing its global anatomical arrangement, the adjacent glands (liver and pancreas) and the previously ignored pyloric caeca, using histological and histochemical techniques with light microscopy and scanning electron microscopy. Three levels of analysis were performed: macroscopic anatomy, histology of the gastrointestinal tract and scanning electron microscopy of mucosal surfaces. The intestinal coefficient (Ci) of P. axelrodi calculated here (0.94) is within the expected range for a carnivorous species. The anatomy of the alimentary and the resulting Ci is consistent with carnivorous diet in line with dietary studies in the wild in this species. The digestive tract, including adjacent glands (liver and pancreas), presented a relatively conserved general pattern within that described for other species of the family Characidae, especially at the histological level.
... Mais tarde, Weitzman e colaboradores consideram as diferenças entre as espécies dos dois gêneros e suas distribuições (rio da costa Atlântica versus rios da costa do Pacífico) e revalidaram Hollandichthys até que uma análise filogenética que demonstrasse as possíveis relações entre estes gêneros estivesse disponível 9 . Estudos realizados com dados moleculares 12,13 resultaram na hipótese de que Hollandichthys é grupo-irmão de Rachoviscus (Figura 9) em vez de Pseudochalceus como proposto por estudos com apenas análises morfológicas 1,14 . As hipóteses com dados moleculares foram corroboradas posteriormente com base em dados de osteologia e morfologia 15 . ...
Resumo: Hollandichthys, popularmente conhecido como lambari-listrado, é um gênero monofilético da família Characidae com duas espécies válidas atualmente, H. multifasciatus e H. taramandahy, ambas endêmicas de riachos do bioma Mata Atlântica. Hollandichthys multifasciatus é encontrada em cursos d'água costeiros entre os estados de Santa Catarina e Rio de Janeiro, e cabeceiras do rio Tietê (bacia do alto rio Paraná), e H. taramandahy é restrita para as bacias dos rios Tramandaí, Mampituba (RS) e Araranguá (SC). São facilmente reconhecidas entre os demais caracídeos deste bioma por apresentar listras pretas horizontais em zigue-zague na lateral do corpo. As duas espécies se diferenciam principalmente pelo padrão de colorido. Hollandichthys taramandahy está ameaçada de extinção na categoria Em Perigo (EN) de acordo com os critérios da IUCN, e corre um risco muito alto de extinção da natureza, principalmente por ser encontrada apenas em ambientes muito específicos e que podem ser impactados pela redução da cobertura vegetal. Aqui são discutidos aspectos relacionados a estratégias reprodutivas, alimentação, diversidade subestimada no grupo e incremento de registros das espécies em coleções científicas. Palavras-chave: Characidae, espécie ameaçada, inseminação, Pseudochalceus, Rachoviscus. Abstract: Hollandichthys, popularly known as "lambari-listrado", is a monophyletic genus of the family Characidae with two valid species, H. multifasciatus and H. taramandahy, both endemic to streams of the Atlantic Forest biome. Hollandichthys multifasciatus is found in coastal watercourses between the states of Santa Catarina and Rio de Janeiro, and waterheads of Tietê River (upper Paraná River basin), and H. taramandahy is restricted to the Tramandaí, Mampituba (RS) and Araranguá (SC) river basins. They are easily recognized among other characids from this biome by possessing zigzag horizontal black stripes on the sides of the body. The two species differ mainly by their color pattern. Hollandichthys taramandahy is threatened with extinction and classified as Endangered (EN) category according to IUCN criteria, with a very high risk of extinction in nature because it is only found in very specific environments that can be impacted by the reduction of vegetation cover. Aspects related to reproductive strategies, feeding, underestimated diversity in the group and increase in species records in scientific collections are discussed.
... Historically, it has been challenging to resolve the phylogenetic relationships in some lineages of the highly diverse Characidae, in part due to their conserved morphologies [14,32,109] or homoplasy of phylogenetic characters [32,78,110]. Our robust phylogenomic hypothesis resolves the placement of H. compressus together with the other trans-Andean species, instead of with the cis-Andean species of the 'rosy tetra' clade where it had been historically grouped [31,32]. ...
