Figure 1
The superfamily Anostomoidea with selected species representing a small amount of its diversity. (a), Leporinus brunneus (Anostomidae), ANSP 190763, 62 mm standard length (SL). (b), Leporinus maculatus (Anostomidae), FMNH ex ANSP 189041, 123.4 mm SL. (c), Ichthyoelephas longirostris (Prochilodontidae), ANSP 192865, ca. 470 mm SL. (d), Semaprochilodus varii (Prochilodontidae), ANSP 187435, 210 mm SL. (e), Curimatopsis crypticus (Curimatidae), male, ANSP 189091, 28.8 mm SL. (f), Caenotropus maculosus (Chilodontidae), ANSP 189147, 70 mm SL. All photographs by Mark Sabaj Pérez and used with permission.
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Although 11 studies have addressed the systematics of the four families and 281 fish species of the ecomorphologically diverse Anostomoidea, none has proposed a global hypothesis of relationships. We synthesized these studies to yield a supermatrix with 463 morphological characters among 174 ingroup species, and inferred phylogeny with parsimony an...
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... intrageneric hypothesis by Netto-Ferreira and Vari (2011). Each of these nodes was supported by a single synapomorphy under the earlier hypotheses, and the collapse under the global analysis ( Fig. 2) results from synonymization of char- acters or character states to maintain coding consis- tency across the supermatrix. Bayesian reconstruction (Fig. S1) recovered the same interfamilial topology as parsimony (see Supporting Information for more details on Bayesian ...
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Atypical Mycobacteriosis is a chronic and progressive disease of fish affecting all species (freshwater, brackish water and saltwater); the disease has been documented in more than 160 species worldwide [[]Chinabut, 1999]. Mycobacterium marinum, M. fortuitum and M. chelonae are the most commonly isolated bacterial species Apart from the mortality i...
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... The FISH data set was obtained from Dillman et al. (2016) and includes information for four families of anostomoid fishes in the order Characiformes (Actinopterygii: Ostariophysi: Characiformes). The original matrix was reduced so as to contain only 10 taxa and retaining 463 characters, each annotated with anatomical terms from the Uberon ontology (see supporting data in the Supplementary Material available on Dryad). ...
... Two or more species representing the four anostomoid families (i.e., Anostomidae, Chilodontidae, Curimatidae, and Prochilodontidae) were selected, along with one outgroup taxon (Parodontidae: Parodon). The taxon sampling represents the diversity among the main lineages of anostomoid fishes (e.g., see Dillman et al. 2016). This data set was analyzed under the ALL alternative of subset construction. ...
Morphology remains a primary source of phylogenetic information for many groups of organisms, and the only one for most fossil taxa. Organismal anatomy is not a collection of randomly assembled and independent ‘parts’, but instead a set of dependent and hierarchically nested entities resulting from ontogeny and phylogeny. How do we make sense of these dependent and at times redundant characters? One promising approach is using ontologies—structured controlled vocabularies that summarize knowledge about different properties of anatomical entities, including developmental and structural dependencies. Here we assess whether evolutionary patterns can explain the proximity of ontology-annotated characters within an ontology. To do so, we measure phylogenetic information across characters and evaluate if it matches the hierarchical structure given by ontological knowledge—in much the same way as across-species diversity structure is given by phylogeny. We implement an approach to evaluate the Bayesian phylogenetic information (BPI) content and phylogenetic dissonance among ontology-annotated anatomical data subsets. We applied this to datasets representing two disparate animal groups: bees (Hexapoda: Hymenoptera: Apoidea, 209 chars) and characiform fishes (Actinopterygii: Ostariophysi: Characiformes, 463 chars). For bees, we find that BPI is not substantially explained by anatomy since dissonance is often high among morphologically related anatomical entities. For fishes, we find substantial information for two clusters of anatomical entities instantiating concepts from the jaws and branchial arch bones, but among-subset information decreases and dissonance increases substantially moving to higher level subsets in the ontology. We further applied our approach to address particular evolutionary hypotheses with an example of morphological evolution in miniature fishes. While we show that phylogenetic information does match ontology structure for some anatomical entities, additional relationships and processes, such as convergence, likely play a substantial role in explaining BPI and dissonance, and merit future investigation. Our work demonstrates how complex morphological datasets can be interrogated with ontologies by allowing one to access how information is spread hierarchically across anatomical concepts, how congruent this information is, and what sorts of processes may play a role in explaining it: phylogeny, development, or convergence.
