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Neotropical Ichthyology

Print version ISSN 1679-6225On-line version ISSN 1982-0224

Neotrop. ichthyol. vol.17 no.1 Maringá  2019  Epub Apr 25, 2019

http://dx.doi.org/10.1590/1982-0224-20180076 

Original article

Revalidation and redescription of Steindachnerina nigrotaenia and redescription of S. insculpta (Characiformes: Curimatidae)

Heraldo A. Britski1 
http://orcid.org/0000-0002-5593-9651

Bruno F. Melo2 
http://orcid.org/0000-0002-0499-567X

Richard P. Vari3 

Claudio Oliveira2 
http://orcid.org/0000-0002-4143-7212

1Museu de Zoologia, Universidade de São Paulo, Av. Nazaré, 481, 04264-000 São Paulo, SP, Brazil. heraldo@usp.br, https://orcid.org/0000-0002-5593-9651

2Departamento de Morfologia, Instituto de Biociências, Universidade Estadual Paulista, IBB/UNESP, R. Prof. Dr. Antonio C. W. Zanin, 250, 18618-689 Botucatu, SP, Brazil. (BFM) melo@ibb.unesp.br, https://orcid.org/0000-0002-0499-567X (corresponding author); (CO) claudio@ibb.unesp.br, https://orcid.org/0000-0002-4143-7212

3Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC, USA. In memoriam.

ABSTRACT

Steindachnerina nigrotaenia is resurrected from the synonym of S. brevipinna and considered a valid species. The previous designation of the lectotype of S. nigrotaenia is considered invalid and a new lectotype is designated herein. Steindachnerina nigrotaenia and S. insculpta are redescribed based on type specimens and on additional material from the rio Paraguai and the upper rio Paraná basins, respectively. The two species can be separated by the number of scales of the lateral line and of the transverse series and by phylogenetic analyses of molecular data.

Keywords: Ostariophysi; Neotropical fishes; Systematics; Taxonomy

RESUMO

Steindachnerina nigrotaenia é retirada da sinonímia de S. brevipinna e considerada uma espécie válida. A designação do lectótipo de S. nigrotaenia é considerada inválida e um novo lectótipo é aqui designado. Steindachnerina nigrotaenia e S. insculpta são redescritas com base no material tipo e em exemplares adicionais das bacias do rio Paraguai e alto rio Paraná, respectivamente. As duas espécies se diferenciam pelo número de escamas da linha lateral e da série transversal e por análises filogenéticas de dados moleculares.

Palavras-chave: Ostariophysi; Peixes neotropicais; Sistemática; Taxonomia

Introduction

The characiform family Curimatidae includes eight genera, two of which concentrate more than half of the species in the family, Cyphocharax Fowler, 1906, and Steindachnerina Fowler, 1906. Of the 24 valid species of Steindachnerina, four species occur in southern South America in the La Plata system: S. biornata (Braga & Azpelicueta, 1987), S. brevipinna (Eigenmann & Eigenmann, 1889), S. conspersa (Holmberg, 1891), and S. insculpta (Fernández-Yépez, 1948). Both S. biornata and S. conspersa appear at the base of the phylogeny of the genus with significant morphological differences when compared to the other two species, whereas S. brevipinna and S. insculpta has similar morphologies and are close related species (Vari, 1991; Melo et al., 2018). Vari (1991) provided a valuable progress in the systematics of the genus by analyzing many type specimens that enable him to complete the taxonomic revision and phylogeny of Steindachnerina. However, there is a putative conflict in the number of valid species in the rio Paraguai (two, S. conspersa and S. brevipinna) and the number of observed morphotypes in that region (three). The available name for the third species is Steindachnerina nigrotaenia (Boulenger, 1902), currently placed in synonym of S. brevipinna by Vari (1991). The history behind the types of Curimatus nigrotaenia explains the reasons of the mistaken synonymy.

Early in the 20th century, the Director of the Museo Civico di Storia Naturale of Genova (MSNG) entrusted to Dr. George A. Boulenger of the British Museum (Natural History) (BMNH) a collection of fishes and reptiles collected by Dr. F. Silvestri in 1900 in Brazil, Paraguay and Argentina. One among several fish species described by Boulenger (1902) based on that material was Curimatus nigrotaenia Boulenger, 1902, a curimatid from the rio Coxipó, state of Mato Grosso, Brazil, upper Paraguai basin. The species was mainly characterized by the possession of 43 or 44 scales in the lateral line, 7.5 to 8.5 series of scales between the lateral line and dorsal-fin origin and six series of scales between the lateral line and pelvic-fin origin, supplemented by a dark, longitudinal stripe along midlateral surface of the body extending onto middle rays of the caudal fin. Boulenger did not mention any dark spot on the basal portion of the middle dorsal-fin rays, a distinct feature present in many species of curimatid fishes. These characters accurately discriminated C. nigrotaenia from other curimatids known at that time from the rio Paraguai basin. Silvestri’s collection was partially returned to the MSNG, but, as reported by Boulenger (1902: 284), some of the specimens were retained at the BMNH. However, Boulenger did not indicate how many individuals of each species were retained at the BMNH versus returned to the MSNG.

Vari (1991) redefined Steindachnerina, to incorporate various species previously assigned to Curimatus Oken, 1817 (= Curimata Bosc, 1817), including Curimatus nigrotaenia. After examining five specimens at the BMNH labeled as syntypes of C. nigrotaenia under catalog number BMNH 1902.2.10:30, Vari (1991:100) observed that the features of these specimens corresponded to those of Steindachnerina brevipinna described from Río de la Plata, Rosario, Argentina. The putative type specimens of C. nigrotaenia had 34 to 35 scales in the lateral line series from the supracleithrum to the hypural joint, 5.5 scales between the lateral line and dorsal-fin origin, 4.5 scales between the lateral line and anal-fin origin, a prominent midlateral dark stripe and a dark spot on the basal portion of the middle rays of the dorsal fin. As a result, Vari (1991:100) treated C. nigrotaenia as a junior synonym of S. brevipinna, designated the largest specimen as the lectotype of C. nigrotaenia, and transferred the other four putative syntypes to BMNH 1989.2.2:3-6 as paralectotypes.

However, the data presented by Boulenger (1902) in the original description disagreed substantially from the data collected by Vari (1991) when he examined the specimens. At that time, Vari (1991:100) considered it impossible to determine whether the original data published by Boulenger contained typographical errors or Boulenger’s counts were incorrect. In addition, he assumed that Boulenger “failed to describe the prominent dark spot on the basal portion of the middle rays of the dorsal fin in the relatively small-sized syntypes” he examined. However, he did not discuss the possibility of a curatorial error at BMNH, resulting in the wrong specimens being placed in the syntype jar for Curimatus nigrotaenia.

After the revision of Vari (1991), one of us (HAB) suspected the occurrence of an error due to several morphological differences observed in the two morphotypes collected in the rio Paraguai. Despite Vari’s conclusions, Britski (1996) maintained Steindachnerina nigrotaenia as a valid species in a brief communication in a regional meeting for biologists, and in the two volumes of the manual of fishes from the Pantanal Matogrossense (Britski et al., 1999; 2007), suspecting that the specimens examined by Vari (1991) at the BMNH were not the syntypes of the species, but actually specimens of S. brevipinna incorrectly labeled as syntypes of S. nigrotaenia. Additionally, the ICZN (Case 3307) received in 2004 a proposal of conservation of usage of the name Curimatus nigrotaenia (= S. nigrotaenia) by designation of a new lectotype, proposed by Britski and Vari, without any resolution.

In this context, we present complete redescriptions of the two species taking into account that specimens of Steindachnerina nigrotaenia and S. insculpta were mixed in the redescription under the name of S. insculpta by Vari (1991), and that the description of Curimatus nigrotaenia by Boulenger (1902), the redescriptions by Britski et al. (1999; 2007), as well as the original description of Cruxentina insculpta by Fernández-Yépez (1948) are very abridged and/or incomplete. In summary, the purpose of this paper is to (1) revalidate Steindachnerina nigrotaenia (Boulenger, 1902), (2) designate a new lectotype for the species, and (3) redescribe both S. nigrotaenia and S. insculpta based on new morphological and molecular data from both species.

