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Geophagus rufomarginatus, UFRJ 9994, holotype, 96.8 mm SL; Brazil: Bahia: Rio Buranhém Basin. Scale bar 10 mm. Photograph by J.L.O. Mattos.

Geophagus rufomarginatus, UFRJ 9994, holotype, 96.8 mm SL; Brazil: Bahia: Rio Buranhém Basin. Scale bar 10 mm. Photograph by J.L.O. Mattos.

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Morphological characters and phylogenetic trees generated by analyses of segments of two mitochondrial genes cytochrome b and cytochrome c oxidase I support recognition of three new species of the ‘Geophagus’ brasiliensis species group from coastal basins of northeast Brazil. All new species were diagnosed by exclusive morphological characters and...

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... These highly abundant substrate-sifting fishes are widely distributed in rivers from the state of Bahia (northeastern Brazil) to Uruguay (Kullander, 2003;Mattos et al., 2015;Souza et al., 2018). However, a comprehensive phylogenetic analysis of the whole species complex showed that most of their diversity is found along Contas River basin and adjacencies (Argolo et al., 2020), encompassing five of 10 species (including three recently described taxa and one yet to be validated) of remarkable morphological, molecular and cytogenetic variation (Argolo et al., 2020;Mattos et al., 2015, Mattos & Costa, 2018Oliveira et al., 2016;Souza et al., 2018). Therefore, evaluating this fish group throughout such a geographical range fits the requirements to assess putative river basin connectivity processes through time. ...
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Once fishes and rivers are intrinsically connected, historical events such as river captures and the reconfiguration of palaeodrainages affect population structure and species distribution. However, direct evidence of these events remains limited. We aimed to directly infer putative connections between adjacent river basins to understand genetic and geographical patterns in populations and species of the ‘ Geophagus ’ brasiliensis complex. For that, we analysed 174 sequences of the mitochondrial gene cytochrome c oxidase subunit I (COI) from five species along 35 sites within and near a coastal basin in northeastern Brazil. Based on haplotype network, palaeodrainage reconstruction, geological inferences of river captures and analyses of molecular variance (AMOVAs), we tested how the genetic structure could be associated with variable patterns of habitat connectivity through time. In general, high levels of genetic variation were identified, including shared haplotypes among drainages for most species. AMOVA suggested that drainage rearrangements between adjacent basins were the main drivers of genetic diversification in the ‘ G .’ brasiliensis complex. Our study refuted dispersal routes via the continental shelf, whereas eight putative river captures suggested historical connections that enabled gene flow among drainages until recent interruption. These results provide novel insights on the role of riverscape evolution on the biogeography of neotropical ichthyofauna.
... We compare the putative new species with other cichlids based on sequences of the mitochondrial cytochrome b oxidase gene, one of the first and widely used molecular markers to propose hypotheses of phylogenetic relationships (Normark et al., 1991;Farias et al., 2001, Pérez et al., 2007Musilová et al., 2008, López-Fernández et al., 2010, species diversification (Kullander et al., 2010;Piálek et al., 2012;Tougard et al., 2017), and diagnoses of genera or species (López-Fernández et al., 2012;Ottoni & Mattos, 2015;Mattos & Costa, 2018). ...
... The integrative analysis of morphological and molecular features has been highly informative and valuable to study the Cichlidae family, allowing to know new candidate species (Kullander et al., 2010;Piálek et al., 2012, Argolo et al., 2020, improving descriptions, recognizing new taxa such as genera (López-Fernández et al., 2012;Ottoni & Mattos, 2015), delimiting species boundaries (Říčan & Kullander, 2006;Mattos & Costa, 2018;Argolo et al., 2020), and identifying presence of invasive species (dos Santos et al., 2016a,b;Benitez et al., 2018). The utilization of a single methodology, i.e., traditional morphology or molecular approach, could hide these taxa, in which are mostly cryptic taxa and with many taxonomy issues, likewise the species described in the present study. ...
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A new species of Geophagus sensu stricto is described from the Tapajos River basin, Brazil, elevating the number of species of the genus to 21. The new species is of commercial importance and is known in the aquarist trade as Geophagus ‘red head’. The new species is diagnosed using an integrative approach, based on mitochondrial DNA analysis along with morphological evidence. The new species is distinguished from all congeners by the absence of markings on the head, the bar pattern composed by nine vertical bars on the flanks and the presence of distinct longitudinal bands in the caudal fin. Additionally, it shows a genetic distance of at least 2.0% in cytochrome b gene sequences from its closest congeners. Molecular analysis including most genera of Cichlidae from South America corroborates that the new species belongs to the group of Geophagus sensu stricto.
