Abstract:

The rio das Cinzas and rio Itararé basins are important tributaries of the rio Paranapanema, a major left-bank tributary of the upper rio Paraná basin. The upper reaches of both drainages intersect the northern portion of the APA Escarpa Devoniana (Devonian Escarpment Environment Protection Area, EPA), an area considered crucial for the preservation of biodiversity. This study aimed to provide an ichthyofaunistic inventory for headwater streams of the rio das Cinzas and rio Itararé basins, both within and off of the borders of the EPA, increasing the knowledge about the species already known from streams in the rio Paranapanema system. We found 32 fish species, representing five orders and 12 families. Siluriformes was the richest order followed by Characiformes. Ten species presented high abundance of sampled individuals, contributing with 85.0% of all specimens. Most species were classified as occasional (captured in less than 25% of the samples) and only two species were classified as constant (captured in more than 50% of samples), which may reflect the altitudinal gradient. We did not record any introduced species in the sampled streams, reinforcing the need of, at least, maintaining the original area of the EPA. We expanded the geographic distribution of Astyanax bifasciatus to the upper rio Paraná system, confirming the cases of faunal exchange among several drainage systems in the limits of the Ponta Grossa Arch. Finally, we identified three putatively undescribed species that show a large biodiversity knowledge shortfall for the region studied.

Keywords:
Altitudinal gradient; Devonian Escarpment; faunal exchange; upper rio Paraná

Resumo:

As bacias do rio das Cinzas e do rio Itararé são afluentes importantes do rio Paranapanema, um dos principais afluentes da margem esquerda do alto rio Paraná. Os trechos superiores de ambas as drenagens interceptam a porção norte da Área de Proteção Ambiental (APA) da Escarpa Devoniana, uma área considerada crucial para a preservação da biodiversidade. Este trabalho teve como objetivo fornecer um inventário ictiofaunístico para riachos de cabeceira das bacias do rio das Cinzas e do rio Itararé, dentro e fora das fronteiras da APA, aumentando o conhecimento sobre as espécies já conhecidas dos riachos do rio Paranapanema. Encontramos 32 espécies de peixes, representando cinco ordens e 12 famílias. Siluriformes foi a ordem mais rica seguida por Characiformes. Dez espécies apresentaram alta abundância de indivíduos amostrados, contribuindo com 85,0% de todos os espécimes. A maioria das espécies foi classificada como ocasional (capturada em menos de 25% das amostras) e apenas duas espécies foram classificadas como constantes (capturadas em mais de 50% das amostras), o que pode refletir o gradiente altitudinal. Não registramos nenhuma espécie introduzida nos riachos amostrados, reforçando a necessidade de, pelo menos, manter a área original do APA. Expandimos a distribuição geográfica de Astyanax bifasciatus para o sistema do alto rio Paraná, confirmando os casos de intercâmbio faunístico entre vários sistemas de drenagem nos limites do Arco de Ponta Grossa. Finalmente, identificamos três espécies supostamente não descritas que mostram um grande déficit de conhecimento da biodiversidade para a região estudada.

Palavras-chave:
Alto rio Paraná; Escarpa Devoniana; gradiente altitudinal; intercâmbio faunístico

Introduction

Freshwater environments provide numerous ecosystem goods and services that will be lost due to the rapid degradation of aquatic bodies by human activity (Azevedo-Santos et al. 2019aAZEVEDO-SANTOS, V.M., FREDERICO, R.G., FAGUNDES, C.K., POMPEU, P.S., PELICICE, F.M., PADIAL, A.A., NOGUEIRA, M.G., FEARNSIDE, P.M., LIMA, L.B., DAGA, V.S., OLIVEIRA, F.J.M., VITULE, J.R., CALLISTO, M., AGOSTINHO, A.A., ESTEVES, F.A., LIMA-JUNIOR, D.P., MAGALHÃES, A.L.B., SABINO, J., MORMUL, R.P., GRASEL, D., ZUANON, J., VILELLA, F.S. & HENRY, R. 2019a. Protected areas: a focus on Brazilian freshwater biodiversity. Divers. Distrib. 25(3): 442-448.). Local biodiversity inventories are essential in environmental management projects, and provide support for studies involving ecological or biogeographic aspects (Oliveira et al. 2014OLIVEIRA, A.G., GOMES, L.C., LATINI, J.D. & AGOSTINHO, A.A. 2014. Implications of using a variety of fishing strategies and sampling techniques across different biotopes to determine fish species composition and diversity. Nat. Conservação 12(2): 112-117.; Frota et al. 2019FROTA, A., MESSAGE, H.J., OLIVEIRA, R.C., BENEDITO, E. & GRAÇA, W.J. 2019. Ichthyofauna of headwater streams from the rio Ribeira de Iguape basin, at the boundaries of the Ponta Grossa Arch, Paraná, Brazil. Biota Neotrop. 19(1): e20180666 http://dx.doi.org/10.1590/1676-0611-BN-2018-0666 (last access 17 June 2019).
http://dx.doi.org/10.1590/1676-0611-BN-2... ). Given that many species in the Neotropical region remain undescribed (Ota et al. 2015OTA, R.R., MESSAGE, H.J., GRAÇA, W.J. & PAVANELLI, C.S. 2015. Neotropical Siluriformes as a model for insights on determining biodiversity of animal groups. PLoS ONE 10(7): e0132913.) or poorly known, species inventories are essential for widening the known geographic range of the species, and also favor the discovery of new species (Carvalho 2009CARVALHO, C.J.B. 2009. Padrões de endemismos e a conservação da biodiversidade. Megadiversidade(1-2): 77-86.; Whittaker et al. 2005WHITTAKER, R.J., ARAÚJO, M.B., JEPSON, P., LADLE, R.J., WATSON, J.E.M. & WILLIS, K.J. 2005. Conservation Biogeography: assessment and prospect. Divers. Distrib. 11(1): 3-23.). In this way, the conservationist practice can only be realized knowing the biodiversity that is intended to be conserved, and inventories are crucial for this finality (Carvalho 2009CARVALHO, C.J.B. 2009. Padrões de endemismos e a conservação da biodiversidade. Megadiversidade(1-2): 77-86.).

The Neotropical region, the most ichthyologically rich biogeographical domain in the world, comprises about 9,000 fish species (Birindelli & Sidlauskas 2018BIRINDELLI, J.L.O. & SIDLAUSKAS, B.L. 2018. Preface: How far has Neotropical Ichthyology in twenty years? Neotrop. Ichthyol. 16(3): e180128.). Of these, many are small and inhabit mostly low-order watercourses (Castro 1999CASTRO, R.M.C. 1999. Evolução da ictiofauna de riachos sul americanos: padrões gerais e possíveis processos casuais. In Ecologia de peixes de riachos: estado atual e perspectivas (E.P. Caramaschi, R. Mazzoni & P.R. Peres-Neto, eds). UFRJ, Rio de Janeiro, p.139-155.; Winemiller et al. 2008WINEMILLER, K.O., AGOSTINHO, A.A. & CARAMASCHI, E.P. 2008. Fish ecology in tropical streams. In Tropical stream ecology (D. Dudgeon, ed). Academic Press, San Diego, p.107-146.), which are estimated to harbor about 50% of the freshwater fish richness in the tropics (Lowe-McConnell 1999LOWE-MCCONNELL, R.H. 1999. Estudos ecológicos de comunidades de peixes tropicais. EDUSP, São Paulo.; Winemiller et al. 2008WINEMILLER, K.O., AGOSTINHO, A.A. & CARAMASCHI, E.P. 2008. Fish ecology in tropical streams. In Tropical stream ecology (D. Dudgeon, ed). Academic Press, San Diego, p.107-146.). Even the highly impacted upper rio Paraná system has an estimated 310 species (236 natives), a number outdated by twelve years (Langeani et al. 2007LANGEANI, F., CASTRO, R.M.C., OYAKAWA, O.T., SHIBATTA, O.A., PAVANELLI, C.S. & CASATTI, L. 2007. Diversidade da ictiofauna do Alto Rio Paraná: composição atual e perspectivas futuras. Biota Neotrop. 7(3): 181-197 http://www.biotaneotropica.org.br/v7n3/pt/abstract?article+bn0340703 (last access 13 June 2019).
http://www.biotaneotropica.org.br/v7n3/p... ).