Material and Methods

Morphology. To resolve the curatorial issues involving Steindachnerina nigrotaenia, we examined the type specimens deposited at the Natural History Museum of London (BMNH) and at the Museo di Storia Naturale of Genova (MSNG), and additional material from recent field collections of S. brevipinna, S. insculpta and S. nigrotaenia. Measurements were made with digital calipers to the nearest 0.1 mm. Counts and measurements were taken on the left side of specimens whenever possible. Lateral line scales were counted from the supracleithrum to caudal fin, including those overlying the hypural plate and extending to the base of median caudal-fin rays, which we also report separately. Counts of total vertebrae were usually taken from radiographs, the fused PU1+U1 was considered a single bone, and the vertebrae incorporated into the Weberian apparatus were counted as four elements.

Subunits of the head are presented as proportions of head length (HL). Head length itself and measurements of body parts are given as proportions of standard length (SL). Greatest body depth was taken at the dorsal-fin origin. In the counts of median-pectoral and pelvic fins, the unbranched rays are indicated by lower case roman numerals, and the branched-fin rays as Arabic numerals. Meristic data are given in the description with parentheses indicating the frequency of each count and asterisks indicating counts of holotype or lectotype specimens.

In the synonymies for each species, localities are presented as in the original citation, followed by the corrected or modern equivalent (in parentheses) if the case. Institutional abbreviations are: Natural History Museum, London (BMNH); California Academy of Sciences, San Francisco (CAS); Laboratório de Biologia e Genética de Peixes, Universidade Estadual Paulista, Botucatu (LBP); Museu Nacional do Rio de Janeiro, Rio de Janeiro (MNRJ); Museo Civico di Storia Naturale de Genova, Genova (MSNG); Museu de Zoologia da Universidade de São Paulo, São Paulo (MZUSP); Núcleo de Pesquisas em Limnologia, Ictiologia e Aquicultura, Universidade Estadual de Maringá, Maringá (NUP); Naturhistorisches Museum Wien, Vienna (NMW).

Molecular analysis. Nineteen specimens of Steindachnerina were included in the molecular analysis: five specimens of S. brevipinna from rio Paraná, five specimens of S. insculpta from the main rio Paraná above the Itaipu dam and rio Tietê, eight specimens of S. nigrotaenia from the upper rio Paraguai, and one specimen of S. elegans from the rio São Francisco to root the tree. Sequences of ten out of nineteen specimens were obtained from Genbank (Tab. 1).

Tab. 1 Taxon, voucher, locality information and Genbank accession numbers of specimens of Steindachnerina analyzed in the molecular approach. 

Taxon Voucher Specimen Locality City, State Country Genbank
Steindachnerina brevipinna LBP 6690 32153 Rio Paraná Marilena, Paraná Brazil GU701503
Steindachnerina brevipinna LBP 6690 32154 Rio Paraná Marilena, Paraná Brazil GU701502
Steindachnerina brevipinna LBP 6690 32155 Rio Paraná Marilena, Paraná Brazil GU701501
Steindachnerina brevipinna LBP 6690 32156 Rio Paraná Marilena, Paraná Brazil GU701505
Steindachnerina brevipinna LBP 6690 32157 Rio Paraná Marilena, Paraná Brazil GU701504
Steindachnerina insculpta LBP 3192 19389 Rio Tietê, Rio Paraná Conchas, São Paulo Brazil JN989237
Steindachnerina insculpta LBP 3192 19390 Rio Tietê, Rio Paraná Conchas, São Paulo Brazil JN989238
Steindachnerina insculpta LBP 5207 26343 Rio Paraná Porto Rico, Paraná Brazil JN989236
Steindachnerina insculpta LBP 6671 32081 Rio Paraná Marilena, Paraná Brazil JN989239
Steindachnerina insculpta LBP 19770 32158 Rio Paraná Marilena, Paraná Brazil GU701506
Steindachnerina nigrotaenia LBP 5636 27395 Rio Paraguai Cuiabá, Mato Grosso Brazil MK495705
Steindachnerina nigrotaenia LBP 8573 43355 Rio Paraguai Barra do Bugres, Mato Grosso Brazil MK495709
Steindachnerina nigrotaenia LBP 9818 45230 Rio Paraguai Miranda, Mato Grosso do Sul Brazil MK495701
Steindachnerina nigrotaenia LBP 9818 45231 Rio Paraguai Miranda, Mato Grosso do Sul Brazil MK495706
Steindachnerina nigrotaenia LBP 9818 45232 Rio Paraguai Miranda, Mato Grosso do Sul Brazil MK495702
Steindachnerina nigrotaenia LBP 9818 45233 Rio Paraguai Miranda, Mato Grosso do Sul Brazil MK495703
Steindachnerina nigrotaenia LBP 9818 45234 Rio Paraguai Miranda, Mato Grosso do Sul Brazil MK495707
Steindachnerina nigrotaenia LBP 9862 45408 Rio Paraguai Miranda, Mato Grosso do Sul Brazil MK495704
Steindachnerina elegans LBP 8272 38329 Rio São Francisco Jaíba, Minas Gerais Brazil MK495708

For the three newly sequenced individuals, genomic DNA was extracted from muscle tissues preserved in 95% ethanol with a DNeasy Tissue kit (Qiagen Inc.; http://www.qiagen.com) according to the manufacturer’s instructions. Partial sequences of the mitochondrial gene cytochrome oxidase c subunit I were obtained by polymerase chain reaction (PCR) using the primers described by Melo et al. (2011). Total volume included 12.5 μl with 9.075 μl of double-distilled water, 1.25 μl 5x buffer, 0.375 μl MgCl2 (50mM), 0.25 μl dNTP mix, 0.25 μl of each primer at 10 μM, 0.05 μl Platinum Taq DNA polymerase enzyme (5 units/μl, Invitrogen; www.invitrogen.com) and 1.0 μl genomic DNA (10-50 ng). The PCR program consisted of an initial denaturation (4 min at 95º C) followed by 30 cycles of chain denaturation (30 s at 95ºC), primer hybridization (30-60 s at 52ºC), and nucleotide extension (45 s at 72ºC). Amplicons were visualized through 1% agarose gel, and used to perform the sequencing reaction using dye terminators (BigDye™ Terminator v 3.1 Cycle Sequencing Ready Reaction Kit, Applied Biosystems), and purified through ethanol precipitation. Samples were then loaded onto an automatic sequencer ABI 3130-Genetic Analyzer (Applied Biosystems) at the São Paulo State University, Botucatu, Brazil.

Consensus sequences were assembled and edited in Geneious v7.1.9 (Kearse et al., 2012) and aligned with Muscle (Edgar, 2004) under default parameters. The index of substitution saturation in asymmetrical (Iss.cAsym) and symmetrical (Iss.cSym) topologies were estimated using Dambe v5.3.38 (Xia, 2013) and the best-fit model of nucleotide evolution was obtained in Mega v7.0 (Tamura et al., 2013). We ran a neighbor-joining tree (NJ) using the Tamura-Nei distance model and generated 1,000 non-parametric bootstrap replicates in Geneious (Kearse et al., 2012), and a maximum likelihood (ML) tree with 1,000 non-parametric bootstraps using GTRGAMMA, the single model implemented in RAxML HPC2 on XSEDE v8.2.10 (Stamatakis, 2014) on the Cipres webserver (Miller et al., 2010).

Results

The analyses of morphological and molecular data strongly support the recognition of three species of the same clade in the La Plata basin: Steindachnerina brevipinna (rio Paraná and rio Paraguai), S. insculpta (upper rio Paraná) and S. nigrotaenia (upper rio Paraguai), supplemented by S. conspersa and S. biornata. The synonymization of S. nigrotaenia was caused by previous curatorial mistakes that guided Vari (1991) to recognize only one species.

After finding the syntype of Curimatus nigrotaenia retained at the BMNH by Boulenger (1902) who assigned it the term “type”, the following notes in literis were received from James Maclaine, curator of the British Museum of Natural History, explaining the erroneous changes that took place with materials in that fish collection: “A) BMNH 1902.2.10.30 Curimatus nigrotaenia, SYNTYPE (1). - mistakenly placed into jar for BMNH 1917.11.2.7 Curimatus morawhannae SYNTYPE (1); B) BMNH 1911.10.31.479-483 Curimatus morawhannae SYNTYPES (5) - mistakenly placed into jar for BMNH 1902.2.10.30 Curimatus nigrotaenia, SYNTYPE (1); subsequently examined by Richard Vari and synonymized and separated into a lectotype (BMNH 1902.2.10.30) and paralectotypes (given new number BMNH 1989.2.23.3-6); the above two lots can be seen in the annotations in Günther’s Catalogue of Fishes; C) BMNH 1917.11.2.7 Curimatus morawhannae SYNTYPE (1) - mistakenly placed into jar for BMNH 1911.10.31.479-482 Curimatus morawhannae SYNTYPES.” This explanation provided by James Maclaine also allowed the correction of some BMNH catalogue numbers published by Vari (1991:87) about syntypes of Curimatus morawhannae.