... Even with 46% missing entries in the molecular partition, the combined analysis yielded better support and resolution of the topology than when morphological and molecular data were analysed separately. Flynn et al. (2005) suggested that excluding missing data might generate a decrease in nodal support and resolution, and many works have demonstrated that the amount of missing data is not a problem as long as enough informative characters (Maddison, 1993;Wiens, 1998Wiens, , 2003Wiens, , 2006 and monophyletic lineages (Graybeal, 1998;Rannala et al., 1998;Dillman et al., 2015;Mirande, 2019) are included in the cladistic analysis. Several authors have argued that the effects of missing data are not absolute and are dependent of the phylogenetic signal, numbers of terminal taxa and characters, and sampling of close relatives of taxa with little information available (e.g. ...
This is the first study to combine morphological and molecular characters to infer the phylogenetic relationships among catsharks. All currently valid genera classified in the family Scyliorhinidae s.l. and representatives of other carcharhinoid families plus one lamnoid and two orectoloboids were included as terminal taxa. A total of 143 morphological characters and 44 NADH2 sequences were analysed. Parsimony analyses under different weighting schemes and strengths were used to generate hypotheses of phylogenetic relationships. The phylogenetic analysis of 78 terminal taxa, using the combined dataset and weighting each column separately (SEP; k = 3) resulted in one most-parsimonious cladogram of 4441 steps with the greatest internal resolution of clades and strongest support. The main changes in nomenclature and classification are the revised definition and scope of Scyliorhinidae, Apristurus and Pentanchus and the revalidation of Atelomycteridae. The monophyly of Pentanchidae is supported, as is that of most catshark genera. Two new subfamilies of the family Pentanchidae are defined: Halaelurinae subfam. nov. and Galeinae subfam. nov. Our analysis emphasizes the relevance of morphological characters in the inference of evolutionary history of carcharhinoids and sheds light on the taxonomic status of some genera in need of further exploration.
... Curimatids are important parts of the fish faunas in the lowland rivers of the Amazon, Orinoco, and La Plata basins, upland rivers of the Guiana and Brazilian shields (e.g., Essequibo, Tocantins, São Francisco, and Paraná basins), and swiftly flowing rivers west of the Andes (e.g., Maracaibo, Magdalena, Atrato and Pacific coastal basins from northern Peru to Costa Rica) (Frable, 2017;Vari, 1988). The alpha taxonomy and higher-level systematics of curimatids have attracted substantial attention (e.g., Vari et al., 2012;Melo, 2017Melo, , 2020Melo & Oliveira, 2017;Bortolo et al., 2018) using both morphological and molecular evidence (Dillman et al., 2016;Dorini et al., 2020;Melo et al., 2016Melo et al., , 2018Vari, 1989). The family is represented by 117 valid species in eight extant genera, representing one of the ten most species-rich families in the Amazon basin (Dagosta & de Pinna, 2019;Fricke et al., 2021;Vari, 1989). ...
The Neotropics harbors a megadiverse ichthyofauna comprising over 6300 species with approximately 80% in just three taxonomic orders within the clade Characiphysi. This highly diverse group has evolved in tropical South America over tens to hundreds of millions of years influenced mostly by re-arrangements of river drainages in lowland and upland systems. In this study, we investigate patterns of spatial diversification in Neotropical freshwater fishes in the family Curimatidae, a species-rich clade of the order Characiformes. Specifically, we examined ancestral areas, dispersal events, and shifts in species richness using spatially explicit biogeographic and macroevolutionary models to determine whether lowlands–uplands serve as museums or cradles of diversification for curimatids. We used fossil information to estimate divergence times in BEAST, multiple time-stratified models of geographic range evolution in BioGeoBEARS, and alternative models of geographic state-dependent speciation and extinction in GeoHiSSE. Our results suggest that the most recent common ancestor of curimatids originated in the Late Cretaceous likely in lowland paleodrainages of northwestern South America. Dispersals from lowland to upland river basins of the Brazilian and Guiana shields occurred repeatedly across independently evolving lineages in the Cenozoic. Colonization of upland drainages was often coupled with increased rates of net diversification in species-rich genera such as Cyphocharax and Steindachnerina. Our findings demonstrate that colonization of novel aquatic environments at higher elevations is associated with an increased rate of diversification, although this pattern is clade-dependent and driven mostly by allopatric speciation. Curimatids reinforce an emerging perspective that Amazonian lowlands act as a museum by accumulating species along time, whereas the transitions to uplands stimulate higher net diversification rates and lineage diversification.