Most of the sampling effort that yielded specimens from the rio Paraná basin is due to projects sponsored by the electrical sector, as a consequence of the intense damming to which the basin is subject (Baumgartner et al. 2012BAUMGARTNER, G., PAVANELLI, C.S., BAUMGARTNER, D., BIFI, A.G., DEBONA, T. & FRANA, V.A. 2012. Peixes do Baixo Rio Iguaçu. Eduem, Maringá.). However, these projects focused mostly on larger rivers and, until recently, low-order streams (especially in the Paraná State) still lacked samplings. This scenario changed in recent years as researchers started to focus ecological and biogeographical aspects of several sub-basins of the alto rio Paraná system, increasing our knowledge about the fish species resident in its streams (e.g.,Castro et al. 2003CASTRO, R.M.C., CASATTI, L., SANTOS, H.F., FERREIRA, K.M., RIBEIRO, A.C., BENINE, R.C., DARDIS, G.Z.P., MELO, A.L.A., ABREU, T.X., BOCKMANN, F.A., CARVALHO, M., GIBRAN, F.Z. & LIMA, F.C.T. 2003. Estrutura e composição da ictiofauna de riachos do Rio Paranapanema, sudeste e sul do Brasil. Biota Neotrop. 3(1): 1-31. http://www.biotaneotropica.org.br/v3n1/pt/abstract?article+BN01703012003 (last access 13 June 2019).
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http://dx.doi.org/10.1590/1676-0611-BN-2... ; Frota et al. 2016aFROTA, A., DEPRÁ, G.C., PETENUCCI, L.M. & GRAÇA, W.J. 2016a. Inventory of the fish fauna from Ivaí River basin, Paraná State, Brazil. Biota Neotrop. 16(3): e20150151 http://dx.doi.org/10.1590/1676-0611-BN-2015-0151 (last access 13 June 2019).
http://dx.doi.org/10.1590/1676-0611-BN-2... ,bFROTA, A., GONÇALVES, E.V.R., DEPRÁ, G.C. & GRAÇA, W.J. 2016b. Inventory of the ichthyofauna from the Jordão and Areia river basins (Iguaçu drainage, Brazil) reveals greater sharing of species than thought. Check List 12(6): 1995.; Larentis et al. 2016LARENTIS, C., DELARIVA, R.L., GOMES, L.C., BAUMGARTNER, D., RAMOS, I.P. & SEREIA, D.A.O. 2016. Ichthyofauna of streams from the lower Iguaçu River basin, Paraná State, Brazil. Biota Neotrop. 16(3): e20150117 http://dx.doi.org/10.1590/1676-0611-BN-2015-0117 (last access 17 June 2019).
http://dx.doi.org/10.1590/1676-0611-BN-2... ; Cavalli et al. 2018CAVALLI, D., FROTA, A., LIRA, A.D., GUBIANI, E.A., MARGARIDO, V.P. & GRAÇA, W.J. 2018. Update on the ichthyofauna of the Piquiri River basin, Paraná, Brazil: a conservation priority area. Biota Neotrop. 18(2): e20170350. http://dx.doi.org/10.1590/1676-0611-BN-2017-0350 (last access 13 June 2019).
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http://dx.doi.org/10.1590/1676-0611-BN-2... ).

The rio Paranapanema covers about 900 km (most of which defining the border between Paraná and São Paulo states) from its headwaters in the Serra de Paranapiacaba (São Paulo State) to its mouth in the rio Paraná, being one of its major left-bank tributaries (Ziesler & Ardizzone 1979ZIESLER, R. & ARDIZZONE, G.D. 1979. The inland waters of Latin America. Copescal Technical Paper No. 1. Food and Agriculture Organization of the United Nations (FAO), Roma, 171 p.). Ichthyofaunistic inventories are available for some of the main sub-basins of the rio Paranapanema system within the Paraná State (Shibatta et al. 2002SHIBATTA, O.A., ORSI, M.L., BENNEMANN, S.T. & SILVA-SOUZA, A.T. 2002. Diversidade e distribuição de peixes na bacia do rio Tibagi. In: A Bacia do Rio Tibagi (M.E. Medri, E. Bianchini, O.A. Shibatta, J.A. Pimenta, eds.). M.E. Medri, Londrina, p. 403-424.; Castro et al. 2003CASTRO, R.M.C., CASATTI, L., SANTOS, H.F., FERREIRA, K.M., RIBEIRO, A.C., BENINE, R.C., DARDIS, G.Z.P., MELO, A.L.A., ABREU, T.X., BOCKMANN, F.A., CARVALHO, M., GIBRAN, F.Z. & LIMA, F.C.T. 2003. Estrutura e composição da ictiofauna de riachos do Rio Paranapanema, sudeste e sul do Brasil. Biota Neotrop. 3(1): 1-31. http://www.biotaneotropica.org.br/v3n1/pt/abstract?article+BN01703012003 (last access 13 June 2019).
http://www.biotaneotropica.org.br/v3n1/p... ; Cetra et al. 2012CETRA, M., BARRELLA, W., LANGEANI NETO, F., MARTINS, A.G., MELLO, B.J. & ALMEIDA, R.S. 2012. Fish fauna of headwater streams that cross Atlantic Forest of south São Paulo state. Check List 8(3): 421-425.; Cionek et al. 2012CIONEK, V.M., SACRAMENTO, P.A., ZANATTA, N., OTA, R.P., CORBETTA, D.F. & BENEDITO, E. 2012. Fishes from first order streams of lower Paranapanema and Ivaí rivers, upper Paraná River basin, Paraná, Brazil. Check List 8(6): 1158-1162.; Pagotto et al. 2012PAGOTTO, J.P.A., VERÍSSIMO, S., GOULART, E. & MISE, F.T. 2012. Fishes (Osteichthyes: Actinopterygii) from the Pirapó River drainage, upper Paraná River basin, Paraná state, Brazil. Check List 8(3): 463-468.; Costa et al. 2013COSTA, A.D.A., FERREIRA, D.G., SILVA, W.F., ZANATTA, A.S., SHIBATTA, O.A. & GALINDO, B.A. 2013. Fishes (Osteichthyes: Actinopterygii) from the Penacho stream, upper Paraná River basin, Paraná State, Brazil. Check List 9(3): 519-523.; Hoffmann et al. 2015HOFFMANN, A.C., NASCIMENTO, R.H.C. & SHIBATTA, O.A. 2015. Fish fauna from tributaries throughout the Tibagi River basin, upper Paraná basin, Brazil. Check List 11(6): 1815.; Cetra et al. 2016CETRA, M., MATTOX, G.M.T., FERREIRA, F.C., GUINATO, R.B., SILVA, F.V. & PEDROSA, M. 2016. Headwater stream fish fauna from the Upper Paranapanema River basin. Biota Neotrop. 16(3): e20150145. http://dx.doi.org/10.1590/1676-0611-BN-2015-0145 (last access 13 June 2019).
http://dx.doi.org/10.1590/1676-0611-BN-2... ; Claro-García et al. 2018CLARO-GARCÍA, A., ASSEGA, F.M. & SHIBATTA, O.A. 2018. Diversity and distribution of ichthyofauna in streams of the middle and lower Tibagi river basin, Paraná, Brazil. Check List 14(1): 43-53.), but the fishes from the rio das Cinzas and rio Itararé basins remain unknown to the scientific literature, except for one tributary of the former (Costa et al. 2013COSTA, A.D.A., FERREIRA, D.G., SILVA, W.F., ZANATTA, A.S., SHIBATTA, O.A. & GALINDO, B.A. 2013. Fishes (Osteichthyes: Actinopterygii) from the Penacho stream, upper Paraná River basin, Paraná State, Brazil. Check List 9(3): 519-523.) and a few streams draining into the latter (Cetra et al. 2016). Thus, this study aimed to provide a new, more inclusive fish inventory for headwater streams of the rio das Cinzas and rio Itararé basins in the Paraná State.