Analysis of the three syntypes of Curimatus nigrotaenia deposited at the MSNG with the BMNH syntypes has confirmed that the material now in London was incorrectly curated. The still well-preserved syntypes at MSNG entirely fit Boulenger’s description and differ from the BMNH “syntypes” studied by Vari (1991). Furthermore, we have successfully found the “type” of C. nigrotaenia, which was wrongly deposited in a jar labeled Curimatus morawhannaeEigenmann, 1912, under the number BMNH 1917.11.2:7 (see paragraph above). Data taken from this “type” agree entirely with the original description of C. nigrotaenia. The specimen of C. nigrotaenia registered under the number BMNH 1902.2.10.30 by Boulenger also had already been cataloged in Günther’s Catalogue of Fishes (museum catalog, vol. 5, pp. 288a) of the BMNH. That finding also revealed that Boulenger retained only one specimen of C. nigrotaenia (labeled as “type”) at the BMNH, and not five syntypes as mentioned by Vari (1991). The number of syntypes sent back to MSNG was not specified by Boulenger, but by Tortonese (1961: 184), in his catalogue of types deposited at the MSNG mentioning three syntype specimens. Overall, the clearly morphological differences between Steindachnerina brevipinna and the syntypes of C. nigrotaenia at MSNG and BMNH illustrate that the synonymization was incorrect, and support the recognition of S. nigrotaenia as a valid species.

Vari’s (1991:100) designation of the lectotype of Curimatus nigrotaenia should be considered invalid because the lot from which he separated the specimen was incorrectly labeled as the syntype series of C. nigrotaenia. As specified in the ICZN (1999: Art. 74.2), “If it is demonstrated that a specimen designated as a lectotype was not a syntype, it loses its status of lectotype”. Günther’s citation of BMNH 1902.2.10.30 as a “type” for C. nigrotaenia is similarly invalid because “the mere citation of ‘Type’ or equivalent expression, … or in an unpublished catalogue of a museum, or on a label, is not necessarily evidence that a specimen is or is fixed as any of the kinds of types referred in this Chapter.” (ICZN, 1999: Art. 72.4.7). Consequently, we herein formally designate the specimen of C. nigrotaenia retained by Boulenger (1902) at the British Museum (BMNH 1902.2.10.30) as the new lectotype for Steindachnerina nigrotaenia, with the three syntypes at the Museo Civico di Storia Naturale of Genova (MSNG 14859) becoming paralectotypes. This is likely the same specimen considered by Günther when assembling his catalog of types, but definitely not the same specimen that Vari previously assigned to that catalog number.

How the “type” of Curimatus nigrotaenia was deposited in a jar labeled as C. morawhannae in the collection at the BMNH is an issue that possibly we will never know, but surely we cannot attribute this action to Boulenger when describing the species, because C. morawhannae was described in 1912 by Carl H. Eigenmann based on specimens from Guyana and, obviously, paratypes were sent to the British Museum after the description of the species (1912). So, this error should be credited to someone who worked with the type specimens of C. nigrotaenia and C. morawhannae afterwards, but before Vari’s examination.

Molecular analysis. Molecular results also support the recognition of Steindachnerina nigrotaenia. The matrix contained 675 bp with 48 polymorphic sites (7.1%), and 98.6% pairwise identity. Nucleotide frequencies corresponded to 22.9% adenine, 28.4% cytosine, 18.2% guanine, and 30.6% thymine/uracil. We did not find any saturation in the dataset. Tab. 1 includes voucher information, locality, and Genbank accession numbers. Both neighbor-joining analysis (not shown) and the maximum likelihood approach returned the same topology (Fig. 1). Results revealed a clear separation among samples of S. insculpta from the rio Paraná and the group S. brevipinna/S. nigrotaenia from rio Paraná and rio Paraguai, respectively. The eight specimens of S. nigrotaenia from the upper rio Paraguai (including one topotype from rio Coxipó) appear genetically closer to the five specimens of S. brevipinna from the rio Paraná. These results demonstrate that S. nigrotaenia clearly differs genetically from the morphologically similar S. insculpta, but much less from S. brevipinna which also occurs in the rio Paraguai. It appears that both S. brevipinna and S. nigrotaenia did accumulate morphological differences but not in the mitochondrial DNA. The sum of evidence allows us to redescribe S. nigrotaenia from the rio Paraguai basin, and S. insculpta from the upper rio Paraná basin.

Fig. 1 Best maximum likelihood tree based on partial sequences of the mitochondrial gene cytochrome oxidase c subunit I among samples of Steindachnerina nigrotaenia, S. insculpta and S. brevipinna. Numbers near nodes represent bootstrap values of maximum likelihood/neighbor-joining analyses. 

Steindachnerina nigrotaenia (Boulenger, 1902)

Fig. 2,Tab. 2

Curimatus nigrotaeniaBoulenger, 1902:2 [type-locality: Mato Grosso, rio Coxipó].-Eigenmann, 1910:421 [reference].-Vari, 1989, tables 2, 3 [assignment to Steindachnerina].-Tortonese, 1961:184 [Catalog: three syntypes].-Litz & Koerber, 2014:9 [Check list; synonym of Steindachnerina brevipinna].

Curimata nigrotaenia.-Fernández-Yépez, 1948:73 [reference].-Fowler, 1950:288 [literature compilation].-Fowler, 1975:369 [reference].-Ringuelet, 1975:72 [Argentina, rio Paraguai system].

Steindachnerina nigrotaenia.-Britski, 1996:74 (previous resurrection).-Britski et al., 1999:72 [diagnose in key; shortened description; text-figure; Pantanal do Mato Grosso].-Britski et al., 2007: 96 [diagnose in key; shortened description; text figure; Pantanal do Mato Grosso].-Polaz et al., 2014:125 [list of species from Parque Nacional do Pantanal Matogrossense].

Steindachnerina insculpta.-Vari, 1991:71 [in part, material from rio Paraguai basin, not from upper rio Paraná basin]. -Aguilera et al., 2018:1018 [Argentina: río San Francisco, Bermejo river basin, Jujuy, and Madrejón El Divisadero, Bermejo river basin, Salta].

Steindachnerina brevipinna.-Vari, 1991:97 [in part, material from Mato Grosso, upper rio Paraguai].

?Curimata nasa.-Géry et al., 1987:425, fig. 41 [material from arroyo Trementina, rio Paraguai basin].

Diagnosis. Steindachnerina nigrotaenia can be distinguished from all congeners, except S. binotata (Pearson, 1924), S. biornata, S. dobula (Günther, 1868), S. hypostoma (Boulenger, 1887), S. insculpta, S. leucisca (Günther, 1868), S. notograptos Lucinda & Vari, 2009, and S. varii Géry, Planquette, Le Bail, 1991, by possessing an unpigmented dorsal fin (vs. a dorsal fin with a spot of dark pigmentation on the basal portion of the middle rays). Steindachnerina nigrotaenia has 43 to 48 perforated scales in the lateral line series running from the supracleithrum to the middle rays of the caudal fin, differing from S. binotata, S. leucisca and S. notograptos, which have 50 or more perforated scales in that series and from S. biornata and S. varii which have fewer than 40 perforated scales in the lateral line series. Steindachnerina nigrotaenia differs from S. hypostoma by possessing a relatively shallower body with greatest body depth 24-27% of SL (vs. 29-34% of SL). Steindachnerina nigrotaenia differs from S. dobula and S. varii by having a dark midlateral stripe extending from the rear of the opercle posteriorly onto middle rays of caudal fin (vs. dark midlateral stripe beginning posterior to a vertical line through the dorsal-fin origin, and continuing posteriorly to the middle rays of the caudal fin). Steindachnerina nigrotaenia differs from the very similar S. insculpta by the generally higher number of perforated scales along the lateral line series (43 to 48, 45 most frequent, vs. 40 to 45, 42 most frequent and 44 and 45 in only 1.3% of specimens examined for this feature, respectively; Fig. 3), by the total number of scale rows in the transverse series from the dorsal-fin origin to the pelvic-fin base (13 to 16, 14 most frequent, vs. 12 to 14, 12 most frequent and 13.5 and 14 in only 6.4% and 1.3% of specimens examined for this feature, respectively; Fig. 4). Steindachnerina nigrotaenia can also be discriminated from congeners by a combination of the following features: multiple lobulated fleshy processes on the roof of the oral cavity, absence of a wide, flattened prepelvic region of the body, possession of 43 to 48 perforated scales in the lateral line series, 13 to 16 scales in the transversal series, lack of a dark spot on the basal portions of the dorsal fin and the presence of a dark midlateral stripe along the lateral surface of the body.