... In the present study, the nominal species "Eigenmannia" goajira was included in the analysis only using external characters and osteological features accessed via radiographs or available in the literature, representing 53.5% of the character list (77 of 144), and 46.5% missing data. In spite of that, as discussed by Dillman et al. (2015), the inclusion of taxa with large amount of missing data in morphological-based phylogenies has provided valuable results to recover the phylogenetic position of taxa with uncertain relationships. These authors, however, reinforced that additional analyses are necessary to determine the threshold of missing data that allows the recovery of a reasonable phylogenetic position. ...
We advance on the knowledge of Eigenmanniinae by proposing a hypothesis of phylogenetic relationships based on the parsimony analysis of a diverse set of 144 anatomical characters, 12% of them treated as quantitative and 88% treated as qualitative (8% external morphology, 51% osteology, 21% myology, and 8% neuroanatomy). Thirty‐seven of 45 valid species of Eigenmanniinae are examined in the study, including the incertae sedis species “Eigenmannia” goajira. The final tree yields new insights on species relationships, thus, producing a new classification to Eigenmanninae. Our analysis recovered the monophyly of Eigenmanniinae, Archolaemus, Eigenmannia, and Rhabdolichops. Eigenmannia is proposed as monophyletic based on four morphological synapomorphies, one of which exclusive to the genus. Japigny is proposed to be the sister group of all remaining Eigenmanniinae and “E.” goajira to be the sister group of Archolaemus. The hypothesis of monophyly of Distocyclus including D. conirostris and D. guchereauae is rejected. Consequently, D. guchereauae is included in Eigenmannia, and a new genus is established to include “E.” goajira. A taxonomic key to all genera is provided. In addition, this study highlights the critical played by a diverse set of anatomical and quantitative characters without discretization on phylogenetic reconstructions. We advance on the taxonomic knowledge of Eigenmanniinae by examining most of its species richness and building a hypothesis of phylogenetic relationships based on the parsimony analysis of 144 anatomical characters. As result, a new classification was proposed. This study highlights the crucial role that a diverse set of anatomical characters play at different hierarchical levels in phylogenetic reconstructions.
... Curimatidae harbor 115 fish species that are extremely abundant in freshwater environments from coastal rivers of Costa Rica to southern La Plata estuary in Argentina (Fricke, Eschmeyer, & Fong, 2019;Vari, 2003) and contains eight extant genera: Curimatopsis, Potamorhina, Curimata, Psectrogaster, Pseudocurimata, Curimatella, Cyphocharax, and Steindachnerina (Vari, 1989). Every genus was subject of comprehensive taxonomic revisions and phylogenies based on morphological data (Dillman, Sidlauskas, & Vari, 2016;Vari, 2003), and the relationships were recently tested by molecular data that revealed new phylogenetic arrangements (Melo, Sidlauskas, et al., 2018). In addition, a previous molecular study of Curimatopsis has challenged the current species diversity in the family by revealing several undescribed and cryptic species that was thought to contain only five species for more than 30 years (Melo, Ochoa, Vari, & Oliveira, 2016), five of them latter described (Dutra, Melo, & Netto-Ferreira, 2018;Melo & Oliveira, 2017;. ...
Potamorhina includes the largest species in the Neotropical fish family Curimatidae. They perform long‐distance migrations in large schools and represent relative importance for regional fisheries in South American lowlands. A morphology‐based phylogenetic study recognized five species and proposed interspecific phylogenetic relationships mostly based on osteology, squamation, and morphology of the gasbladder. Subsequent cytogenetic studies revealed extreme variability in diploid numbers and other cytomolecular structures and hypothesized multiple events of chromosome rearrangements with centric fissions followed by reversed fusions. However, neither the taxonomic revision and phylogeny nor the cytogenetic hypothesis of chromosome evolution in Potamorhina was tested using molecular phylogenetic approaches. Here, we use mitochondrial and nuclear DNA sequences to delimit species of Potamorhina with an extensive sampling across the Amazon basin and use phylogenetic methods to test prior hypothesis of multiple events of chromosome rearrangements during the evolution of the genus. Phylogenetic and species delimitation methods clearly support the presence of five species but reveal novel interspecific relationships allowing a reinterpretation of the morphological characters relative to the number of vertebrae, caudal peduncle pigmentation, and modifications in the gasbladder chambers. With the new phylogenetic arrangement, we propose a novel hypothesis of occurrence of a single chromosome fission in the lineage of P. latior followed by an extraordinary event that involved more than 20 chromosome‐pair fissions during the evolution of the ancestor of P. altamazonica and P. squamoralevis. This novel hypothesis represents a simpler and more conceivable explanation for the achievement of these elevated chromosome numbers during the evolution of Potamorhina. Previous studies hypothesized multiple events of chromosome rearrangements with centric fissions and reversed fusions during the evolution of Neotropical fishes of the genus Potamorhina. We use phylogenetic and delimitation methods, and results reveal novel relationships allowing a reinterpretation of the morphological characters. We also propose a hypothesis involving a single chromosome fission followed by a major event of >20 chromosome‐pair fissions, which represent a simpler and more conceivable explanation for these elevated chromosome numbers, very unusual among diploid teleosts.