Material and Methods

Study area

The rio das Cinzas basin is the main watercourse of the Norte Pioneiro mesoregion of the Paraná State. Its headwaters are located in the Serra de Furnas, in the municipality of Piraí do Sul, west of the Devonian Escarpment. With a 9,613 km² watershed encompassing about 5% of the area and 3% of the population of Paraná State (Paraná 2010PARANÁ. Secretaria do Estado do Meio Ambiente e Recursos Hídricos - SEMA. 2010. Bacias hidrográficas do Paraná. SEMA, Curitiba http://www.meioambiente.pr.gov.br/arquivos/File/corh/Revista_Bacias_Hidrograficas_do_Parana.pdf (last access 16 June 2019).
http://www.meioambiente.pr.gov.br/arquiv... ) and a 240 km course, it forms the border between the municipalities of Santa Mariana and Itambaracá and is one of the main tributaries of the Capivara dam, situated in the rio Paranapanema main channel. The ribeirão Grande, ribeirão Jaboticabal, ribeirão Vermelho, and the rio Laranjinha are the main tributaries of the rio das Cinzas.

The rio Itararé basin is located in the Paraná and São Paulo states. It is born on the border between the municipalities of Bom Sucesso de Itararé (São Paulo State), and Sengés (Paraná State), and runs through much of Northeast of Paraná State until it empties into the rio Paranapanema between the municipalities of Ribeirão Claro (Paraná State) and Timburi (São Paulo State). This watershed has a total area of approximately 4,845 km², encompassing about 3% of the area and 1% of the population of the Paraná State. Its main tributaries situated in the Paraná State are the rio Jaguariaíva and rio Jaguaricatu, which flow into its left bank (Paraná 2010PARANÁ. Secretaria do Estado do Meio Ambiente e Recursos Hídricos - SEMA. 2010. Bacias hidrográficas do Paraná. SEMA, Curitiba http://www.meioambiente.pr.gov.br/arquivos/File/corh/Revista_Bacias_Hidrograficas_do_Parana.pdf (last access 16 June 2019).
http://www.meioambiente.pr.gov.br/arquiv... ).

Within the rio das Cinzas and rio Itararé basins some protected areas are guaranteed by the National System of Conservation Units of Nature in Brazil, despite of imminent political attacks that are underway (see Alves et al. 2019ALVES, G.H., SANTOS, R.S., FIGUEIREDO, B.R.S., MANETTA, G.I., MESSAGE, H.J., PAZIANOTO, L.H.R., GUIMARÃES, G.B., BENEDITO, E. & COUTO, E.V. 2019. Misguided policy may jeopardize a diverse South brazilian environmental protection area. Biota Neotrop. 19(1): e20180574. http://dx.doi.org/10.1590/1676-0611-BN-2018-0574 (last access 16 June 2019).
http://dx.doi.org/10.1590/1676-0611-BN-2... ). Between the municipalities of Cornélio Procópio and Santa Mariana is the Mata São Francisco State Park with 833 ha of Seasonal Forest, in addition to the Mina Velha State Park, in the municipality of Ibaiti, which has remnants of Araucaria forest (Paraná 2010PARANÁ. Secretaria do Estado do Meio Ambiente e Recursos Hídricos - SEMA. 2010. Bacias hidrográficas do Paraná. SEMA, Curitiba http://www.meioambiente.pr.gov.br/arquivos/File/corh/Revista_Bacias_Hidrograficas_do_Parana.pdf (last access 16 June 2019).
http://www.meioambiente.pr.gov.br/arquiv... ). The Southern portion of the basin, in the municipalities of Piraí do Sul and Jaguariaíva, composes the Environment Protection Area (EPA) of Devonian Escarpment, whose Northern portion includes the rio Itararé, between the municipalities of Jaguariaíva and Sengés. This EPA is a crucial area for biodiversity preservation because it comprises within its limits important State and National Parks, Private Reserve of Natural Heritage, and integrates important Brazilian river basins such as the rio Paranapanema system (Alves et al. 2019ALVES, G.H., SANTOS, R.S., FIGUEIREDO, B.R.S., MANETTA, G.I., MESSAGE, H.J., PAZIANOTO, L.H.R., GUIMARÃES, G.B., BENEDITO, E. & COUTO, E.V. 2019. Misguided policy may jeopardize a diverse South brazilian environmental protection area. Biota Neotrop. 19(1): e20180574. http://dx.doi.org/10.1590/1676-0611-BN-2018-0574 (last access 16 June 2019).
http://dx.doi.org/10.1590/1676-0611-BN-2... ).

Data collection

In all, 24 streams of low orders (less than three) were sampled (Table 1; Figure 1) on two collection trips: June 2017, and July 2018. Of these, seven streams are located in the rio das Cinzas basin, and 17 in the rio Itararé basin (Table 1; Figure 1). A 100-m stretch (the wadeable stream stretches have at least one sequence of riffle-run-pool meso-habitats) was defined at each site, which were georeferenced (Table 1) and sampled by electrofishing. This technique employs two electrified dip nets, producing an electric discharge between 200 and 400 V and current of 2 A, that stun the fish temporarily (Lobón-Cervia 1991LOBÓN-CERVIÁ, J. 1991. Dinámica de poblaciones de peces en rios: pesca eléctrica y métodos de capturas sucesivas en la estima de abundancias. Monografia, Consejo Superior de Investigaciones Científicas, Museo Nacional de Ciencias Naturales, Madrid.). The specimens were anesthetized in benzocaine hydrochloride and fixed in 10% formalin in the field. After a few days, the fishes were transferred to 70% ethanol in the laboratory, identified in species-level, and deposited in the Coleção Ictiológica do Núcleo de Pesquisas em Limnologia, Ictiologia e Aquicultura (NUP).