Fig. 2 a. Steindachnerina nigrotaenia, lectotype of Curimatus nigrotaenia, BMNH 1902.2.10.30, 43.5 mm SL, Brazil, Mato Grosso, rio Coxipó, upper rio Paraguai; b. LBP 1477, 43.4 mm SL, Brazil, Mato Grosso do Sul, Coxim, rio Taquari, upper rio Paraguai. 

Fig. 3 Histogram of number of lateral-line scales against number of specimens of Steindachnerina brevipinna, S. insculpta and S. nigrotaenia

Fig. 4 Histogram of number of series of scales from dorsal-fin origin to pelvic-fin base against number of specimens of Steindachnerina brevipinna, S. insculpta and S. nigrotaenia

Description. Morphometric data of lectotype, paralectotypes and other specimens are summarized in Tab. 2. Body relatively elongate, somewhat compressed. Dorsal profile of head slightly convex anteriorly, straight from the line above orbit to rear of head. Dorsal profile of body straight or very slightly convex from rear of head to origin of dorsal fin; straight at dorsal-fin base; straight from base of last dorsal-fin ray to caudal peduncle. Dorsal surface of body transversely rounded anteriorly, with indistinct median keel immediately anterior to dorsal fin, smoothly rounded transversely posterior to dorsal fin. Ventral profile of body gently curved from tip of lower jaw to caudal peduncle. Prepelvic region obtusely flattened, with median series of scales proximate to pelvic-fin origin, median series irregularly arranged anteriorly. Barely discernable median keel posterior to pelvic-fin origin.

Tab. 2 Morphometric data for Steindachnerina nigrotaenia and S. insculpta. N = number of specimens. SD = Standard deviation. Range of each species includes the lectotype of S. nigrotaenia (BMNH 1902.2.10.30) and the paratype of S. insculpta (MZUSP 1376), respectively. 

Steindachnerina nigrotaenia Steindachnerina insculpta
Lectotype N Range Mean SD Paratype N Range Mean SD
Standard length (mm) 43.5 44 37.0-90.3 63.4 - 105.0 78 42.6-108.1 75.6 -
Percentages of standard length
Body depth 26.5 44 25.0-33.8 28.8 1.5 29.5 78 25.6-34.1 30.1 1.7
Caudal peduncle depth 11.0 44 10.3-11.7 11.0 0.3 11.7 78 10.8-12.4 11.7 0.4
Head length 32.0 44 25.7-32.5 29.0 1.7 18.9 78 25.6-30.2 28.1 0.8
Predorsal distance 47.2 44 44.3-49.7 46.8 1.2 46.8 78 43.3-49.7 46.2 1.2
Prepelvic distance 51.9 44 48.8-55.4 52.4 1.4 54.9 78 48.8-58.0 52.1 1.6
Pectoral-fin length 15.2 44 15.0-20.0 18.1 0.9 17.9 78 15.6-19.8 18.0 0.9
Pelvic-fin length 19.9 44 18.0-22.3 19.7 0.9 19.6 78 15.5-21.8 19.0 1.2
Percentages of head length
Eye diameter 32.4 44 27.6-34.9 30.5 1.8 27.7 78 26.3-33.5 29.8 1.7
Snout length 34.5 44 30.1-35-6 32.9 1.3 35.1 78 29.7-38.9 34.1 1.7
Interorbital distance 33.1 44 31.7-43.2 38.0 2.2 39.7 78 38.3-44.8 41.0 1.8

Dorsal-fin margin rounded; anteriormost rays three to 3.5 times the length of ultimate ray. Pectoral-fin margin acute, extending one-half to two-third the distance to origin of pelvic fin in smaller adults, barely beyond that point in largest specimens examined. Pelvic-fin margin acute; pelvic fin extending about one half the distance to origin of anal fin. Caudal fin forked. Adipose fin well developed. Anal fin emarginate, anteriormost branched rays 2.5 to three times the length of ultimate ray.

Head profile pointed; upper jaw distinctly longer, mouth inferior; buccopharyngeal complex on roof of the oral cavity in adults consisting of multiple lobulated fleshy bodies; nostrils very close, anterior circular, posterior crescent-shaped with aperture closed by thin flap of skin separating nares; adipose eyelid present, more developed anteriorly, with broad, vertically ovoid opening over center of eye.

Pored lateral-line scales from supracleithrum to medial caudal-fin rays 43(1), 44(9), 45*(22), 46(10), 47(1) or 48(1), 2 to 3 over caudal-fin base; series of scales from dorsal-fin origin to lateral line 6(8), 7*(31) or 8(5); series of scales from lateral line to pelvic-fin base 6*(39), 6.5(3) or 7(2); total series of scales from dorsal fin to base pelvic-fin base 13(4), 14*(36), 14.5(5), 15(3) or 16(1); series of scales from lateral line to first ray of anal fin 5*(6), 5.5(8) or 6(29); circumpeduncular scales 16*(36), 17(1), 18(3) or 20(3); predorsal scales 11*(13), 12(19) or 13(10); scales from dorsal fin to adipose fin 12*(7), 13(17), 14(16) or 15(3); scales from adipose fin to caudal fin 8(3), 9(10), 10(19), 11(8) or 12*(2); prepelvic scales 18(1), 19(8), 20*(10), 21(12), 22(5), 23(4) or 24(1); scales from pelvic fin to anus 8*(13), 9(21), 10(7) or 11(2); scales from anus to anterior margin of anal fin 2(4), 3(31) or 4*(9); scales from posterior margin of anal fin to anteroventral margin of caudal fin 8(1), 9(6), 10(16), 11*(16) or 12(2). All scales of lateral line pored with canals in scales straight.

Dorsal-fin rays ii,8,ii*; pectoral-fin rays i,12(3), i,13(7), i,14*(20), i,15(9) or i,16(4); pelvic-fin rays i,8*(41) or i,9(3); anal-fin rays ii-iii,5,ii(2) or ii-iii,6,ii*(42); caudal-fin rays i,8,9,i*(38). Total vertebrae 28(8) or 29 (2).

Coloration in alcohol. Overall coloration of specimens retaining guanine on scales silvery to silvery golden, darker on dorsal portions of head and body. Ground coloration of specimens lacking guanine on scales tan to yellow, darker dorsally, with irregular patch of dark pigmentation extending from rear of orbit across opercle; degree of intensity of dark pigmentation and extent of patch variable among individuals. Irregular, dark longitudinal stripe extending along lateral line from supracleithrum to base of middle caudal-fin rays. Stripe slightly wider posteriorly, continuous posteriorly with dusky stripe on middle caudal-fin rays. Stripe in adults continuous, about one scale wide; not as well developed in juveniles, consisting of discrete spots surrounding pores of lateral line. Caudal-fin with stripe more prominent on the anterior portion of rays. Anterior margin and distal portions of dorsal fin typically dusky in adults. Ventral lobe and dorsal rays of dorsal lobe of caudal fin dusky. All caudal-fin rays outlined by small chromatophores on membranes. Adipose fin dusky distally. Dorsal fin with dusky anterior region in many specimens, without any discrete spot on middle rays. Other fins hyaline.

Sexual dimorphism. Secondary sexual characters were not found.

Geographical distribution. Steindachnerina nigrotaenia is distributed along the rio Paraguai basin (Fig. 5).

Fig. 5 Distribution map of Steindachnerina brevipinna in both rio Paraná and rio Paraguai (blue), S. nigrotaenia in the rio Paraguai (red), and S. insculpta in the upper rio Paraná (yellow). Pentagons indicate type localities. Some symbols represent more than one collecting locality. 