Hyphessobrycon is one of the most species rich and widely distributed genera in the family Characidae, with more than 160 species ranging from Veracruz, Mexico to Mar Chiquita Lagoon in Buenos Aires, Argentina. The majority of Hyphessobrycon diversity shows a cis -Andean distribution; only nine species are trans -Andean including H . compressus (Meek 1908). It is well established that Hyphessobrycon is not monophyletic but it has been suggested that natural groups can be identified within the larger Hyphessobrycon species group. In this study, we tested the monophyly of trans -Andean species of Hyphessobrycon and investigated the placement of H . compressus . We inferred the first phylogenomic hypothesis of trans -Andean Hyphessobrycon that includes nearly complete taxonomic sampling (eight of nine valid species) using ultraconserved elements (UCEs). We analyzed 75% (1682 UCEs), 90% (1258 UCEs), and 95% (838 UCEs) complete data matrices, and inferred phylogenomic hypotheses under concatenation and coalescent approaches. In all cases, we recovered the monophyly of trans -Andean Hyphessobrycon inclusive of H . compressus , strong support for three species groups, and evidence of cryptic diversity within the widespread H . compressus and H . condotensis . We used our phylogenomic hypothesis to investigate the biogeographic history of Hyphessobrycon in Middle America. Our ancestral range estimation analysis suggests a single event of cis - to trans -Andean colonization followed by stepwise colonization from the Pacific slope of northwestern South America (Chocó block) to northern Middle America (Maya block). Our work supports the recognition of the trans -Andean species as Hyphessobrycon sensu stricto and provides an evolutionary template to examine morphological characters that will allow us to better understand the diversity of Hyphessobrycon in Middle America.
... Phylogenetic hypotheses recently published based on molecular data or combined morphological and molecular data have confirmed the validity of the genera Knodus and Bryconamericus (Mirande, 2010(Mirande, , 2019Thomaz et al., 2015;García-Melo et al., 2019;Ferreira et al., 2021). Thomaz et al. (2015) presented a phylogeny of Stevardiinae based on molecular data, with the largest number of Knodus species included to date, where the genus was recovered as non-monophyletic, with for example, K. hypopterus (Fowler) being more related to the species of Bryconacidnus Myers (although the type species of Bryconacidnus was not included in the study). ...
Knodus ytuanama, new species, is described from the upper rio Juruena, rio Tapajós drainage, Amazon basin, Mato Grosso, Brazil. The new species differs from its congeners by presenting the interradial membranes of the caudal fin thickened, forming folds, and also differs from most congeners by the presence of a dark, wide midlateral stripe extending from the posterior margin of opercle to the middle caudal-fin rays, the absence of a humeral blotch in adults, and by having four rows of scales between the lateral line and the pelvic-fin origin, among another features. We also provide a discussion on the presence of membranous flaps on the fins as an adaptation for living in fast-water environments in Knodus ytuanama n. sp. as well as in a congener, K. tiquiensis.
Based on ten specimens, using morphometric, meristic and osteological characters, Astyanax leoni n. sp. is described from the upper Madre de Dios River drainage, Amazon River Basin of Peru. A. leoni n. sp. is a member of the Astyanax orthodus species group and is distinguished (except from A. multidens, A. villwocki, and A. gandhiae) in having an inconspicuous rod-shaped stripe on the caudal peduncle that extends anteriorly on top of a silver lateral stripe. It differs from the species mentioned above as follows: from A. multidens by having shorter chevron marks that cover up to only two series of scales above and below the apex of the chevron (vs. chevrons taller, with extensions that cover three series of scales); from A. villwocki by having chevron markings along the entire body (vs. no chevrons on the caudal peduncle); It differs from Astyanax gandhiae in having the caudal peduncle spot extending anteriorly to beyond the tip of the dorsal fin (vs. caudal peduncle spot not reaching anteriorly to tip of dorsal fin), and not of a uniform width as in A. villwocki. A dichotomous key to the species of Astyanax orthodus group is included.
Resumo É descrito um novo peixe caracídeo (Pisces, Cypriniformes, Characoidei) pertencente a um novo gênero Inpaichthys kerri n.g.n sp., coletado perto da Estação do Núcleo Aripuanã do Instituto Nacional de Pesquisas da Amazônia (INPA), ao Norte de Mato Grosso, Brasil. A espécie nova possui linha lateral incompleta e nadadeira caudal nua. Os dentes são tricuspidados, em duas séries muito irregulares no pré-maxilar; a série interna geralmente composta de somente dois dentes medianos em cada lado. A maxila é dentada em metade de sua borda, geralmente com 7 a 8 dentes tricuspidados e cônicos. Esta dentição, em combinação com outras características morfológicas descritas no trabalho, difere o Inpaichthys dos outros gêneros da subfamília Tetragonopterinae.