Fish species were classified based on Van der Laan et al. (2019)VAN DER LAAN, R., FRICKE, R. & ESCHMEYER, W.N. (eds). 2019. Eschmeyer’s Catalog of Fishes: Classification. http://www.calacademy.org/scientists/catalog-of-fishes-classification/ (last access 13 June 2019).
http://www.calacademy.org/scientists/cat... , species validity was based on Fricke et al. (2019)FRICKE, R., ESCHMEYER, W.N. & FONG, J.D. 2019. Eschmeyer’s Catalog of Fishes: Genera, Species, References. http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp (last access 13 June 2019).
http://researcharchive.calacademy.org/re... , and species determinations were accomplished by comparison of the specimens with original descriptions or with specimens reported from the streams of rio Paranapanema system in Paraná State (Castro et al. 2003CASTRO, R.M.C., CASATTI, L., SANTOS, H.F., FERREIRA, K.M., RIBEIRO, A.C., BENINE, R.C., DARDIS, G.Z.P., MELO, A.L.A., ABREU, T.X., BOCKMANN, F.A., CARVALHO, M., GIBRAN, F.Z. & LIMA, F.C.T. 2003. Estrutura e composição da ictiofauna de riachos do Rio Paranapanema, sudeste e sul do Brasil. Biota Neotrop. 3(1): 1-31. http://www.biotaneotropica.org.br/v3n1/pt/abstract?article+BN01703012003 (last access 13 June 2019).
http://www.biotaneotropica.org.br/v3n1/p... ; Cetra et al. 2012CETRA, M., BARRELLA, W., LANGEANI NETO, F., MARTINS, A.G., MELLO, B.J. & ALMEIDA, R.S. 2012. Fish fauna of headwater streams that cross Atlantic Forest of south São Paulo state. Check List 8(3): 421-425.; Cionek et al. 2012CIONEK, V.M., SACRAMENTO, P.A., ZANATTA, N., OTA, R.P., CORBETTA, D.F. & BENEDITO, E. 2012. Fishes from first order streams of lower Paranapanema and Ivaí rivers, upper Paraná River basin, Paraná, Brazil. Check List 8(6): 1158-1162.; Pagotto et al. 2012PAGOTTO, J.P.A., VERÍSSIMO, S., GOULART, E. & MISE, F.T. 2012. Fishes (Osteichthyes: Actinopterygii) from the Pirapó River drainage, upper Paraná River basin, Paraná state, Brazil. Check List 8(3): 463-468.; Costa et al. 2013COSTA, A.D.A., FERREIRA, D.G., SILVA, W.F., ZANATTA, A.S., SHIBATTA, O.A. & GALINDO, B.A. 2013. Fishes (Osteichthyes: Actinopterygii) from the Penacho stream, upper Paraná River basin, Paraná State, Brazil. Check List 9(3): 519-523.; Hoffmann et al. 2015HOFFMANN, A.C., NASCIMENTO, R.H.C. & SHIBATTA, O.A. 2015. Fish fauna from tributaries throughout the Tibagi River basin, upper Paraná basin, Brazil. Check List 11(6): 1815.; Cetra et al. 2016CETRA, M., MATTOX, G.M.T., FERREIRA, F.C., GUINATO, R.B., SILVA, F.V. & PEDROSA, M. 2016. Headwater stream fish fauna from the Upper Paranapanema River basin. Biota Neotrop. 16(3): e20150145. http://dx.doi.org/10.1590/1676-0611-BN-2015-0145 (last access 13 June 2019).
http://dx.doi.org/10.1590/1676-0611-BN-2... ; Claro-García et al. 2018CLARO-GARCÍA, A., ASSEGA, F.M. & SHIBATTA, O.A. 2018. Diversity and distribution of ichthyofauna in streams of the middle and lower Tibagi river basin, Paraná, Brazil. Check List 14(1): 43-53.) and from the rio Paraná floodplain (Ota et al. 2018OTA, R.R., DEPRÁ, G.C., GRAÇA, W.J. & PAVANELLI, C.S. 2018. Peixes da planície de inundação do alto rio Paraná e áreas adjacentes: revised, annotated and updated. Neotrop. Ichthyol. 16(2): e170094.). Whenever possible, the determinations of the species were checked by specialists of each taxonomic group. The species were categorized by their frequency of occurrence (hereafter, constancy), which was calculated for all sampled streams according to Dajoz (1983)DAJOZ, R. 1983. Ecologia geral. Vozes, Petrópolis.. Species with a constancy up to 25% were classified as occasional, between 25 and 50% as accessory, and higher than 50% as constant. In addition, the species were categorized by origin to distinguish native and introduced species to the upper rio Paraná system, following Langeani et al. (2007)LANGEANI, F., CASTRO, R.M.C., OYAKAWA, O.T., SHIBATTA, O.A., PAVANELLI, C.S. & CASATTI, L. 2007. Diversidade da ictiofauna do Alto Rio Paraná: composição atual e perspectivas futuras. Biota Neotrop. 7(3): 181-197 http://www.biotaneotropica.org.br/v7n3/pt/abstract?article+bn0340703 (last access 13 June 2019).
http://www.biotaneotropica.org.br/v7n3/p... , Frota et al. (2016a)FROTA, A., DEPRÁ, G.C., PETENUCCI, L.M. & GRAÇA, W.J. 2016a. Inventory of the fish fauna from Ivaí River basin, Paraná State, Brazil. Biota Neotrop. 16(3): e20150151 http://dx.doi.org/10.1590/1676-0611-BN-2015-0151 (last access 13 June 2019).
http://dx.doi.org/10.1590/1676-0611-BN-2... , Cavalli et al. (2018)CAVALLI, D., FROTA, A., LIRA, A.D., GUBIANI, E.A., MARGARIDO, V.P. & GRAÇA, W.J. 2018. Update on the ichthyofauna of the Piquiri River basin, Paraná, Brazil: a conservation priority area. Biota Neotrop. 18(2): e20170350. http://dx.doi.org/10.1590/1676-0611-BN-2017-0350 (last access 13 June 2019).
http://dx.doi.org/10.1590/1676-0611-BN-2... and Ota et al. (2018)OTA, R.R., DEPRÁ, G.C., GRAÇA, W.J. & PAVANELLI, C.S. 2018. Peixes da planície de inundação do alto rio Paraná e áreas adjacentes: revised, annotated and updated. Neotrop. Ichthyol. 16(2): e170094..

Results

A total of 1,823 individuals were collected, distributed in five orders, 12 families, and 32 species (Tables 1 and 2; Figures 2 and 3). Siluriformes (Figure 2) was the most representative order with 16 species (50.0% of all collected species) and four families, followed by Characiformes (Figure 3) with 12 species and four families. Among the families, Characidae and Loricariidae (both with seven species) presented higher richness contributing together with approximately 43.8% of all collected species, followed by Heptapteridae (five species) with approximately 15.6% of all collected species, Crenuchidae and Trichomycteridae (three species each) with approximately 8.8% of all collected species, and the other families (Table 2) that were represented by only one species.

Regarding the abundance of the sampled individuals (Figure 4), Astyanax aff. paranae Eigenmann, 1914 (447 individuals), Phalloceros harpagos Lucinda, 2008 (321 individuals), Neoplecostomus selenae Zawadzki, Pavanelli & Langeani, 2008 (208 individuals), Geophagus aff. brasiliensis (Quoy & Gaimard, 1824) (115 individuals), Cambeva davisi (Haseman, 1911) (112 individuals), Bryconamericus aff. iheringii (Boulenger, 1887) (106 individuals), Characidium schubarti Travassos, 1955 (67 individuals), Astyanax bockmanni Vari & Castro, 2007 (63 individuals), Apareiodon ibitiensis Amaral Campos, 1944 (59 individuals), and Rhamdia quelen (Quoy & Gaimard, 1824) (54 individuals) were the most abundant species, contributing with approximately 85.0% of all collected individuals. On the other hand, the other 22 species counted less than 50 individuals, representing approximately 15.0% of all collected individuals (Figure 4).

As for constancy (Table 2) most of the species were categorized as occasional (25 species). Five species (C. schubarti, R. quelen, C. davisi, G. aff. brasiliensis, and P. harpagos) were categorized as accessory, and only two species (A. aff. paranae and N. selenae) were categorized as constant. Regarding the origin of species (Table 2) none was considered as introduced to the rio Paranapanema system. It is noteworthy that the capture of Astyanax bifasciatusGaravello & Sampaio, 2010GARAVELLO, J.C. & SAMPAIO, F.A.A. 2010. Five new species of genus Astyanax Bird & Girard, 1854 from Rio Iguaçu, Paraná, Brazil (Ostariophysi, Characiformes, Characidae). Braz. J. Biol. 70(3): 847-865. represents the first record of this species from the upper rio Paraná system. Lastly, three putatively undescribed species (approximately 9.4% of all collected species) were also collected: A. aff. paranae (sensuOta et al. 2018OTA, R.R., DEPRÁ, G.C., GRAÇA, W.J. & PAVANELLI, C.S. 2018. Peixes da planície de inundação do alto rio Paraná e áreas adjacentes: revised, annotated and updated. Neotrop. Ichthyol. 16(2): e170094.), Characidium sp., and Eigenmannia sp.

