Accelerated Diversification Explains the Exceptional Species Richness of Tropical Characoid Fishes.

The Neotropics harbor the most species-rich freshwater fish fauna on the planet, but the timing of that exceptional diversification remains unclear. Did the Neotropics accumulate species steadily throughout their long history, or attain their remarkable diversity recently? Biologists have long debated the relative support for these museum and cradle hypotheses, but few phylogenies of megadiverse tropical clades have included sufficient taxa to distinguish between them. We used 1288 ultraconserved element loci spanning 293 species, 211 genera, and 21 families of characoid fishes to reconstruct a new, fossil-calibrated phylogeny and infer the most likely diversification scenario for a clade that includes a third of Neotropical fish diversity. This phylogeny implies paraphyly of the traditional delimitation of Characiformes because it resolves the largely Neotropical Characoidei as the sister lineage of Siluriformes (catfishes), rather than the African Citharinodei. Time-calibrated phylogenies indicate an ancient origin of major characoid lineages and reveal a much more recent emergence of most characoid species. Diversification rate analyses infer increased speciation and decreased extinction rates during the Oligocene at around 30 Ma during a period of mega-wetland formation in the proto-Orinoco-Amazonas. Three species-rich and ecomorphologically diverse lineages (Anostomidae, Serrasalmidae, and Characidae) that originated more than 60 Ma in the Paleocene experienced particularly notable bursts of Oligocene diversification and now account collectively for 68% of the approximately 2150 species of Characoidei. In addition to paleogeographic changes, we discuss potential accelerants of diversification in these three lineages. While the Neotropics accumulated a museum of ecomorphologically diverse characoid lineages long ago, this geologically dynamic region also cradled a much more recent birth of remarkable species-level diversity. [Biodiversity; Characiformes; macroevolution; Neotropics; phylogenomics; ultraconserved elements.].

Bunocephalus hertzi, a new banjo catfish from the upper Rio Paraná basin, Brazil (Siluriformes: Aspredinidae), with the redescription of Bunocephalus larai Ihering, 1930.

Bunocephalus hertzi, new species, is described from two tributaries of the upper Rio Paraná basin. Because the type series of B. larai (the only other species of Bunocephalus in the basin) is lost, a redescription of B. larai is also provided herein. Bunocephalus larai and B. hertzi can be distinguished from all its congeners, except B. minerim, by the absence of an epiphyseal bar (vs. present). From B. minerim, both species can be distinguished by the presence of 10 principal caudal-fin rays (vs. 9). Bunocephalus hertzi can be differentiated from B. larai by the longer coracoid process size (11.9-16.2% vs. 6.7-9.8% SL), shorter maxillary barbel (just reaching the pectoral fin origin; 54.6-67.4% vs. 73.7-84.6% HL) and skull ornamentation represented by five bony protuberances between the posterior end of the occipital and the origin of the dorsal fin (vs. four bony protuberances).

The brain of Brycon orbignyanus (Valenciennes, 1850) (Teleostei: Characiformes: Bryconidae): gross morphology and phylogenetic considerations

The brain of Brycon orbignyanus is described as a model for future studies of the gross morphology of the central nervous system in Characiformes. The study of brain gross morphology of 48 distinct taxa of Characiformes, one of Cypriniformes, two of Siluriformes and two of Gymnotiformes, allowed us to propose, for the first time, six putative brain synapomorphies for the Characiformes and also two possibly unique gross brain morphology characters for the Siluriformes. A detailed protocol for the extraction of the brain in Characiformes is also provided.(AU)

O encéfalo de Brycon orbignyanus é descrito como um modelo para futuros estudos da anatomia externa do Sistema Nervoso Central de Characiformes. O estudo da morfologia externa de 48 táxons distintos de Characiformes, um de Cypriniformes, dois de Siluriformes e dois de Gymnotiformes, permitiu-nos propor, pela primeira vez, seis prováveis sinapomorfias encefálicas e também duas possíveis características encefálicas para Siluriformes. Um protocolo detalhado para a dissecção e extração do encéfalo de Characiformes é também apresentado.(AU)

Taxonomic revision and molecular phylogeny of Gymnocorymbus Eigenmann, 1908 (Teleostei, Characiformes, Characidae).