Material examined. All from Brazil, rio Paraguai basin. BMNH 1902.2.10.30, 43.5 mm SL, Mato Grosso, rio Coxipó, F. Silvestri. Lectotype of Curimatus nigrotaenia (designated herein). MSNG 14859, 3, 42.7, 51.2, 52.2 mm SL, same data as lectotype. Paralectotypes of Curimatus nigrotaenia (designated herein). MNRJ 11206, 3, 59.8-85.7 mm SL, Mato Grosso, Acorizal, rio Acorizal at bridge along road from Acorizal to Baús. MNRJ 11207, 1, 102.0 mm SL, Mato Grosso, Acorizal, ribeirão Baús. MNRJ 11214, 1, not measured, Mato Grosso, Acorizal, ribeirão Tangará, along road from Acorizal to Cuiabá. MZUSP 4383, 3; MZUSP 4449, 1, 39.2 mm SL, Mato Grosso, Santo Antonio do Leverger, rio Cuiabá. MZUSP 21512, 16, Mato Grosso, Cuiabá, rio Coxipó da Ponte, 15°38’S 56°03’W. MZUSP 21656, 6, Mato Grosso, Santo Antonio do Leverger, rio Cuiabá. MZUSP 21662, 2, Mato Grosso, Barão do Melgaço, rio Cuiabá, Boca do Croará, 16°11’S 55°57’W. MZUSP 38015, 2, Mato Grosso, Cáceres, Porto Limão. MZUSP 38079, 1, Mato Grosso do Sul, Coxim, rio Taquari, Ilha da Goiaba, 18°30’00”S 54°45’00”W. MZUSP 40068, 4, Mato Grosso do Sul, Aquidauana, rio Aquidauana, Baia da Onça or Jatobá, Fazenda Alegrete, 20°28’00”S 55°48’00”W. MZUSP 45476, 9, Mato Grosso, rio Paraguai at Cáceres and vicinities. MZUSP 47762, 130, Mato Grosso, Barão de Melgaço, rio Cuiabá. MZUSP 54139, 1, rio Apa, small bay of the river. MZUSP 56234, 1, Mato Grosso, Chapada dos Guimarães, rio Casca. MZUSP 59541, 41, Mato Grosso do Sul, Corumbá, rio Vermelho, Fazenda Xaroes, 19°36’73”S 56°55’42”W. MZUSP 74361, 4, Mato Grosso, Santo Antonio do Leverger, rio Cuiabá, 3 km from the town. MZUSP 75480, 2, Mato Grosso, Cáceres, ribeirão das Flexas, along road from Cáceres to Cuiabá at about 69 km E of Cáceres. MZUSP 78750, 3, Mato Grosso, Porto Estrela, rio Saloba, along road at Estação Ecológica da Serra das Araras, 15°39’03”S 57°13’29”W. MZUSP 78845, 1, Mato Grosso, Diamantino, downstream of waterfall in the rio Paraguai, 14°28’32”S 56°23’32”W. MZUSP 78946, 1, Mato Grosso, Santo Antonio do Leverger, rio Cuiabá. MZUSP 87785, 2, Mato Grosso, Tangará da Serra, rio Sepotuba, salto Maciel, 15°55’59”S 57°39’00” W. MZUSP 89954, 2, Mato Grosso, rio Sepotuba. MZUSP 90075, 8, Mato Grosso, Curvelândia, lower rio Sepotuba, 15°53’34”S 57°38’44”W. MZUSP 90181, 14, Mato Grosso, lower rio Sepotuba. MZUSP 90422, 27, Mato Grosso, Curvelândia, rio Sepotuba, 15°46’07”S 57°38’54”W. MZUSP 90702, 3, Lambari d’Oeste, rio Sepotuba, at mouth of a creek, 15°10’00”S 57°41’00”W. MZUSP 91014, 1, Barra do Bugres, rio Sepotuba, 15°06’52”S 57°39’57”W. MZUSP 91060, 2, Cáceres, rio Sepotuba, artificial lagoon downstream of the bridge, 15°14’08”S 57°41’46”W. LBP 5636, 1 (tissue 27395), Mato Grosso, Cuiabá, rio Coxipó, rio Paraguai. LBP 8573, 1 (tissue 43355), Mato Grosso, Barra do Bugres, rio Paraguai. LBP 9818, 6 (tissues 45230-45234, Mato Grosso do Sul, Miranda, rio Paraguai. LBP 9862, 1 (tissue 45408), Mato Grosso do Sul, Miranda, rio Paraguai.

Steindachnerina insculpta (Fernández-Yépez, 1948)

Fig. 6,Tab. 2

Cutimata elegansAmaral Campos, 1945:46 [Brazil: rio Mogi Guaçu].-Britski, 1972:83 [Brazil, São Paulo, rio Paraná basin]. -Foresti et al., 1974:249 [karyotypes].-Nomura, 1977:727 [Brazil: rio Mogi-Guaçu; meristic].-Nomura, Taveira, 1979:331 [life history].

Cruxentina insculptaFernández-Yépez, 1948:53, figs. 27, 28 [type-locality: Brazil: São Paulo, rio Tatuhy (= Tatuí); autorship cited as Amaral Campos].-Britski, 1969:200, 203 [correction of originally cited authorship, depositary and collector; meristic and morphometrics].-Fowler, 1975:368 [reference].-Vari, 1989, tables 2, 3 [assignment to Steindachnerina].

Curimata elegans.-Gomes, Monteiro, 1955:88, 103, 129 [São Paulo, Pirassununga; occurrence in flowing and still waters].-Oliveira et al., 1988:594 [in part, Brazil, São Paulo, rio Mogi-Guaçu and Botucatu; not Minas Gerais, Três Marias; chromosome counts].

Pseudocurimata elegans elegans.-Godoy, 1975: 88, 103, 129, fig. 132A, 133 [Brazil: São Paulo, rio Mogi-Guassu; life history data; meristic and morphometrics; not cited occurrence in drainage basins other than upper rio Paraná].

Steindachnerina insculpta.-Venere, Galetti Júnior, 1989:18, 19, fig. 19 [upper rio Paraná basin, rio Mogi-Guaçu and rio Passa-Cinco; karyotype information].-Vari, 1991:71 [in part Brazil, material from upper rio Paraná basin above Sete Quedas; not from rio Paraguai system and from Goiás]. -Koerber et al., 2017:16 [reference; type locality mistakenly referred to as being in Venezuela].

?Curimata nasa.-Géry et al., 1987:425, fig. 41 [material from arroyo Porooco and Itaipu Lake, rio Paraná basin].

Fig. 6 Steindachnerina insculpta, LBP 11381, 78.7 mm SL, Brazil, São Paulo, Birigui, Rio Tietê, upper Rio Paraná. 

Diagnosis. Steindachnerina insculpta can be distinguished from congeners, except S. binotata, S. biornata, S. dobula, S. hypostoma, S leucisca, S. nigrotaenia, S. notograptos, and S. varii by possessing a plain dorsal fin (vs. dorsal fin with a spot of dark pigmentation on the basal portion of the middle rays). Steindachnerina insculpta has 40 to 45 scales in the lateral line series, differing from S. binotata, S. leucisca, S. hypostoma, S. notograptos, and S. gracilis which have 46 or more scales and from S. biornata which has 31 to 34 scales in the lateral line series. Steindachnerina insculpta differs from S. dobula and S. varii by having a dark midlateral stripe extending from rear of opercle posteriorly onto middle rays of caudal fin (vs. dark midlateral stripe beginning posterior to vertical line from dorsal-fin origin). Steindachnerina insculpta differs from the very similar S. nigrotaenia by the lower modal number of perforated scales in the lateral line series (40 to 45, 42 most frequent vs. 43 to 48, 45 most frequent and 43 in only 2.27% of specimens examined for this feature; Fig. 3), and by the total number of scale rows in the transverse series from the dorsal-fin origin to the pelvic-fin base (12 to 14, 12.5 most frequent, 13.5 and 14 less frequent vs. 13 to 16, 14 most frequent; 13 in only 8.16% of specimens examined for this feature; Fig. 4). Steindachnerina insculpta can also be discriminated from its congeners by a combination of the following features: multiple lobulated fleshy processes on the roof of the oral cavity, absence of a wide, flattened prepelvic region of the body, 40 to 45 scales on the lateral line, 12 to 14 scales on the transversal series, lack of a dark spot on basal portions of the dorsal fin and the presence of a dark midlateral stripe along the body.

Description. Morphometric data of Steindachnerina insculpta are summarized in Tab. 2. Body relatively elongate, somewhat compressed. Dorsal profile of head very slightly convex anteriorly, straight from above orbit to rear of head. Dorsal profile of body straight or very slightly convex from rear of head to origin of dorsal fin; straight at base of dorsal fin; straight from base of last dorsal-fin ray to caudal peduncle. Dorsal surface of body transversely rounded anteriorly, with indistinct median keel immediately anterior to dorsal fin, smoothly rounded transversely posterior to dorsal fin. Ventral profile of body gently curved from tip of lower jaw to caudal peduncle. Prepelvic region obtusely flattened, with median series of scales proximate to pelvic-fin origin, median series less regularly arranged anteriorly. Median keel barely discernable posterior to pelvic-fin origin.