Discussion

With the predominance of Siluriformes and Characiformes, the taxonomic composition of the 24 sampled streams follows the pattern described for the Neotropical freshwater fish fauna (Castro 1999CASTRO, R.M.C. 1999. Evolução da ictiofauna de riachos sul americanos: padrões gerais e possíveis processos casuais. In Ecologia de peixes de riachos: estado atual e perspectivas (E.P. Caramaschi, R. Mazzoni & P.R. Peres-Neto, eds). UFRJ, Rio de Janeiro, p.139-155.; Lowe-McConnell 1999LOWE-MCCONNELL, R.H. 1999. Estudos ecológicos de comunidades de peixes tropicais. EDUSP, São Paulo.; Albert et al. 2011ALBERT, J., PETRY, P. & REIS, R.E. 2011. Major biogeographic and phylogenetic patterns. In Historical biogeography of Neotropical freshwater fishes (J.S. Albert & R.E. Reis, eds). University of California Press, Berkeley, p.21-56.). The number of species recorded in Characidae, Loricariidae, and Heptapteridae also follows the pattern of the most species-rich families in the upper rio Paraná system (Langeani et al. 2007LANGEANI, F., CASTRO, R.M.C., OYAKAWA, O.T., SHIBATTA, O.A., PAVANELLI, C.S. & CASATTI, L. 2007. Diversidade da ictiofauna do Alto Rio Paraná: composição atual e perspectivas futuras. Biota Neotrop. 7(3): 181-197 http://www.biotaneotropica.org.br/v7n3/pt/abstract?article+bn0340703 (last access 13 June 2019).
http://www.biotaneotropica.org.br/v7n3/p... ). Similarly, several studies have indicated the prevalence of these families in other streams of this drainage system (e.g.,Hoffmann et al. 2015HOFFMANN, A.C., NASCIMENTO, R.H.C. & SHIBATTA, O.A. 2015. Fish fauna from tributaries throughout the Tibagi River basin, upper Paraná basin, Brazil. Check List 11(6): 1815.; Bifi et al. 2017BIFI, A.G., DIAS, A.C. & FROTA, A. 2017. Fish species (Osteichthyes: Actinopterygii) from two tributaries of the Rio do Peixe basin, Tupã municipality, São Paulo state, Brazil. Check List 13(2): 2063.; Batista-Silva et al. 2018BATISTA-SILVA, V.F., FROTA, A., KASHIWAQUI, E.A.L., ABELHA, M.C.F., BAILLY, D., GUBIANI, E.A. & GRAÇA, W.J. 2018. Ichthyofauna from three streams of the lower Iguatemi River in the upper Paraná river basin, Brazil. Check List 14(2): 363-378.; Claro-García et al. 2018CLARO-GARCÍA, A., ASSEGA, F.M. & SHIBATTA, O.A. 2018. Diversity and distribution of ichthyofauna in streams of the middle and lower Tibagi river basin, Paraná, Brazil. Check List 14(1): 43-53.; Azevedo-Santos et al. 2019bAZEVEDO-SANTOS, V.M., BRITSKI, H.A., OLIVEIRA, C. & BENINE, R.C. 2019b. Ichthyofauna of streams of the Rio Sapucaí basin, upper Rio Paraná system, Minas Gerais, Brazil. Biota Neotrop. 19(1): e20180617 http://dx.doi.org/10.1590/1676-0611-BN-2018-0617 (last access 18 June 2019).
http://dx.doi.org/10.1590/1676-0611-BN-2... ), especially for streams of the rio Paranapanema system, such as the rio das Cinzas (Costa et al. 2013COSTA, A.D.A., FERREIRA, D.G., SILVA, W.F., ZANATTA, A.S., SHIBATTA, O.A. & GALINDO, B.A. 2013. Fishes (Osteichthyes: Actinopterygii) from the Penacho stream, upper Paraná River basin, Paraná State, Brazil. Check List 9(3): 519-523.) and the rio Itararé (Cetra et al. 2016CETRA, M., MATTOX, G.M.T., FERREIRA, F.C., GUINATO, R.B., SILVA, F.V. & PEDROSA, M. 2016. Headwater stream fish fauna from the Upper Paranapanema River basin. Biota Neotrop. 16(3): e20150145. http://dx.doi.org/10.1590/1676-0611-BN-2015-0145 (last access 13 June 2019).
http://dx.doi.org/10.1590/1676-0611-BN-2... ) basins.

Our results showed that occasional species dominated the sampled streams. This may be related to the influence of the altitudinal gradient on the species distribution since the species richness tends to decrease with the altitude increase (Fu et al. 2004FU, C., WU, J., WANG, X., LEI, G. & CHEN, J. 2004. Patterns of diversity, altitudinal range and body size among freshwater fishes in the Yangtze River Basin, China. Global Ecol. Biogeogr. 13(6): 43-552.; Jaramillo-Villa et al. 2010JARAMILLO-VILLA, U., MALDONADO-OCAMPO, J.A. & ESCOBAR, F. 2010. Altitudinal variation in fish assemblage diversity in streams of the central Andes of Colombia. J. Fish Biol. 76(10): 2401-2417.; Askeyev et al. 2017ASKEYEV, A., ASKEYEV, O., YANYBAEV, N., ASKEYEV, I., MONAKHOV, S., MARIĆ, S. & HULSMAN, K. 2017. River fish assemblages along an elevation gradient in the eastern extremity of Europe. Environ. Biol. Fish. 100(5): 585-596.; Frota et al. 2019FROTA, A., MESSAGE, H.J., OLIVEIRA, R.C., BENEDITO, E. & GRAÇA, W.J. 2019. Ichthyofauna of headwater streams from the rio Ribeira de Iguape basin, at the boundaries of the Ponta Grossa Arch, Paraná, Brazil. Biota Neotrop. 19(1): e20180666 http://dx.doi.org/10.1590/1676-0611-BN-2018-0666 (last access 17 June 2019).
http://dx.doi.org/10.1590/1676-0611-BN-2... ). As verified in other studies relating fish species richness with the altitudinal gradient (e.g.,Askeyev et al. 2017ASKEYEV, A., ASKEYEV, O., YANYBAEV, N., ASKEYEV, I., MONAKHOV, S., MARIĆ, S. & HULSMAN, K. 2017. River fish assemblages along an elevation gradient in the eastern extremity of Europe. Environ. Biol. Fish. 100(5): 585-596.), above 650 m of altitude few species were repeated in the streams sampled (Table 1). These species were classified as constant or accessory, in addition to being among the most abundant species such as A. aff. paranae, N. selenae, C. davisi, R. quelen, and C. schubarti. In fact, members of Heptapteridae, Loricariidae, and Trichomycteridae have morphological structures adapted to fixation in underwater structures, such as fortified pectoral fin rays, subterminal sucker mouths and/or spines in the operculum. These adaptations can guarantee the conquest of higher lands, such as headwater streams (Oyakawa et al. 2006OYAKAWA, O.T., AKAMA, A., MAUTARI, K.C. & NOLASCO, J.C. 2006. Peixes de riachos da Mata Atlântica. Neotrópica, São Paulo.). The great abundance and constancy of A. aff. paranae may be related to a short-distance migratory character (<100 km) already evidenced for some species of this genus (Agostinho et al. 2004AGOSTINHO, A.A., GOMES, L.C., VERÍSSIMO, S. & OKADA, E.K. 2004. Flood regime, dam regulation and fish in the Upper Paraná River: effects on assemblage attributes, reproduction and recruitment. Rev. Fish Biol. Fisher. 14(1): 11-19.).