Characidae, one of the most diverse families of Characiformes and one of the largest clades of fishes in the world, has a complex taxonomic background, with one issue being the placement of various genera, including Gymnocorymbus. Herein, we generate the first molecular phylogeny for the genus using three nuclear and two mitochondrial loci and review the systematics of Gymnocorymbus. This genus includes the black tetra, a well-known and popular species among aquarists. Molecular phylogeny strongly supports the monophyly of Gymnocorymbus, with this hypothesis corroborated by the presence of three morphological synapomorphies. Of the six previously known nominal species of Gymnocorymbus, three are considered valid herein: Gymnocorymbus bondi from the Río Orinoco basin, Gymnocorymbus ternetzi from the Rio Paraguay basin, and Gymnocorymbus thayeri from the Corantijn and Amazon basin and rivers of northeastern Brazil. A fourth species, Gymnocorymbus flaviolimai sp. n., is described from the Rio Madeira, Amazon basin. Lectotypes are designated for G. ternetzi and G. thayeri. Our results support previous hypotheses of the alignment of Gymnocorymbus close to the subfamily Stethaprioninae and also support the sister relationship between G. ternetzi and G. thayeri, and of that clade as sister to G. bondi.

Taxonomy of probolodus eigenmann, 1911 (characiformes: characidae) with description of two new species, and comments about the phylogenetic relationships and biogeography of the genus

Probolodus is a genus of tetras distributed along the coastal basins of southeastern Brazil, from Espírito Santo on the north, to São Paulo states. It was proposed by Eigenmann in 1911, to include the single species P. heterostomus. However, examining of material recently collected we identified two new species: P. oyakawai, new species, from the rio Ribeira de Iguape basin, and P. sazimai, new species, from the rio Itapemirim (Espírito Santo), and drainages of the rio Doce basin (Espírito Santo and Minas Gerais states). Therefore, the distribution of P. heterostomus was restricted to the rio Paraíba do Sul basin, southeastern Brazil. The genus Probolodus was redefined based on osteological characters, in particular details of oral dentition. Probolodus heterostomus differs from its congeners by a combination of meristic and osteological characters. An identification key for the species of the genus is presented. The possible evolutionary relationships of Probolodus with other characids, especially some lepidophagous taxa are discussed. Regarding the biogeography of the group, some geological events are hypothesized as responsible for the cladogenetic events among species of the genus.

Probolodus é um gênero de lambari distribuído ao longo das bacias costeiras do sudeste brasileiro, do Espírito Santo até São Paulo. Foi proposto originalmente por Eigenmann em 1911, para abrigar a única espécie P. heterostomus. Entretanto, através do exame de material coletado recentemente, identificamos duas novas espécies do gênero: P. oyakawai, espécie nova, da bacia do rio Ribeira de Iguape, e P. sazimai, espécie nova, das drenagens das bacias dos rios Itapemirim (Espírito Santo) e Doce (Espírito Santo e Minas Gerais). Portanto, a distribuição de P. heterostomus foi restringida à bacia do rio Paraíba do Sul, sudeste do Brasil. O gênero Probolodus foi redefinido a partir de caracteres osteológicos, em especial detalhes da dentição. Probolodus heterostomus difere de suas duas congêneres por uma combinação de caracteres merísticos e osteológicos. Uma chave de identificação do gênero é apresentada. As possíveis relações evolutivas de Probolodus com outros caracídeos, em especial alguns táxons lepidófagos examinados neste estudo, são discutidas. Com relação à biogeografia do gênero, alguns eventos geológicos são postulados como possivelmente responsáveis pela sua cladogênese.

A new species of utiaritichthys miranda ribeiro (characiformes: serrasalmidae) from the serra dos parecis, Tapajós drainage

Utiaritichthys esguiceroi is described from the upper portion of the rio Juruena, rio Tapajós drainage, Mato Grosso State, Central Brazil. The new species distinguished from its two congeners, U. sennaebragai Miranda Ribeiro and U. longidorsalis Jégu, Tito de Morais & Santos, by having 99 to 101 perforated scales on lateral line (vs. 69 to 83), presence of 17 to19 prepelvic spines (vs. 9-13 in U. sennaebragai and 28-31 in U. longidorsalis), 20 to 21 postpelvic spines (vs. 15 to 19 in U. sennaebragai, and 14 in U. longidorsalis), and 23 to 25 circumpeduncular scales (vs. 30-48 in U. sennaebragai, and 33-35 in U. longidorsalis). Furthermore, the new species differs from U. longidorsalis by having larger interdorsal width, and adipose-fin base length (11.8-15.6 vs. 7.1-7.9% of SL, and 4.2-5.8 vs. 3.7-3.8% of SL, respectively).