Astyanax correntinus (Holmberg, 1891) is redescribed with recently collected material from the río Paraná near Corrientes city, northeast of Argentina. This species is distinguished by a combination of characters: (1) the possession of 8/6–7 transverse scales; (2) the presence of 39–42 perforated scales in the lateral series; (3) the presence of iv–v, 29–33 anal-fi n rays; (4) the absence of circuli on posterior fi eld of scales; (5) the broad silvery lateral band; (6) a small rounded humeral spot, and (7) the presence of pentacuspidate outer premaxillary teeth. Several characters are compared with those of Astyanax pelegrini Eigenmann, 1907 from the río Paraguay, in Bahía Negra, Paraguay. Resumen. Astyanax correntinus (Hol mber g, 1891) es redescripta con material recientemente colectado en el Río Paraná cerca de la ciudad de Corrientes, noreste de Argentina. Esta especie es distinguide por una combinación de caracteres: (1) la posesión de 8/6–7 escamas transversas, (2) la presencia de 39–42 escamas perforadas en la serie lateral, (3) la presencia de iv–v, 29–33 radios en la aleta anal, (4) la ausencia de circuli en el campo posterior de las escamas, (5) la banda lateral plateada y ancha, (6) una pequeña mancha humeral redondeada, y (7) la presencia de dientes premaxilares externos pentacúspides. Varios caracteres son comparados con los de Astyanax pelegrini Eigenmann, 1907 procedentes del río Paraguay, en Bahía Negra, Paraguay. Kurzfassung. Astyanax correntinus (HOLMBERG, 1891) wird wieder beschrieben anhand vonneuem Material aus dem río Paraná nahe der Stadt Corrientes im Nordosten Argentiniens. Die Art ist charakterisiert durch eine Kombination von Merkmalen: (1) 8/6–7 Schuppen in einer Querreihe; (2) 39–42 durchbohrte Schuppen in der Seitenlinie; (3) Anale mit iv–v, 29–33 Flossenstrahlen; (4) dem Fehlen von Circuli im hinteren Teil der Schuppen; (5) ein breites silbernes Längsband; (6) ein kleiner runder Schulterfl eck und (7) fünfspitzige Zähne in der äußeren Reihe des Praemaxillare. Einzelne Merkmale werden verglichen mit denen von Astyanax pelegrini EIGENMANN, 1907 vom río Paraguay, Bahía Negra, Paraguay.
Astyanax puka, new species from the endorheic RRo SalR basin, is diagnosed by the following combination of characters: shallow body (33.5–39.8 % SL), 36–38 perforated scales in lateral line, 6/5 transverse scales, 9–10 gradually decreasing dentary teeth, distally expanded premaxillary teeth, one distally expanded maxillary tooth with 7–9 cusps, short upper-jaw length (34.5–37.8 % HL), short maxilla (20.7–23.3 % HL) expanded abruptly under maxillary tooth position, short postorbital distance (36.6–46.9 % HL), short distance between pelvic and anal-fin origins (17.1–20.6 % SL), 22–27 branched anal-fin rays, long caudal peduncle (13.3–16.5 % SL), bony hooks restricted to pelvic-fin rays 2 to 6 and anterior 6–13 branched anal-fin rays on mature males, and the presence of broad denticles on anterior, lateral, and posterior edges of first ceratobranchial gill rakers. Resumen Astyanax puka, nueva especie procedente de la cuenca endorreica del del Río Salí, es diagnosticada por la siguiente com-binaci\n de caracteres: cuerpo bajo (33,5–39,8 % LS), 36–38 escamas perforadas en la línea lateral, 6/5 escamas trans-versas, 9–10 dientes en el dentario gradualmente decrecientes, dientes premaxilares expandidos distalmente, un diente maxilar expandido distalmente con 7–9 cúspides, quijada superior corta (34.5–37.8 % LC), maxilar corto (20.7–23.3 % LC) expandido abruptamente bajo la posici\n del diente maxilar, distancia post-orbitaria corta (36.6–46.9 % LC), distan-cia entre el origen de las aletas pJlvica y anal corta (17.1–20.6 % LS), 22–27 radios ramificados en la aleta anal, pedún-culo caudal largo (13.3–16.5 % LS), ganchos \ seos restringidos a los radios pJlvicos 2 a 6 y los primeros 6–13 radios anales ramificados en machos adultos, y la presencia de dentículos anchos en los bordes anterior, lateral y posterior de los rastrillos branquiales del primer ceratobranquial.
Astyanax puka, new species from the endorheic RRo SalR basin, is diagnosed by the following combination of characters: shallow body (33.5–39.8 % SL), 36–38 perforated scales in lateral line, 6/5 transverse scales, 9–10 gradually decreasing dentary teeth, distally expanded premaxillary teeth, one distally expanded maxillary tooth with 7–9 cusps, short upperjaw length (34.5–37.8 % HL), short maxilla (20.7–23.3 % HL) expanded abruptly under maxillary tooth position, short postorbital distance (36.6–46.9 % HL), short distance between pelvic and anal-fin origins (17.1–20.6 % SL), 22–27 branched anal-fin rays, long caudal peduncle (13.3–16.5 % SL), bony hooks restricted to pelvic-fin rays 2 to 6 and anterior 6–13 branched anal-fin rays on mature males, and the presence of broad denticles on anterior, lateral, and posterior edges of first ceratobranchial gill rakers.