Costa et al. (2013)COSTA, A.D.A., FERREIRA, D.G., SILVA, W.F., ZANATTA, A.S., SHIBATTA, O.A. & GALINDO, B.A. 2013. Fishes (Osteichthyes: Actinopterygii) from the Penacho stream, upper Paraná River basin, Paraná State, Brazil. Check List 9(3): 519-523. found a few introduced species already inhabiting the waters of the rio das Cinzas basin, such as Cyprinus carpio Linnaeus, 1768, Oreochromis niloticus (Linnaeus, 1758), and Poecilia reticulata Peters, 1859. In contrast, the absence of such species from the headwater streams sampled herein suggests a greater integrity of that fish community, since biological invasions are among the main threats to freshwater environments (Daga et al. 2015DAGA, V.S., SKÓRA, F., PADIAL, A.A., ABILHOA, V., GUBIANI, É.A. & VITULE, J.R. S. 2015. Homogenization dynamics of the fish assemblages in Neotropical reservoirs: comparing the roles of introduced species and their vectors. Hydrobiologia 746(1): 327-347.; Ruaro et al. 2018RUARO, R., MORMUL, R.P., GUBIANI, É.A., PIANA, P.A., CUNICO, A.M. & GRAÇA, W.J. 2018. Non-native fish species are related to the loss of ecological integrity in Neotropical streams: a multimetric approach. Hydrobiologia 817(1): 413-430.), and especially detrimental for the fish fauna of Neotropical streams (Winemiller et al. 2008WINEMILLER, K.O., AGOSTINHO, A.A. & CARAMASCHI, E.P. 2008. Fish ecology in tropical streams. In Tropical stream ecology (D. Dudgeon, ed). Academic Press, San Diego, p.107-146.; Ruaro et al. 2018RUARO, R., MORMUL, R.P., GUBIANI, É.A., PIANA, P.A., CUNICO, A.M. & GRAÇA, W.J. 2018. Non-native fish species are related to the loss of ecological integrity in Neotropical streams: a multimetric approach. Hydrobiologia 817(1): 413-430.). The Devonian Escarpment EPA is directly linked to the maintenance of this integrity, especially in the case of the rio Itararé basin, and conservation efforts are increasingly necessary to at least maintain the original area of this natural reserve (Alves et al. 2019ALVES, G.H., SANTOS, R.S., FIGUEIREDO, B.R.S., MANETTA, G.I., MESSAGE, H.J., PAZIANOTO, L.H.R., GUIMARÃES, G.B., BENEDITO, E. & COUTO, E.V. 2019. Misguided policy may jeopardize a diverse South brazilian environmental protection area. Biota Neotrop. 19(1): e20180574. http://dx.doi.org/10.1590/1676-0611-BN-2018-0574 (last access 16 June 2019).
http://dx.doi.org/10.1590/1676-0611-BN-2... ).

Astyanax bifasciatus was thought to be endemic to the rio Iguaçu basin (Garavello & Sampaio 2010GARAVELLO, J.C. & SAMPAIO, F.A.A. 2010. Five new species of genus Astyanax Bird & Girard, 1854 from Rio Iguaçu, Paraná, Brazil (Ostariophysi, Characiformes, Characidae). Braz. J. Biol. 70(3): 847-865.; Baumgartner et al. 2012BAUMGARTNER, G., PAVANELLI, C.S., BAUMGARTNER, D., BIFI, A.G., DEBONA, T. & FRANA, V.A. 2012. Peixes do Baixo Rio Iguaçu. Eduem, Maringá.), a drainage system of the lower rio Paraná that encompasses a large area of southern Paraná State and northern Santa Catarina State. In a recent inventory (Frota et al. 2019FROTA, A., MESSAGE, H.J., OLIVEIRA, R.C., BENEDITO, E. & GRAÇA, W.J. 2019. Ichthyofauna of headwater streams from the rio Ribeira de Iguape basin, at the boundaries of the Ponta Grossa Arch, Paraná, Brazil. Biota Neotrop. 19(1): e20180666 http://dx.doi.org/10.1590/1676-0611-BN-2018-0666 (last access 17 June 2019).
http://dx.doi.org/10.1590/1676-0611-BN-2... ), this species had its geographical distribution extended to the rio Ribeira de Iguape basin, a coastal drainage bordered to the south by the rio Iguaçu basin and to the north by the rio Itararé basin. Another example of the interesting biogeographic (Ribeiro 2006RIBEIRO, A.C. 2006. Tectonic history and the biogeography of the freshwater fishes from the coastal drainages of eastern Brazil: an example of faunal evolution associated with a divergent continental margin. Neotrop. Ichthyol. 4(2): 225-246.; Frota et al. 2016aFROTA, A., DEPRÁ, G.C., PETENUCCI, L.M. & GRAÇA, W.J. 2016a. Inventory of the fish fauna from Ivaí River basin, Paraná State, Brazil. Biota Neotrop. 16(3): e20150151 http://dx.doi.org/10.1590/1676-0611-BN-2015-0151 (last access 13 June 2019).
http://dx.doi.org/10.1590/1676-0611-BN-2... ) and phylogeographic (Morais-Silva et al. 2018MORAIS-SILVA, J.P., OLIVEIRA, A.V., FABRIN, T.M.C., DIAMANTE, N.A., PRIOLI, S.M.A.P., FROTA, A., GRAÇA, W.J. & PRIOLI, A.J. 2018. Geomorphology influencing the diversification of fish in small-order rivers of neighboring basins. Zebrafish 15(4): 389-397.) patterns at the boundaries of the Ponta Grossa Arch. In fact, this arch is the most prominent geological structure in the headwater regions of the Paranapanema, Iguaçu and Ribeira de Iguape river basins (Ribeiro 2006RIBEIRO, A.C. 2006. Tectonic history and the biogeography of the freshwater fishes from the coastal drainages of eastern Brazil: an example of faunal evolution associated with a divergent continental margin. Neotrop. Ichthyol. 4(2): 225-246.).