Utiaritichthys esguiceroi é descrita da porção superior do rio Juruena, bacia do rio Tapajós, estado do Mato Grosso, Brasil Central. A espécie nova diferencia-se de suas congêneres, U. sennaebragai Miranda Ribeiro e U. longidorsalis Jégu, Tito de Morais & Santos, pela presença de 99 a 101 escamas perfuradas na linha lateral (vs. 69 a 83), presença de 17 a 19 espinhos pré-pélvicos (vs. 9-13 em U. sennaebragai e 28-31 em U. longidorsalis), 20 a 21 espinhos pós-pélvicos (vs. 15 a 19 em U. sennaebragai e 14 em U. longidorsalis), e 23-25 escamas da série circumpeduncular (vs. 30-48 em U. sennaebragai e 33-35 em U. longidorsalis). Em adição, a espécie nova difere de U. longidorsalis por possuir a largura interdosal e comprimento da base da nadadeira adiposa maiores (11,8-15,6 vs. 7,1-7,9% do CP e 4,2-5,8 vs. 3,7-3,8% do CP, respectivamente).

Multilocus molecular phylogeny of Gasteropelecidae (Ostariophysi: Characiformes) reveals the existence of an unsuspected diversity.

The characiform family Gasteropelecidae, the so-called freshwater hatchetfishes, is comprised of three genera and nine species found in Panama and all South American countries except Chile. Our goal was to investigate the molecular characteristics, phylogenetic relationships among the species and genera of Gasteropelecidae and phylogenetic relationships between the Gasteropelecidae family with other Characiformes. DNA fragments from two mitochondrial (16S rRNA and Cytochrome B) and three nuclear genes (Rag1, Rag2 and Myh6) were sequenced. Our results corroborate the morphology-based hypothesized monophyly of the Gasteropelecidae family and most of the relationships among its genera. However, the genus Gasteropelecus is polyphyletic because G. maculatus is placed as the sister group to all other gasteropelecids, whereas G. sternicla is more closely related to species of Carnegiella. Similarly, the species Carnegiella strigata is not monophyletic, which suggests that the family needs a taxonomic review. Moreover, the species Thoracocharax stellatus was composed by four distinct lineages suggesting the this species may represents a species complex.

Anatomy and evolution of the mandibular, hyopalatine, and opercular muscles in characiform fishes (Teleostei: Ostariophysi).

The Characiformes are distributed throughout large portions of the freshwaters of Africa and America. About 90% of the almost 2000 characiform species inhabit the American rivers, with their greatest diversity occurring in the Neotropical region. As in most other groups of fishes, the current knowledge about characiform myology is extremely poor. This study presents the results of a survey of the mandibular, hyopalatine, and opercular musculature of 65 species representing all the 18 traditionally recognized characiform families, including the 14 subfamilies and several genera incertae sedis of the Characidae, the most speciose family of the order. The morphological variation of these muscles across the order is documented in detail and the homologies of the characiform adductor mandibulae divisions are clarified. Accordingly, the mistaken nomenclature previously applied to these divisions in some characiform taxa is herein corrected. Contradicting some previous studies, we found that none of the examined characiforms lacks an A3 section of the adductor mandibulae, but instead some taxa have an A3 continuous with A2. Derived myological features are identified as new putative synapomorphies for: the Characoidei; the clade composed of the Alestidae, Characidae, Gasteropelecidae, Cynodontoidea, and Erythrinoidea; the clade Cynodontoidea plus Erythrinoidea; the clade formed by Ctenoluciidae and Erythrinidae; the Serrasalminae; and the Triportheinae. Additionally, new myological data seems to indicate that the Agoniatinae might be more closely related to cynodontoids and erythrinoids than to other characids.