... This fact is more evident in freshwater fi shes, since freshwater systems are isolated, hampering the migration of species from one river system into another. Such a situation favors speciation, in many cases resulting in cryptic species, which is common among Neotropical fi sh lineages, as evidenced and revealed by work dealing with species delimitation molecular approaches (e.g., Costa & Amorim 2011;Costa et al. 2012Costa et al. , 2014Costa et al. , 2017Pereira et al. 2011;Benzaquem et al. 2015;Melo et al. 2014Melo et al. , 2016aMelo et al. , 2016bMelo et al. , 2016cAmorim 2018;Mattos & Costa 2018;Carvalho et al. 2018;Guimarães et al. 2018Guimarães et al. , 2019Rosso et al. 2018;Ottoni et al. 2019;de Santana et al. 2019). Recently, the presence of cryptic species and a greater biodiversity have been also evidenced for the "Rosy tetra" clade (e.g., Castro-Paz et al. 2014;Guimarães et al. 2018Guimarães et al. , 2019, which is herein corroborated, with the description of two new species belonging to this clade, based on seven different species delimitations methods. ...
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Two new species, Hyphessobrycon frickei Guimarães, Brito, Bragança, Katz & Ottoni sp. nov. and H. geryi Guimarães, Brito, Bragança, Katz & Ottoni sp. nov., are herein described, based on seven different and independent species delimitation methods, and on molecular and morphological characters, making the hypothesis of these new species supported from an integrative taxonomy perspective. They belong to the "Rosy tetra" clade, which is mainly characterized by the presence of a dark brown or black blotch on the dorsal fi n and the absence of a midlateral stripe on the body. These two new species are distinguished from the other members of this clade mainly by the arrangement, shape and color pattern of humeral and dorsal-fi n spots, as well as by other characters related to scale counts and body pigmentation. The placement of the new species within the "Rosy tetra" clade was based on the combination of morphological character states mentioned above and corroborated by a molecular phylogenetic analysis using the mitochondrial gene cytochrome oxidase subunit 1. In addition, a new clade (here termed Hyphessobrycon copelandi clade) within the "Rosy tetra" clade is proposed based on molecular data, comprising H. copelandi, H. frickei sp. nov., H. geryi sp. nov. and a still undescribed species. Our results corroborate the occurrence of hidden species within the "Rosy tetra" clade, as suggested by previous studies.
... Current evidence of the lack of knowledge about the ichthyofauna in this region is the large number of taxa considered "sp." or accompanied by the terms "cf." or "aff." in taxonomic inventories (e.g., Sarmento-Soares et al. 2009a, Cetra et al. 2010, Burger et al. 2011, Ramos et al. 2014. Although numerous freshwater fish species from the Bahia river basins have been described in the last two decades (e.g., Bertaco & Lucena 2006, Ribeiro & Lucena 2006, Lima & Britski 2007, Benine et al. 2007, Zanata & Camelier 2008, 2010, 2015, Bichuette et al. 2008, Sarmento-Soares et al. 2009b, Sarmento-Soares et al. 2011, Ferreira et al. 2014, Vari et al. 2010, Bichuette & Rizzato 2012, Oliveira et al. 2013, Sarmento-Soares & Martins-Pinheiro 2013, Camelier & Zanata 2014b, 2017, 2018, Mattos et al. 2015, Peixoto & Wosiacki 2016, Zanata & Pitanga 2016, Zawadzki et al., 2017Barreto et al. 2018, Mattos & Costa, 2018, Burger et al. 2019, Lucena & Lucena 2019, ichthyological explorations continue to reveal unknown species, especially in small tributaries and in upper drainage areas. The increase in the number of species recently described from the São Francisco river basin has already been documented in the literature (see Alves et al. 2011 andBarbosa et al. 2017). ...
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This work was carried out from the assessment of the conservation status of the freshwater ichthyofauna from Bahia State. The inventory data and species distribution were obtained from the specialized scientific literature and representative ichthyological collections. A total of 281 native species was recorded in Bahia State, distributed in the Northeastern Mata Atlantica (NMA) and São Francisco (SFR) freshwater ecoregions. There was a larger number of species in the NMA (187 spp.), composed by several coastal basins, than in the SFR (134 spp.), composed by São Francisco river basin. Among the 30 families recorded, Characidae and Rivulidae were the most representative, with 53 and 48 species, respectively. The conservation status of 214 species was assessed and 33 of them (15%) were included in the IUCN threat categories. Of these, 11 species were classified as vulnerable (VU), 12 as endangered (EN), and 10 as critically endangered (CR). Most threatened species (n = 14) belongs to the family Rivulidae. The larger number of threatened species in the NMA: (n = 23) is mainly related to the high endemism of restricted-range species associated with the human occupation impacts along the coastal regions. In the SFR, most of threatened species are annual killifishes, which are locally disappearing due to increasing degradation of their temporary habitats.