... These were found in Citharinus latus Müller & Troschel, 1844 with a morphology remarkably similar to that present in Prochilodus. However, the Prochilodontidae, together with the Curimatidae, Anostomidae and Chilodontidae, constitute the Anostomoidea (Vari, 1983;Buckup, 1998;Dillman et al., 2016), while the African characiform families Citharinidae and Distichodontidae form the Citharinoidei, a monophyletic lineage that is traditionally considered the sister group to all remaining Characiformes (Vari, 1979;Ortí & Meyer, 1997;Buckup, 1998;Calcagnotto et al., 2005;Oliveira et al., 2011;Dillman et al., 2016). Monophyly of both Anostomoidea and Citharinoidei are strongly supported by both morphological and molecular data (Vari, 1979(Vari, , 1983Oliveira et al., 2011). ...
... These were found in Citharinus latus Müller & Troschel, 1844 with a morphology remarkably similar to that present in Prochilodus. However, the Prochilodontidae, together with the Curimatidae, Anostomidae and Chilodontidae, constitute the Anostomoidea (Vari, 1983;Buckup, 1998;Dillman et al., 2016), while the African characiform families Citharinidae and Distichodontidae form the Citharinoidei, a monophyletic lineage that is traditionally considered the sister group to all remaining Characiformes (Vari, 1979;Ortí & Meyer, 1997;Buckup, 1998;Calcagnotto et al., 2005;Oliveira et al., 2011;Dillman et al., 2016). Monophyly of both Anostomoidea and Citharinoidei are strongly supported by both morphological and molecular data (Vari, 1979(Vari, , 1983Oliveira et al., 2011). ...
... Phylogenetically, bony tubes rostral to the preopercle are found in both Prochilodontidae and Anostomidae. The monophyly of Prochilodontidae + Curimatidae, and of Anostomidae + Chilodontidae is strongly supported by morphological data (Vari, 1983;Dillman et al., 2016). Therefore, with a phylogenetic perspective, the distribution of autogenous ossification of the preopercular canal is most parsimoniously interpreted as being independently acquired by both families (Fig. 20). ...
The lateral-line system has been traditionally recognized as an important source of phylogenetic information for different groups of fishes. Although extensively studied in Siluriformes and Cypriniformes, the lateral-line system of Characiformes remained underexplored. In the present study, the anatomy of the cephalic lateral-line canals of characiforms is described in detail and a unifying terminology that considers the ontogeny and homologies of the components of this system is offered. Aspects of the arrangement of lateral-line canals, as well as the number, location and size of canal tubules and pores, resulted in the identification of novel putative synapomorphies for Characiformes and several of its subgroups. The study also revised synapomorphies previously proposed for different characiform families and provided comments on their observed distribution across the order based on extensive taxon sampling. Information from the ontogenetic studies of the cephalic lateral-line canal system and a proposal for the proper use of these data to detect truncations in the development of the lateral-line canals across the order is also offered.
... Curimatidae is a monophyletic family within Characiformes and is supported by synapomorphies related to the branchial apparatus, buccopharyngeal complex, hyoid arch, jaws, palatine arch, and neurocranium (Vari 1989, Dillman et al. 2016, which are associated with specialized food acquisition and processing (Netto-Ferreira and Vari 2011). Vari (1989) proposed the monophyly of Steindachnerina Fowler 1906 on the basis of 4 synapomorphies related to modifications in the first and second infrapharyngobranchials, the ventral process of the third hypobranchial (H3), the basihyal, and associated basihyal tooth-plate. ...
Recent expeditions to northwestern Argentina revealed the presence of an unknown species of Curimatidae for the Bermejo river basin. The morphometric and meristic analyses of these specimens allow us to identify them as Stein-dachnerina insculpta (Fernández-Yépez, 1948), which is here reported for the first time in Argentina.