Dorsal-fin margin rounded; anteriormost rays three to three and half times length of ultimate ray. Pectoral-fin margin acute, extending one-half to two-third distance to origin of pelvic fin in smaller adults, barely beyond that point in the largest examined specimens. Pelvic-fin margin acute; pelvic fin extending about one-half distance to origin of anal fin. Caudal fin forked. Adipose fin well developed. Anal fin emarginate, anteriormost branched rays two and one-half to three times the length of ultimate ray.

Head pointed in profile; upper jaw distinctly longer, mouth inferior; buccopharyngeal complex on roof of oral cavity in adults consisting of multiple lobulated fleshy bodies; nostrils very close, anterior circular, posterior crescent-shaped with aperture closed by thin flap of skin separating nares; adipose eyelid present, more developed anteriorly, with broad, vertically ovoid opening over center of eye.

Total pored lateral-line scales 40 (3), 41(13), 42(31), 43(28), 44(1) or 45(1); two to three scales over caudal fin. Longitudinal series of scales from dorsal-fin origin to lateral line 6(51), 6.5(23) or 7(4); longitudinal series of scales from lateral line to base of pelvic fin 5(26), 5.5(49) or 6(3); total lateral series from dorsal-fin origin to pelvic-fin base 12(23), 12.5(29), 13(20), 13.5(5) or 14(1); longitudinal series of scales from lateral line to first ray of anal fin 4.5(9) or 5(69); circumpeduncular scales 16(77) or 17(1); predorsal scales 10(7), 11(38), 12(17) or 13(5); scales from end of dorsal fin to adipose fin 12(6), 13(34) or 14(37); scales from end of adipose fin to anteriormost dorsal caudal-fin ray 8(3), 9(14), 10(35), 11(16) or 12(4); prepelvic scales 19(3), 20(11), 21(26), 22(11), 23(15) or 24(4); scales from pelvic fin to anus 7(5), 8(23), 9(38), 10(10) or 11(1); scales from anus to anal-fin origin 2(8), 3(51) or 4(2); scales from end of anal fin to anteriormost ventral caudal-fin ray 9(13), 10(37), 11(19) or 12(4); all scales of lateral line pored, canals in scales of lateral line straight.

Dorsal-fin rays ii-iii,8,ii(76); pectoral-fin rays i,12(2), i,13(17), i,14(40), i,15(18) or i,16(1); pelvic-fin rays i,7(1), i,8(76) or i,9(1); anal-fin rays ii-iii,6,ii(76) or iii5,ii(1); caudal-fin rays i,8,9,i(78). Total vertebrae 28(8) or 29(3).

Coloration in alcohol. Overall coloration of specimens retaining guanine on scales silvery to silvery golden, darker on dorsal portions of head and body. Ground coloration of specimens lacking guanine on scales tan to yellow, darker dorsally, with irregular patch of dark pigmentation extending from rear of orbit across opercle; degree of intensity of dark pigmentation and extent of patch variable among individuals. Irregular, dark longitudinal stripe extending along lateral line from supracleithrum to base of middle caudal-fin rays. Stripe slightly wider posteriorly, continuous posteriorly with dusky stripe on middle caudal-fin rays. Stripe in adults continuous, about one scale wide; not as well developed in juveniles, consisting of discrete spots surrounding pores of lateral line. Caudal-fin stripe more prominent on anterior portion of rays. Anterior margin and distal portions of dorsal fin typically dusky in adults. Ventral lobe and dorsal rays of dorsal lobe of caudal fin dusky. All caudal-fin rays outlined by small chromatophores on membranes. Adipose fin dusky distally. Dorsal fin with dusky anterior region in many specimens, without any discrete spot on middle rays. Other fins hyaline.

Sexual dimorphism. Secondary sexual characters were not found.

Geographical distribution. Steindachnerina insculpta is restricted to the upper rio Paraná basin (Fig. 5).

Material examined. All from Brazil, upper rio Paraná. CAS 20312, holotype of Cruxentina insculpta, 95.3 mm SL, São Paulo, rio Tatuhy (= Tatuí, SP). MZUSP 1376, 1, paratype of Cruxentina insculpta, 105.6 mm SL, rio Tatuhy (= Tatuí). USNM 295275, 5, 76.3-104.0 mm SL; MZUSP 20381, 98, Alfredo de Castilho, córrego do Moinho. USNM 295272, 5 (4, 66.7-78.4 mm SL); MZUSP 21515, 26, Miguelópolis, Volta Grande reservoir. USNM 295271, 22 (5, 51.8-87.8); MNRJ 5669, 5 (3, 74.4-81.7); MZUSP 20700, 15; MZUSP 20704, 33; MZUSP 20739, 19; MZUSP 20741, 1; MZUSP 20744, 2; MZUSP 20750, 74 (10, 80.9-90.4); MZUSP 20791, 1; BMNH 1946.12.23:97-111, 11, 69.9-79.5 mm SL, rio Mogi-Guaçu, Emas. MZUSP 20672, 36; MZUSP 20691, 17, rio Mogi-Guaçu, Cachoeira de Emas. USNM 295274, 9; MZUSP 20711,10, rio Mogi-Guaçu, Pirassununga. CAS 41729, 4 66.0-91.9 mm SL; NWW 67002, 3; NMW 68914, 2, Piracicaba, rio Piracicaba. MZUSP 20755, 64; MZUSP 20758, 37; MZUSP 20768, 1, Corumbataí, rio Corumbataí. MZUSP 21317, 10, 88.7-101.4 mm SL; USNM 295266, 2, Borborema, rio Tietê. MZUSP 21525, 1, Marimbondo, rio Grande. MZUSP 21429, 106, Ilha Solteira, rio Paraná. MZUSP 21603, 2, Pedreira, rio Jaguari. MZUSP 20865, 1, rio Pardo, usina de Limoeiro. MZUSP 21471, 1, Botucatu. Minas Gerais: MZUSP 21505, 2, Alfenas, rio Grande, Furnas reservoir. MZUSP 21502, 1, Boa Esperança, rio Grande. Mato Grosso do Sul: MZUSP 20683,15; MZUSP 20714, 84; MZUSP 20853, 18, Três Lagoas, usina de Jupiá, rio Paraná. MZUSP 20824, 1; MZUSP 20894, 1, Três Lagoas, rio Sucuriú. Paraná: MZUSP 21620, 103, rio Paraná, Guaíra, above Sete Quedas. LBP 3192, 4 (tissues 19389-19390), São Paulo, Conchas, rio Tietê; LBP 5207, 8 (tissue 26343), Paraná, Porto Rico, rio Paraná; LBP 6671, 1 (tissue 32081); LBP 19770, 1 (tissue 32158), Paraná, Marilena, rio Paraná; NUP 1424, 4; NUP 4576, 6, Guaíra, rio Paraná, Itaipu reservoir. NUP 1753, 7, Santa Helena, rio Paraná, Itaipu reservoir. NUP 7513, 4, Foz do Iguaçu, rio Paraná, Reservatório Itaipu.

Discussion

Populations of Steindachnerina from the rio Paraguai basin and from the upper rio Paraná basin were considered by Vari (1991:71) as belonging to a single species (S. insculpta) but studies carried out on these populations have allowed us to conclude that they belong to two distinct species: S. insculpta (upper rio Paraná basin) and S. nigrotaenia (rio Paraguai basin), supplemented by the very distinct S. brevipinna (rio Paraguai basin) (Fig. 7). Although the first two species are very similar in many morphological aspects, they might be diagnosed from each other firstly on the basis of the number of scales in the lateral line and number of scales in the transverse line, and they can be further diagnosed by S. brevipinna by the lack of the dark spot on basal portions of the dorsal fin, and by modal differences in the number of scales of the lateral line and of the transverse line (Figs. 3-4). Furthermore, a molecular analysis strongly supports the recognition of the three species.

Fig. 7  Steindachnerina brevipinna , LBP 9753, 51.6 mm SL, Brazil, Mato Grosso do Sul, Nova Andradina, upper Rio Paraná. 

Vari (1991) interpreted the presence of Steindachnerina insculpta (= S. nigrotaenia) populations in the rio Paraguai basin as a result of a possible introduction of specimens originated from the upper rio Paraná. This interpretation was first a consequence of the supposition that S. nigrotaenia was a synonym of S. brevipinna, and secondarily because Vari (1991) had only a small number of specimens from the rio Cuiabá collected at that time. That small number of lots concentrated in a small area of the rio Paraguai basin and geographically distant from populations of the upper rio Paraná led him to hypothesize a human-mediated introduction to explain the disjoint distribution of what he considered to be populations of the same species (S. insculpta). Obviously, the present findings disprove his hypothesis, showing that he was working with two distinct species, S. insculpta and S. nigrotaenia.