Mainly due to their recent tectonic activities (Franco-Magalhães et al. 2010FRANCO-MAGALHAES, A.O.B., HACKSPACHER, P.C., GLASMACHER, U.A. & SAAD, A.R. 2010. Rift to post-rift evolution of a “passive” continental margin: the Ponta Grossa Arch, SE Brazil. Int. J. Earth Sci. 99(7): 1599-1613.), the Ponta Grossa Arch is thought to be responsible for a particularly accelerated river dynamism between these adjacent drainage systems, enabling the faunal exchange between them (Ribeiro 2006RIBEIRO, A.C. 2006. Tectonic history and the biogeography of the freshwater fishes from the coastal drainages of eastern Brazil: an example of faunal evolution associated with a divergent continental margin. Neotrop. Ichthyol. 4(2): 225-246.). This same scenario can be found for other fish species. For example, adjacent tributaries of the Iguaçu, Paranapanema and Ribeira de Iguape river basins share the presence of C. davisi (see Ingenito et al. 2004INGENITO, L.F.S., DUBOC, L.F. & ABILHOA, V. 2004. Contribuição ao conhecimento da ictiofauna da bacia do alto rio Iguaçu, Paraná, Brasil. Arquivos de Ciências Veterinárias e Zoológicas da UNIPAR 7(1): 23-36.; Ribeiro 2006RIBEIRO, A.C. 2006. Tectonic history and the biogeography of the freshwater fishes from the coastal drainages of eastern Brazil: an example of faunal evolution associated with a divergent continental margin. Neotrop. Ichthyol. 4(2): 225-246.; Morais-Silva et al. 2018MORAIS-SILVA, J.P., OLIVEIRA, A.V., FABRIN, T.M.C., DIAMANTE, N.A., PRIOLI, S.M.A.P., FROTA, A., GRAÇA, W.J. & PRIOLI, A.J. 2018. Geomorphology influencing the diversification of fish in small-order rivers of neighboring basins. Zebrafish 15(4): 389-397.), Hyphessobrycon griemi Hoedeman, 1957, H. bifasciatus Ellis, 1911, H. reticulatus Ellis, 1911 (see Ingenito et al. 2004INGENITO, L.F.S., DUBOC, L.F. & ABILHOA, V. 2004. Contribuição ao conhecimento da ictiofauna da bacia do alto rio Iguaçu, Paraná, Brasil. Arquivos de Ciências Veterinárias e Zoológicas da UNIPAR 7(1): 23-36.; Ribeiro 2006RIBEIRO, A.C. 2006. Tectonic history and the biogeography of the freshwater fishes from the coastal drainages of eastern Brazil: an example of faunal evolution associated with a divergent continental margin. Neotrop. Ichthyol. 4(2): 225-246.), Mimagoniates microlepis (Steindachner, 1877) (see Ingenito et al. 2004INGENITO, L.F.S., DUBOC, L.F. & ABILHOA, V. 2004. Contribuição ao conhecimento da ictiofauna da bacia do alto rio Iguaçu, Paraná, Brasil. Arquivos de Ciências Veterinárias e Zoológicas da UNIPAR 7(1): 23-36.; Ribeiro 2006RIBEIRO, A.C. 2006. Tectonic history and the biogeography of the freshwater fishes from the coastal drainages of eastern Brazil: an example of faunal evolution associated with a divergent continental margin. Neotrop. Ichthyol. 4(2): 225-246.; Menezes et al. 2008MENEZES, N.A., RIBEIRO, A.C. WEITZMAN, S. & TORRES, R.A. 2008. Biogeography of Glandulocaudinae (Teleosei: Characiformes: Characidae) revisited: phylogenetic patterns, historical geology and genetic connectivity. Zootaxa 1726(1): 33-48.; Hoffmann et al. 2015HOFFMANN, A.C., NASCIMENTO, R.H.C. & SHIBATTA, O.A. 2015. Fish fauna from tributaries throughout the Tibagi River basin, upper Paraná basin, Brazil. Check List 11(6): 1815.), and Rhamdiopsis moreirai Haseman, 1911 (see Ferraris 2007FERRARIS, C.J. 2007. Checklist of catfishes, recent and fossil (Osteichthyes: Siluriformes), and catalogue of siluriform primary types. Zootaxa 1418(1): 1-628.; Hoffmann et al. 2015HOFFMANN, A.C., NASCIMENTO, R.H.C. & SHIBATTA, O.A. 2015. Fish fauna from tributaries throughout the Tibagi River basin, upper Paraná basin, Brazil. Check List 11(6): 1815.). The slopes of the upper rio Paraná system and the rio Ribeira de Iguape basin share the presence of Hypostomus ancistroides (Ihering, 1911) (Oyakawa et al. 2005OYAKAWA, O.T., AKAMA, A. & ZANATA, A.M. 2005. Review of the genus Hypostomus Lacépède, 1803 from rio Ribeira de Iguape basin, with description of a new species (Pisces, Siluriformes, Loricariidae). Zootaxa 921(1): 1-27.). The Iguaçu and Ribeira de Iguape river basins share the presence of Characidium travassosiMelo, Buckup & Oyakawa, 2016MELO, M.R.S., BUCKUP, P.A. & OYAKAWA, O.T. 2016. A new species of Characidium Reinhardt, 1867 (Characiformes: Crenuchidae) endemic to the Atlantic Forest in Paraná State, Southern Brazil. Neotrop. Ichthyol. 14(3): e160014. (Frota et al. 2016bFROTA, A., GONÇALVES, E.V.R., DEPRÁ, G.C. & GRAÇA, W.J. 2016b. Inventory of the ichthyofauna from the Jordão and Areia river basins (Iguaçu drainage, Brazil) reveals greater sharing of species than thought. Check List 12(6): 1995.; Melo et al. 2016MELO, M.R.S., BUCKUP, P.A. & OYAKAWA, O.T. 2016. A new species of Characidium Reinhardt, 1867 (Characiformes: Crenuchidae) endemic to the Atlantic Forest in Paraná State, Southern Brazil. Neotrop. Ichthyol. 14(3): e160014.). Now, the geographic distribution of A. bifasciatus is added to the mentioned examples since this species also occurs in all drainage systems mentioned.

Nine of the 32 fish species records presented herein are new for the Itararé and Cinzas river basins. Those are Bryconamericus turiubaLangeani, Lucena, Pedrini & Tarelho-Pereira, 2005LANGEANI, F., CASTRO, R.M.C., OYAKAWA, O.T., SHIBATTA, O.A., PAVANELLI, C.S. & CASATTI, L. 2007. Diversidade da ictiofauna do Alto Rio Paraná: composição atual e perspectivas futuras. Biota Neotrop. 7(3): 181-197 http://www.biotaneotropica.org.br/v7n3/pt/abstract?article+bn0340703 (last access 13 June 2019).
http://www.biotaneotropica.org.br/v7n3/p... , Corydoras ehrhardti Steindachner, 1910, Hypostomus iheringii (Regan, 1908), Hypostomus tietensis (Ihering, 1905), Cambeva iheringi (Eigenmann, 1917), Gymnotus inaequilabiatus Valenciennes, 1839, and the three putatively new species A. aff. paranae (sensuOta et al. 2018OTA, R.R., DEPRÁ, G.C., GRAÇA, W.J. & PAVANELLI, C.S. 2018. Peixes da planície de inundação do alto rio Paraná e áreas adjacentes: revised, annotated and updated. Neotrop. Ichthyol. 16(2): e170094.), Characidium sp., and Eigenmannia sp. (compare with Costa et al. 2013COSTA, A.D.A., FERREIRA, D.G., SILVA, W.F., ZANATTA, A.S., SHIBATTA, O.A. & GALINDO, B.A. 2013. Fishes (Osteichthyes: Actinopterygii) from the Penacho stream, upper Paraná River basin, Paraná State, Brazil. Check List 9(3): 519-523.; Cetra et al. 2016CETRA, M., MATTOX, G.M.T., FERREIRA, F.C., GUINATO, R.B., SILVA, F.V. & PEDROSA, M. 2016. Headwater stream fish fauna from the Upper Paranapanema River basin. Biota Neotrop. 16(3): e20150145. http://dx.doi.org/10.1590/1676-0611-BN-2015-0145 (last access 13 June 2019).
http://dx.doi.org/10.1590/1676-0611-BN-2... ). Except for the latter two species, which are being reported for the first time herein, all of these species have a wide distribution range within the upper rio Paraná drainage. On the other hand, Costa et al. (2013)COSTA, A.D.A., FERREIRA, D.G., SILVA, W.F., ZANATTA, A.S., SHIBATTA, O.A. & GALINDO, B.A. 2013. Fishes (Osteichthyes: Actinopterygii) from the Penacho stream, upper Paraná River basin, Paraná State, Brazil. Check List 9(3): 519-523. reported 11 species not collected by us (introduced species excluded): Piabina argentea Reinhardt, 1867, Apareiodon affinis (Steindachner, 1879), Gymnotus sylviusAlbert & Fernandes-Matioli, 1999ALBERT, J., PETRY, P. & REIS, R.E. 2011. Major biogeographic and phylogenetic patterns. In Historical biogeography of Neotropical freshwater fishes (J.S. Albert & R.E. Reis, eds). University of California Press, Berkeley, p.21-56., Gymnotus sp., Imparfinis schubarti (Gomes, 1956), Hypostomus albopunctatus (Regan, 1908), H. nigromaculatus (Schubart, 1964), H. paulinus (Ihering, 1905), Rineloricaria pentamaculata Langeani & Araujo, 1994, Crenicichla jaguarensis Haseman, 1911, and Synbranchus marmoratus Bloch, 1795. Cetra et al. (2016)CETRA, M., MATTOX, G.M.T., FERREIRA, F.C., GUINATO, R.B., SILVA, F.V. & PEDROSA, M. 2016. Headwater stream fish fauna from the Upper Paranapanema River basin. Biota Neotrop. 16(3): e20150145. http://dx.doi.org/10.1590/1676-0611-BN-2015-0145 (last access 13 June 2019).
http://dx.doi.org/10.1590/1676-0611-BN-2... recorded 19 (introduced species excluded): Astyanax biotae Castro & Vari, 2004, A. scabripinnis (Jenyns, 1842), Hyphessobrycon anisitsi (Eigenmann, 1907), Oligosarcus paranensis Menezes & Géry, 1983, Piabina argentea, Characidium gomesi Travassos, 1956, Parodon nasus Kner, 1859, Gymnotus sylvius, Eigenmannia virescens (Valenciennes, 1836), Pimelodella avanhandavae Eigenmann, 1917, Hypostomus nigromaculatus, Hypostomus sp. 1, Hypostomus sp. 2, Otothyropsis sp., Rineloricaria pentamaculata, Trichomycterus sp. 1, Trichomycterus sp. 2, Phalloceros reisi Lucinda, 2008, and Synbranchus marmoratus.