... More recently, several studies reported high levels of genetic and morphological variation in 'G.' brasiliensis from various Brazilian river basins (e.g. Alves-Silva and Dergam, 2015;Ferreira et al., 2016;Oliveira et al., 2016;Souza et al., 2018), including the description of four new species: 'G.' diamantinensis Mattos et al., 2015;'G.' santosi Mattos and Costa, 2018;'G.' rufomarginatus Mattos and Costa, 2018;and'G.' multiocellus Mattos andCosta, 2018 (Mattos et al., 2015;Mattos and Costa, 2018), bringing the genus to the eight currently recognized species. ...
... In recent species descriptions, the mitochondrial DNA (mtDNA) gene Cytochrome C Oxidase Subunit I (COI) has been used as barcodes to differentiate species (Hebert et al., 2003), including analyses within the 'G.' brasiliensis complex (Mattos and Costa, 2018;Souza et al., 2018). This approach, along with the development of several singlelocus delimitation methods (e.g. ...
... The type series of 'G.' brasiliensis is supposed to have been collected at a type locality in Rio de Janeiro city, but the site has been filled for land reclamation, so we sampled several locations throughout Rio de Janeiro state to increase the representation of its variation. Finally, the data gathered from online databases (GenBank and BOLD) encompassed sequences from type localities used in the descriptions of 'G.' rufomarginatus, 'G.' diamantinensis, 'G.' multiocellus and 'G.' santosi (Mattos et al., 2015;Mattos and Costa, 2018) as well as other species and lineages from several localities (Supplementary Table S1). ...
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The ‘Geophagus’ brasiliensis complex is one of the most abundant groups of cichlids from eastern coastal basins in South America. Traditionally, this fish group has been recognized as incertae sedis because of phylogenetic uncertainties and unclear taxonomy. In addition, the remarkable morphological, chromosomal, and DNA variation reported over recent years in several populations of these cichlids has increased the debate about their species richness and their distributional range. Here, we tested the presence of independent evolutionary lineages within the ‘G.’ brasiliensis complex, addressing their taxonomic status and evolutionary relationships, including a comparative analysis of genetic and morphological patterns, based on an extensive dataset, comprising 172 sampling sites along most of their known range using a mitochondrial marker, RADseq data and geometric morphometrics. The number of putative species in the present study varied from 9 to 11 depending on the molecular species delimitation methods used. Our results revealed at least two putative new taxa (‘Geophagus’ sp. Doce and ‘Geophagus’ sp. Upper Contas). Morphometric analyses, particularly those based on Canonical Variate Analysis (CVA), revealed significant morphological differentiation between species within the main clades. On the other hand, analyses of morphological phylogenetic signal and phylomorphospace provided no evidence of adaptive differentiation among these species. Thus, diversification in the ‘G.’ brasiliensis complex seems to have been influenced by hydrogeological events that promoted allopatry, such as the presence of paleodrainages and distributional reconfiguration through river captures. We propose major changes in the known distribution of some species within the complex and conservatively suggest the recognition of 10 species within the ‘Geophagus’ brasiliensis complex, with the potential for further dividing ‘G.’ rufomarginatus after additional taxonomic evaluation.
... Geophagus gr. brasiliensis fits in the G. brasiliensis species group, which currently includes eight species according to Mattos & Costa (2018), being six of them known from NMAF ecoregion. Geophagus obscurus (Castelnau 1855) and G. itapicuruensis Haseman 1911 were the first two species of the genus described from this region. ...
... Geophagus obscurus (Castelnau 1855) and G. itapicuruensis Haseman 1911 were the first two species of the genus described from this region. Recently, four new species included in the G. brasiliensis complex were also described, G. diamantinensis Mattos Mattos & Costa 2018. Herein, the species analyzed is tentatively identified as G. gr. ...
... brasiliensis due to the character overlap with the aforementioned species, for example, the number of branched rays of all fins. In addition, some of these species were described by Mattos & Costa (2018) using mainly color pattern in life and molecular data, which makes it difficult to perform morphological comparisons based on the literature. ...