Several pieces of evidence indicate a gap separating the distributions of Steindachnerina nigrotaenia (upper rio Paraguai and lower rio Paraná) from that of S. insculpta (upper rio Paraná, upriver Itaipu). Inventories in the rio Paraná below Itaipu reported neither S. insculpta nor S. nigrotaenia, and indicated, instead, the presence of S. brevipinna, S. biornata and S. conspersa (e.g., López et al., 2003; 2005). In addition, many species of the middle rio Paraná were incorporated to the ichthyofauna of the upper rio Paraná after the construction of the Itaipu dam (Agostinho et al., 1992, 1997; Júlio Júnior et al., 2009), including S. brevipinna that that is genetically distinct of the sympatric S. insculpta (Oliveira et al., 2002), but no evidence of S. nigrotaenia in that section has been reported recently (Graça, Pavanelli, 2007; Ota et al., 2018).

Specimens identified as Curimata nasa (Steindachner, 1882) by Géry et al. (1987:425) from Paraguay were considered by Vari (1991:71) to belong to S. insculpta (sensuVari, 1991). However, the five specimens mentioned by Géry et al. (1987) are from different localities: two of them collected in the rio Paraguai (possibly S. nigrotaenia), and three other specimens in the rio Paraná basin (S. insculpta). Nevertheless, any definitive decision on the status of these specimens without a direct examination is premature. More recently, Koerber et al. (2017:16) in the checklist of fishes from Paraguay cited S. brevipinna, S. conspersa and S. insculpta, the latter being S. nigrotaenia according to the concept herein established; they also erroneously refer to the type locality of S. insculpta as being in Venezuela, disregarding that Britski (1969:201), and Vari (1991:72) had already clarified this question. Aguilera et al. (2018) also reported the first occurrence of S. insculpta in Argentina, which, based on their description, the species in question also represents S. nigrotaenia.

After the revision of Steindachnerina (Vari, 1991), two other species were described in the genus and another species is validated herein, then accumulating 25 species of Steindachnerina. Another species, Cyphocharax corumbae (Pavanelli & Britski, 1999) was originally described in Steindachnerina but recently transferred to Cyphocharax (Melo et al., 2018). Presently, five Steindachnerina species are known from the rio La Plata basin (Paraná, Paraguai and Uruguai rivers): S. brevipinna (rio Paraná, rio Paraguai, and rio Uruguai), S. conspersa (rio Paraguai and lower rio Paraná), S. nigrotaenia (upper rio Paraguai), S. insculpta (upper rio Paraná), and S. biornata (rio Uruguai, lower rio Paraná and coastal rivers of southern Brazil).

This study resolves a long-standing question in the systematics of Steindachnerina and adds one more species to the ichthyofauna of the rio Paraguai. Further research might investigate the phylogenetic placement of S. nigrotaenia within the recently proposed S. dobula clade (sensuMelo et al., 2018) and also compare morphological features of the three related species occurring in the La Plata system.

Comparative material examined. Steindachnerina brevipinna: LBP 659, 2, Mato Grosso, Poconé, rio Paraguai; LBP 3728, 1, Mato Grosso do Sul, Aquidauana, rio Paraguai; LBP 4665, 4, Mato Grosso do Sul, Rio Baia, rio Paraguai; LBP 5804, 15, Mato Grosso, Santo Antônio do Laverger, rio Paraguai; LBP 6690, 34 (tissues 32153-32157), Paraná, Marilena, rio Paraná; LBP 8433, 9, Mato Grosso, Tangará da Serra, rio Paraguai; LBP 8448, 5, Mato Grosso, Cáceres, rio Paraguai; LBP 9753, 45, Mato Grosso do Sul, Nova Andradina, rio Paraná; LBP 13338, 2, Mato Grosso, Poconé, rio Paraguai; NUP 1431, 4 (102.2-114.2 mm SL), Paraná, Guaíra, rio Paraná, Itaipu reservoir; NUP 2895, 62 (5, 88.3-104.5 mm SL), Paraná, Porto Rico, Ilha Porto Rico, rio Paraná, Lagoa Jenipapo. Steindachnerina elegans: LBP 8272, 2, Minas Gerais, Jaíba, rio Verde Grande, rio São Francisco.

Acknowledgments

We thank James Maclaine, Oliver Crimmen and Ralf Britz (BMNH) for assistance during collection visits. We also thank James Maclaine for sending photographs of the lectotype of Curimatus nigrotaenia and for the explanations regarding the exchanges of type material that took place at BMNH. We are indebted to Giuliano Doria of the Museo Civico di Storia Naturale of Genova, for the loan of the syntypes of C. nigrotaenia, and to Heiko Bleher who facilitated the loan of the specimens. We thank Marcelo R. Britto who provided some data on material from MNRJ collection, Carla S. Pavanelli for sending specimens from the Itaipu area, Beatriz F. Dorini for helping with molecular data and Veronica Slobodian who provided important radiographs. We are grateful to Naércio A. Menezes for comments and suggestions on the manuscript, to Fernando Dagosta for assistance with tables, figures and other suggestions, and to Brian L. Sidlauskas and two anonymous reviewers for improving the manuscript. The project was supported by a research grant from the CNPq (HAB) and CNPq Universal 404991/2018-1 (BFM), and by FAPESP 16/11313-8 (BFM), and FAPESP 14/26508-3 (CO).

References

Agostinho AA, Júlio Júnior HF, Borghetti JR. Considerações sobre os impactos dos represamentos na ictiofauna e medidas para sua atenuação. Um estudo de caso: reservatório de Itaipu. Rev Unimar. 1992; 14(Suppl.):89-107. [ Links ]

Agostinho AA, Júlio Júnior HF, Gomes LC, Bini LM, Agostinho CS. Composição, abundância e distribuição espaço-temporal da ictiofauna. In: Vazzoler AEAM, Agostinho AA, Hahn NS, editors. A planície de inundação do alto rio Paraná: aspectos físicos, biológicos e socioeconômicos. Maringá: EDUEM & UEM-Nupelia; 1997. p.179-208. [ Links ]

Aguilera G, Terán GE, Alonso F, Mirande JM. First record of Steindachnerina insculpta (Fernández-Yépez, 1948) (Characiformes, Curimatidae) in Argentina. Check List [serial on the Internet]. 2018; 14(6):1017-20. Available from: https://doi.org/10.15560/14.6.1017Links ]

Amaral Campos A. Sobre os caracídeos do rio Mogi-Guaçu (Estado de São Paulo). Arq Zool Est São Paulo. 1945; 4(11):431-66. [ Links ]

Boulenger GA. Descriptions of new fishes and reptiles discovered by Dr. F. Silvestri in South America. Ann Mag Nat Hist. 1902; 9(52):284-88. (Series 7). [ Links ]

Britski HA. Lista dos tipos de peixes das coleções do Departamento de Zoologia da Secretaria da Agricultura do Estado de São Paulo. Pap Avulsos Zool. 1969; 22(19):197-215. [ Links ]

Britski HA. Peixes de água-doce do estado de São Paulo: Sistemática. In: Comissão Interestadual da bacia do Paraná-Uruguai. Poluição e piscicultura. São Paulo: Faculdade de Saúde Pública da USP. 1972; p.83-108. [ Links ]

Britski HA. Revalidação de Steindachnerina nigrotaenia (Boulenger, 1902) (Pisces, Characiformes). In: 7º Encontro de Biólogos. Ribeirão Preto: CRB-1 (SP, MT, MS); 1996. p. 03-21. [ Links ]

Britski HA, Silimon KZS, Lopes BS. Peixes do Pantanal: manual de identificação. 1st ed. Brasília/Corumbá: Embrapa; 1999. [ Links ]

Britski HA, Silimon KZS, Lopes BS. Peixes do Pantanal: manual de identificação . 2nd ed. Brasília: Embrapa; 2007. [ Links ]

Edgar R. Muscle: a multiple sequence alignment method with reduced time and space complexity. BMC Bioinformatics [serial on the Internet]. 2004; 5(113):1-19. Available from: https://doi.org/10.1186/1471-2105-5-113Links ]

Eigenmann CH. Catalogue of the freshwater fishes of tropical and South Temperate America. In: Scott WB (editor). Reports of the Princeton University Expeditions to Patagonia, 1896-1899. Stuttgart: E. Schweizerbart’sche. Verlagsbuchhandlung ; 1910. p.375-511. (Zöology, vol 3, part 4). [ Links ]

Eigenmann CH. The freshwater of British Guiana, including a study of the ecological grouping of species, and the relation of the fauna of the plateau to that of the lowlands. Mem Carnegie Mus. 1912; 5(1):1-578. [ Links ]