... Through these triumphs and challenges, the field of Neotropical Ichthyology has advanced dramatically in the past 20 years. Morphological phylogenetic analyses have become larger and more comprehensive (Dillman et al., 2016;Mirande, 2018) or have delved into little explored character systems like those of the gas-bladder, muscles, pseudotympanum, sensory canals or ultrastructure of sperm (Di Dario, 2004;Birindelli et al., 2009;Datovo, Castro, 2012;Quagio-Grassiotto et al., 2012;Dutra et al. 2015). Molecular approaches have become widespread and even ubiquitous in their application with their phylogenetic results alternately challenging, expanding or affirming earlier osteological analyses Lujan et al., 2015;Tagliacollo et al., 2016;Carvalho et al., 2018;Melo et al., 2018) and providing rich data with which to examine hypotheses of speciation (Barluenga, Meyer 2004;Pereira et al., 2011;Ramirez et al., 2017). ...
... We have also advanced greatly in our knowledge of the phylogeny and evolution of Neotropical fishes, with recent molecular, morphological and combined studies frequently resolving stable reconstructions among families and genera within each order Birindelli, 2014;Lujan et al., 2015;Lopez-Fernandez et al., 2010;Dillman et al., 2016;Tagliacollo et al., 2016;Ilves et al., 2018;Mirande, 2018;Carvalho et al., 2018). Yet, not all studies agree, and new data have challenged existing classifications (Covain et al., 2016), refined longstanding phylogenetic hypotheses (Melo et al., 2018) or identified recalcitrant nodes that challenge even the most current analyses (Ilves et al., 2018). ...
Studies on the diversity, taxonomy, phylogeny, and biogeography of Neotropical Fishes have thrived over the twenty years that have elapsed since the first symposium on their phylogeny and classification in Porto Alegre, Brazil. Here, we review recent advances in the study of Neotropical fishes and assess the known diversity of freshwater species in that region. 6,255 valid freshwater species have been discovered in the Neotropics so far, and we estimate that over 9,000 species will be known when the inventory is complete. We also summarize the events of the second Symposium on Phylogeny and Classification of Neotropical Fishes that took place last year in Londrina, Brazil. Along with invited talks on the biodiversity of all major groups of Neotropical fishes, a series of presentations on the development of fish collections, and numerous contributed talks, the meeting included a special session to honor Dr. Richard Vari, who was one of the most prolific and beloved members of our community.
... Some species are found only in rivers of white, black, or clear water, while others are eurytopic and live in more than one type of water (Vari et al., 2010). A morphological phylogenetic analysis of the family was unable to recover synapomorphies supporting the monophyly of the genus (Vari, 1989(Vari, , 1992, and in fact, a recent supermatrix-based phylogeny for Anostomoidea based on morphological data failed to recover Cyphocharax as a monophyletic taxon (Dillman et al., 2015). Even after the extensive revision of the genus by Vari (1992), new collections in some regions of South America continue to reveal the existence of undescribed species (Melo and Vari, 2014;Melo, 2017). ...
A new species of Cyphocharax (Characiformes: Curimatidae) is described from the lower Rio Tapajós basin, including its tributary, the Rio Arapiuns, Pará, Brazil. The new species is diagnosed from congeners by a combination of characters that includes overall body light-colored, with a silvery hue, lacking any pattern of stripes or blotches, a distinctly straight body profile, and some meristic and morphometric features. Comparisons with similar congeners as Cyphocharax festivus, Cyphocharax leucostictus, Cyphocharax nigripinnis, and Cyphocharax plumbeus, some of which occur sympatrically with the new species, are presented. © 2018 by the American Society of Ichthyologists and Herpetologists.
... The most comprehensive phylogenetic study of the family to date (Vari, 1989a) hypothesized a monophyletic Curimatidae supported by 19 morphological synapomorphies. A recent reanalysis of a supermatrix linking data from that study to data for relationships in the superfamily Anostomoidea expanded that list to 27 synapomorphies (Dillman et al., 2016). The vast majority of these features involve modifications of the branchial apparatus, buccopharyngeal complex, hyoid arch, jaws, palatine arch, and neurocranium, many of which presumably adapt the family to its unusual detritivorous niche. ...
... That phylogeny resolved as a ladderized topology (Fig. 2a) with a terminal polytomy among Curimatella, Cyphocharax, Pseudocurimata and Steindachnerina. Dillman et al. (2016) recovered a similar result, corroborated the non-monophyly of Cyphocharax with relationships within that genus as an unresolved comb, and hinted at the possibility of a polyphyletic Curimatella in the Bayesian portion of their morphological analysis. ...
... To investigate the evolutionary implications of the molecular phylogeny reconstructed herein, we performed ancestral state reconstructions on nine morphological characters (numbers 28, 37, 193, 194, 241, 329, 346, 406, and 459) extracted from the supermatrix of Dillman et al. (2016). These nine characters represent putative synapomorphies of clades not recovered in the molecular analysis, and thus characters whose interpretation may change. ...