The identification of A. biotae, H. anisitsi, P. argentea, O. paranensis, C. gomesi, A. affinis, H. albopunctatus, H. nigromaculatus, R. pentamaculata, P. avanhandavae, P. reisi, C. jaguarensis, and S. marmoratus is pretty straightforward, meaning that at least 13 species, other than those presented by us, could be added to a complete list of the native stream fish fauna from the upper rio Paranapanema system. However, a careful analysis of the remaining species is necessary in order to verify whether they are in fact distinct from the species listed herein. Gymnotus species are hardly distinguishable from one another, especially at small sizes. Imparfinis schubarti has been considered a junior synonym of I. mirini Haseman, 1911, which suggests that the usual diagnostic characters may not be sufficient to distinguish those species, in the case that they are in fact two species. Hypostomus paulinus represents a species complex and is easily confused with H. tietensis (Zawadzki, personal communication).

Some species considered to be new by other authors (Costa et al. 2013COSTA, A.D.A., FERREIRA, D.G., SILVA, W.F., ZANATTA, A.S., SHIBATTA, O.A. & GALINDO, B.A. 2013. Fishes (Osteichthyes: Actinopterygii) from the Penacho stream, upper Paraná River basin, Paraná State, Brazil. Check List 9(3): 519-523.; Cetra et al. 2016CETRA, M., MATTOX, G.M.T., FERREIRA, F.C., GUINATO, R.B., SILVA, F.V. & PEDROSA, M. 2016. Headwater stream fish fauna from the Upper Paranapanema River basin. Biota Neotrop. 16(3): e20150145. http://dx.doi.org/10.1590/1676-0611-BN-2015-0145 (last access 13 June 2019).
http://dx.doi.org/10.1590/1676-0611-BN-2... ) may be variants of other, described species, which cannot be verified presently without the examination of the specimens. For example, regarding the two supposedly new species of Trichomycterus (= Cambeva), we found among our specimens a wide color-pattern variation, which nonetheless fall within the range of variation observed by Nascimento et al. (2017)NASCIMENTO, R.H.C., FRANTINE-SILVA, W., SOUZA-SHIBATTA, L., SOFIA, S.H., FERRER, J. & SHIBATTA, O.A. 2017. Intrapopulational variation in color pattern of Trichomycterus davisi (Haseman, 1911) (Siluriformes: Trichomycteridae) corroborated by morphometrics and molecular analysis. Zootaxa 4290(3): 503-518. for C. davisi and by da Silva et al. (2010)DA SILVA, C.C.F., MATTA, L.S.F., HILSDORF, A.W.S., LANGEANI, F. & MARCENIUK, A.P. 2010. Color pattern variation in Trichomycterus iheringi (Eigenmann, 1917) (Siluriformes: Tricomycteridae) from Rio Itatinga and Rio Claro, São Paulo, Brazil. Neotrop. Ichthyol. 8(1): 49-56. for C. iheringi. Thus, we cautiously hypothesize Trichomycterus sp. 1 and Trichomycterus sp. 2 of Cetra et al. (2016)CETRA, M., MATTOX, G.M.T., FERREIRA, F.C., GUINATO, R.B., SILVA, F.V. & PEDROSA, M. 2016. Headwater stream fish fauna from the Upper Paranapanema River basin. Biota Neotrop. 16(3): e20150145. http://dx.doi.org/10.1590/1676-0611-BN-2015-0145 (last access 13 June 2019).
http://dx.doi.org/10.1590/1676-0611-BN-2... to represent variants of these species.

In any case, in addition to the results of Cetra et al. (2016)CETRA, M., MATTOX, G.M.T., FERREIRA, F.C., GUINATO, R.B., SILVA, F.V. & PEDROSA, M. 2016. Headwater stream fish fauna from the Upper Paranapanema River basin. Biota Neotrop. 16(3): e20150145. http://dx.doi.org/10.1590/1676-0611-BN-2015-0145 (last access 13 June 2019).
http://dx.doi.org/10.1590/1676-0611-BN-2... , our results point to the gap between existing evolutionary knowledge and current biological knowledge, which characterizes a large biodiversity shortfall (Hortal et al. 2015HORTAL, J., BELLO, F., DINIZ-FILHO, J.A.F., LEWINSOHN, T.M., LOBO, J.M. & LADLE, R.J. 2015. Seven shortfalls that beset large-scale knowledge of biodiversity. Annu. Rev. Ecol. Evol. S. 46: 523-549.), especially for the stream fish fauna from the rio Paranapanema system. Future taxonomic studies with morphological and molecular methodologies can be conclusive on these issues. Here we have improved knowledge of the fish fauna of the rio das Cinzas and rio Itararé basins, upper rio Paranapanema system. However, we recognize that this inventory likely represents a small fraction of what remains to be sampled within these basins. Once again, sampling at headwater regions of high altitudes revealed possible unknown species to science, interesting biogeographic patterns and alarming biodiversity shortfalls, as well as inherent concerns about anthropogenic actions in Brazilian freshwater environments.

Acknowledgements

We are deeply grateful to Francisco Alves Teixeira (UEM), Valmir Alves Teixeira (UEM), and Wladimir Marques Domingues (UEM) for the aid in the field trip; to Cláudio Zawadzki (UEM), Carlos Oliveira (UEM), and Fernando Carvalho (UFMS) for help in fish identification; to Marli Campos (Nupélia) by cataloging the vouchers. We thanks to Committee of Ethics for the Use of Animals in Experimentation, State University of Maringá (UEM) by the approval of samplings (process number 5680160117), and to Brazilian Institute of Environment and Renewable Natural Resources (IBAMA, in Portuguese) by the permanent license to collect zoological material (number 14028-1) conceived to WJG. AF and WJG received grants from Fundação Araucária. AF, MJMG, and RRO has been supported by scholarship from Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq). WJG received personal grants from CNPq (305200/2018-6).

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