Fernández-Yépez A. Los Curimatidos (peces fluviales de Sur América): Catalogo descriptivo con nuevas adiciones genericas y especificas. Boletín Taxónomica del Laboratorio de Pesqueria de Caiguire. 1948; 1:1-86. [ Links ]

Foresti F, Oliveira LM, Angeleli WA. Caracterização cromossômica em peixes do gênero Curimatus (Cypriniformes: Curimatidae). Cienc Cult. 1974; 26:249. [ Links ]

Fowler HW. Os peixes de água doce do Brasil. Arq Zool Est São Paulo . 1950; 6:205-404. [ Links ]

Fowler HW. A catalog of world fishes (XXII). Volume (III) Orders Mormyrida, Characida, Gymnotida, Silurida and Anguillida. Quat J Taiwan Mus. 1975; 28(1/2):1-124. [ Links ]

Géry J, Mahnert V, Dlouhy C. Poissons Characoides non Characidae du Paraguay (Pisces, Ostariophysi). Rev Suisse Zool. 1987; 94(2):357-464. [ Links ]

Godoy MP. Peixes do Brasil, subordem Characoidei; bacia do Rio Mogi Guassu. Piracicaba: Editora Franciscana; 1975. [ Links ]

Gomes AL, Monteiro FP. Estudo da população total de peixes da represa da Estação Experimental de Biologia e Piscicultura, em Pirassununga, São Paulo. Rev Biol Marina Valpso. 1955; 6:82-154. [ Links ]

Graça WJ, Pavanelli CS. Peixes da planície de inundação do alto rio Paraná e áreas adjacentes. Maringá: Eduem; 2007. [ Links ]

Günther ACLG. Catalogue of the fishes in the British Museum. Catalogue of the Physostomi, containing the families Siluridae, Characinidae, Haplochitonidae, Stemoptychidae, Scopelidae, Stomiatidae in the collection of the British Museum. London: The Trustees; 1864. [ Links ]

International Commission on Zoological Nomenclature (ICZN). International Code of Zoological Nomenclature. 4th ed. London: International Trust for Zoological Nomenclature Natural History Museum; 1999. [ Links ]

Júlio Júnior HF, Dei Tós C, Agostinho AA, Pavanelli CS. A massive invasion of fish species after eliminating a natural barrier in the upper rio Paraná basin. Neotrop Ichthyol [serial on the Internet]. 2009; 7(4):709-18. Available from: http://dx.doi.org/10.1590/S1679-62252009000400021Links ]

Kearse M, Moir R, Wilson A, Stones-Havas S, Cheung M, Sturrock S et al. Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics [serial on the Internet]. 2012; 28(12):1647-49. Available from: https://doi.org/10.1093/bioinformatics/bts199Links ]

Koerber S, Vera-Alcaraz HS, Reis RE. Checklist of the Fishes of Paraguay (CLOFPY). Ichthyol Contrib Peces Criollos. 2017; 53:1-99. [ Links ]

Litz TO, Koerber S. Check list of the freshwater fishes of Uruguay (CLOFF-UY). Ichthyol Contrib Peces Criollos . 2014; 28:1-40. [ Links ]

López HL, Miquelarena AM, Menni RC. Lista comentada de los peces continentales de la Argentina. Buenos Aires: Facultad de Ciencias Naturales y Museo UNLP; 2003. (Série Técnica y Didáctica. N. 5). [ Links ]

López HL, Miquelarena AM, Pontes Gomes J. Biodiversidad y distribución de la ictiofauna Mesopotámica. INSUGEO. 2005; 14:311-54. [ Links ]

Melo BF, Benine RC, Mariguela TC, Oliveira C. A new species of Tetragonopterus Cuvier, 1816 (Characiformes: Characidae: Tetragonopterinae) from the rio Jari, Amapá, northern Brazil. Neotrop Ichthyol [serial on the Internet]. 2011; 9(1):49-56. Available from: http://dx.doi.org/10.1590/S1679-62252011000100002Links ]

Melo BF, Sidlauskas BL, Hoekzema K, Vari RP, Dillman CB, Oliveira C. Molecular phylogenetics of Neotropical detritivorous fishes of the family Curimatidae (Teleostei: Characiformes). Mol Phylog Evol [serial on the Internet]. 2018; 127:800-12. Available from: https://doi.org/10.1016/j.ympev.2018.06.027Links ]

Miller MA, Pfeiffer W, Schwartz T. Creating the CIPRES Science gateway for inference of large phylogenetic trees [Internet]. New Orleans: Proceedings of the Gateway Computing Environments Workshop (GCE); 2010. p.1-8. Available from: https://doi.org/10.1109/GCE.2010.5676129Links ]

Nomura H. Caracteres merísticos do saguiru, Curimatus elegans Steindachner, 1874 do rio Mogi Guaçu, São Paulo (Osteichthyes, Curimatidae). Rev Bras Biol. 1977; 37(4):727-29. [ Links ]

Nomura H, Taveira ACD. Biologia do saguiru, Curimatus elegans Steindachner, 1874 do Mogi Guaçu, São Paulo (Osteichthyes, Curimatidae). Rev Bras Biol . 1979; 39(2):331-39. [ Links ]

Oliveira C, Almeida Toledo LF, Foresti F, Britski HA, Toledo Filho SA. Chromosome formulae of Neotropical freswater fishes. Rev Bras Genét. 1988; 11(3):577-624. [ Links ]

Oliveira AV, Prioli AJ, Prioli SMAP, Pavanelli CS, Júlio Jr HF, Panarari RS. Diversity and genetic distance in populations of Steindachnerina in the upper Paraná river foodplain of Brazil. Genetica. 2002; 115:259-267. [ Links ]

Ota RR, Deprá GC, Graça WJ, Pavanelli CS. Peixes da planície de inundação do alto rio Paraná e áreas adjacentes: revised, annotated and updated. Neotrop Ichthyol [serial on the Internet]. 2018; 16(2):e170094. Available from: http://dx.doi.org/10.1590/1982-0224-20170094Links ]

Pavanelli CS, Britski HA. Description of a new species of Steindachnerina(Teleostei: Characiformes: Curimatidae) from the upper Rio Paraná basin, Brazil. Ichthyol Explor Freshw. 1999; 10(3):211-16. [ Links ]

Polaz CNM, Melo BF, Britzke R, Resende EK, Machado FA, Lima JAF, Petrere Júnior M. Fishes from the Parque Nacional do Pantanal Matogrossense, upper Paraguai River basin, Brazil. Check List [serial on the Internet]. 2014; 10(1):122-30. Available from: http://dx.doi.org/10.15560/10.1.122Links ]

Ringuelet RA. Zoogeografía y ecología de los peces de aguas continentales de Argentina y consideraciones sobre las áreas ictiológicas de América del Sur. Ecosur. 1975; 2(3):1-122. [ Links ]

Stamatakis A. RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenenies. Bioinformatics [serial on the Internet]. 2014; 30(9):1312-13. Available from: https://doi.org/10.1093/bioinformatics/btu033Links ]

Tamura K, Stecher G, Peterson D, Filipski A, Kumar S. MEGA6: molecular evolutionary genetics analysis version 6.0. Mol Biol Evol [serial on the Internet]. 2013; 30(12):2725-29. Available from: https://doi.org/10.1093/molbev/mst197Links ]

Tortonese E. Catalogo del tipi de pesci del Museo Civico di Storia Naturale di Genova. (Parte I). Annali del Museo Civico di Storia Naturale ‘Giacomo Doria’. 1961; 72:179-91. [ Links ]

Vari RP. A Phylogenetic study of the tropical Characiform Family Curimatidae (Pisces: Ostariophysi). Smithson Contrib Zool. 1989; 47:1-73. [ Links ]

Vari RP. Systematics of the Neotropical Characiform genus Steindachnerina Fowler (Pisces: Ostariophysi). Smithson Contrib Zool . 1991; 507:1-118. [ Links ]

Venere PC, Galetti Júnior PM. Chromosome evolution and phylogenetic relationships of some neotropical Characiformes of the Family Curimatidae. Rev Bras Gen. 1989; 12(1):17-25. [ Links ]

Xia X. DAMBE5: a comprehensive software package for data analysis in molecular biology and evolution. Mol Biol Evol [serial on the Internet]. 2013; 30(7):1720-28. Available from: https://doi.org/10.1093/molbev/mst064Links ]

Received: June 10, 2018; Accepted: February 20, 2019

Edited by

Paulo Lucinda

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