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... Similar to the A. autrani group, in that several species occur along the latitudinal zonation of eastern Brazil, a congruent distribution pattern and endemism is also shared by other freshwater fish groups in this region, such as: Delturinae (family Loricariidae) (Reis et al. 2006;Buckup 2011), Neoplecostominae (family Loricariidae) (Abell et al. 2008;Buckup 2011;Roxo et al. 2014), the genera Mimagoniates Regan, 1907 and Oligosarcus Günther, 1864 (family Characidae) (Menezes et al. 2007(Menezes et al. , 2008Buckup 2011;Ribeiro and Menezes 2015), the genus Brycon Müller & Troschel, 1844 (Bryconidae) (Menezes et al. 2007;Abe et al. 2014;Travenzoli et al. 2015), the genus Characidium Reinhardt, 1867 (Crenuchidae) (Menezes et al. 2007;Poveda-Martínez et al. 2016), the genera Listrura de Pinna, 1988, Microcambeva Costa &Bockmann, 1994, andTrichomycterus Valenciennes, 1832 (family Trichomycteridae) (Barbosa and Costa 2010;Abell et al. 2008;Buckup 2011;Mattos and Lima 2012;Villa-Verde et al. 2013;Katz and Barbosa 2014), the genus Microglanis Eigenmann, 1912 (family Pseudopimelodidae) (Sarmento-Soares et al. 2006;Ruiz and Shibatta 2010;Mattos et al. 2013), several killifish genera (family Aplocheiliidae) (Abell et al. 2008;Buckup 2011;Costa 2014, Costa and, the genus Phalloceros Eigenmann, 1907 (family Poeciliidae) (Lucinda 2008), and the "Geophagus brasiliensis" species group and the genus Crenicichla Heckel, 1840 (family Cichlidae) (Kullander and Lucena 2006;Mattos 2014;Mattos et al. 2015;Mattos and Costa 2018). ...
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Three distinct and independent molecular-based species delimitation analyses were performed among the species and populations included within the Australoheros autrani group, based on sequences of the mitochondrial gene Cytochrome b: a tree-based method proposed by Wiens and Penkrot (WP), a Character-based DNA Barcoding (CBB) and co-alescent species delimitation method termed the Bayesian Implementation of the Poisson tree processes (bPTP). The congruence of WP and CBB delimited 11 independent lineages (species), while the bPTP delimited just nine lineages. We did not favour any of the methods , and we considered the possibility of two slightly variant scenarios. A time-calibrated phylogenetic analysis is proposed based on the predominant congruence of the results of these three species delimitation methods herein applied. The monophyly of the A. autrani species group was highly supported with maximum node support value and diagnosed by 11 nucleotide substitutions. The sister clade of the A. autrani species group is the clade comprising A. sp. Timbé do Sul and A. minuano. The phylogenetic analysis supports three main clades within the A. autrani species group, supported by maximum node support value, with the Southern Mata Atlântica clade as the most basal clade. Divergence time estimates indicate that the diversification of the Australoheros originated during the early Neogene, but only in the late Neogene did the processes of diversification in the southeast and north regions occur. Diversification within the Australoheros autrani species group occurred synchronically for the three main clades during the beginning of the Quaternary. It is demonstrated that molecular characters are valuable tools for species recognition, particularly in speciose groups with inconspicuous or difficult to record morphological characters. The resulting phylogeny of the Australoheros autrani group is highly compatible with the geological and biogeographic scenarios proposed for the Neogene and Quarternary shaping of the extant river basins of eastern Brazil. Despite the origin of the A. autrani group being dated to the late Miocene, species level diversification occurred in the Pleistocene and was probably driven by headwater capture events and sea-level fluctuations.
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A new species of Astyanax from tributaries of the rio de Contas, Bahia, Brazil, is described. The new species differs from congeners by having three horizontal series of scales from lateral line to pelvic-fin origin and the distal margin of third infraorbital distinctly separated from vertical and horizontal limbs of preopercle, leaving a broad area not covered by superficial bones. The new species further differs from most congeners by the presence of bony hooks on all fins of mature males. Particularly from congeners occurring in rivers of the Northeastern Mata Atlântica freshwater ecoregion, it further differs by having the highest body depth just anterior to the dorsal-fin origin, 34-37 pored lateral line scales, a vertically elongated conspicuous dark humeral blotch reaching below the lateral line and a conspicuous dark wide midlateral stripe extending from the clear area on the rear of the humeral blotch to the end of middle caudal-fin rays and forming an inconspicuous blotch on caudal peduncle.