The blind catfish from the caves of Chapada Diamantina, Bahia, Brazil (Siluriformes: Heptapteridae): description, anatomy, phylogenetic relationships, natural history, and biogeography

Rhamdiopsis krugi, a new troglobitic heptapterid catfish, is described from the caves of Chapada Diamantina, State of Bahia, northeastern Brazil. This species, although frequently cited in the scientific literature along the last seventeen years, remained undescribed largely due to its uncertain phylogenetic affinities. The generic assignment of R. krugi was clouded largely by its high number of unusual morphological features (some related to cave life), for instance: absence of eyes and body pigmentation; presence of a widely exposed pseudotympanum; posterior border of the anterior branch and anterior margin of the arborescent portion of the posterior branch of the transverse process of fourth vertebra joined; dorsal hypural plate commonly with seven rays; ventral caudal plate usually with six rays; dorsal and ventral caudal-fin lobes typically with six branched fin rays each; 38-39 vertebrae; anal fin with 14-17 rays; and lateral line very short. Rhamdiopsis krugi can be easily distinguished from its congeners, R. microcephala and R. moreirai, by its troglomorphic features and by the presence of a shorter lateral line, fewer vertebrae and anal-fin rays, pattern of branching of caudal-fin rays, and several attributes of skeletal system. The affinities of this new species are discussed in light of current phylogenetic knowledge of the family Heptapteridae. Incongruent derived characters do not allow selection of a particular hypothesis of sister group relationships among species of Rhamdiopsis. The occurrence of R. krugi in the rio Paraguaçu basin is possibly due to an event of hydrological capture from a section of the middle portion of the rio São Francisco basin, caused by tectonic events. The semi-arid region where R. krugi presently lives was probably covered by a wide forested area during a humid cycle in Quaternary. A summary of natural history and ecology data of R. krugi, as well as notes on its conservation, are provided. We also offer comments on the morphological plasticity of R. krugi.

Rhamdiopsis krugi, um novo bagre heptapterídeo troglóbio, é descrito de cavernas da Chapada Diamantina, Estado da Bahia, nordeste do Brasil. Esta espécie, embora frequentemente citada na literatura ao longo dos últimos dezessete anos, não foi descrita antes em função das suas afinidades filogenéticas incertas. A posição genérica de R. krugi foi obscurecida principalmente pelo seu alto número de caracteres morfológicos incomuns (parte deles relacionada à vida nas cavernas), como por exemplo: ausência de olhos e de pigmentação corporal; presença de um pseudotímpano amplamente exposto; borda posterior do ramo anterior e margem anterior da porção arborescente do ramo posterior do processo transverso da quarta vértebra conectados um ao outro; placa hipural dorsal normalmente com sete raios; placa caudal ventral usualmente com seis raios; lobos dorsal e ventral da nadadeira caudal tipicamente com seis raios ramificados cada; 38-39 vértebras; nadadeira anal com 14-17 raios; e linha lateral muito curta. Rhamdiopsis krugi pode ser facilmente distinguida de seus congêneres, R. microcephala e R. moreirai, por seus caracteres troglomórficos e pela presença de uma linha lateral mais curta, menos vértebras e raios na nadadeira anal, padrão de ramificação dos raios da nadadeira caudal e vários atributos do sistema esquelético. As afinidades desta nova espécie são discutidas à luz do conhecimento atual sobre a filogenia da família Heptapteridae. Caracteres derivados incongruentes não permitem optar por uma hipótese particular de relação de grupo-irmão entre as espécies de Rhamdiopsis. A ocorrência de R. krugi na bacia do rio Paraguaçu é devida, possivelmente, a um evento de captura hidrológica de uma seção da porção média da bacia do rio São Francisco, ocasionada por eventos tectônicos. A região semi-árida onde R. krugi atualmente vive estava provavelmente coberta por uma ampla floresta durante um ciclo úmido no Quaternário. Um sumário das informações sobre a história natural e ecologia de R. krugi, assim como observações sobre sua conservação, são apresentadas. Apresentamos também comentários sobre a plasticidade morfológica de R. krugi.

A new Moenkhausia Eigenmann, 1903 (Ostariophysi: Characiformes) from Chapada Diamantina, rio Paraguaçu Basin, Bahia, Northeastern Brazil

Moenkhausia diamantina, new species, is described from tributaries of the rio Paraguaçu, BA, northeastern Brazil. This species is distinguished from all congeners by features of body color pattern, the presence of scales on the predorsal median line and the number of anal-fin rays

Moenkhausia diamantina, nova espécie, é descrita de tributário do rio Paraguaçu, BA, nordeste do Brasil. Esta espécie é distinguida das demais congêneres por caracteres relacionados ao padrão de coloração do corpo, pela presença de escamas na linha mediana pré-dorsal e número de raios da nadadeira anal