Freshwater fishes of the Río de la Plata: current assemblage structure

Tomás Maiztegui^1,2 , Ariel Hernán Paracampo², Jorge Liotta³, Eva Cabanellas², Carlos Bonetto² and Darío César Colautti²

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Abstract

Few studies have addressed the composition of fish assemblages of the freshwater Río de la Plata (RdlP) and have only been limited to species lists gathered over the last two centuries. As such inventories have never been reviewed or validated by fish sampling, the richness and structure of RdlP fish assemblage are poorly known. Hence, we conducted an exhaustive literature review and a fieldwork in six coastal points of Argentina to update the species composition and determine the hierarchical structure of the fish assemblage. From the 206 species registered in the literature, 48 were not confirmed, 13 were absent, five were taken as synonymized species, 29 were supported by literature and 107 were confirmed; one was an established exotic species, and three were a non-established exotic species. The findings reported here suggest that the fish assemblage currently comprises 141 species, including four new records. Analysis of fieldwork data in number and weight of fish captured resulted in an assemblage hierarchical structure of five dominant, 22 frequent, and 45 rare species; 16 dominant, 11 frequent, and 45 rare taxa, respectively. These results could be used as baseline to monitor, manage, and preserve neotropical fish species in their southern distribution boundary.

Keywords: Biodiversity, Conservation, Ichthyofauna, Neotropics, Richness.

El conocimiento de los ensambles de peces del sector dulceacuícola del Río de la Plata (RdlP) es escaso y limitado a listas de especies de compilaciones realizadas en los últimos dos siglos. Como esos inventarios nunca han sido revisados o validados mediante muestreos de peces, el conocimiento respecto de la riqueza y la estructura de los ensambles del RdlP resulta deficiente. Para ordenar y mejorar la información acerca de la riqueza y estructura de los ensambles de peces del área se realizó una revisión bibliográfica y un muestreo de campo para actualizar la composición de especies y determinar la estructura jerárquica del ensamble de peces. De las 206 especies colectadas de acuerdo a la bibliografía 48 se categorizaron como no reconfirmadas, 13 como ausentes, cinco como especies sinonimizadas, 29 como soportadas por la literatura, 107 como confirmadas, una exótica establecida y tres exóticas no establecidas. Los resultados sugieren que el ensamble de peces actualmente está compuesto por 141 especies, incluyendo cuatro nuevos registros. El análisis de los datos en número y en peso de los peces colectados mostró 5 especies dominantes, 22 frecuentes y 45 raras; y 16 dominantes, 11 frecuentes y 45 raras, respectivamente. Los resultados de este trabajo pueden ser usados como línea de base para el monitoreo, manejo y conservación de las especies de peces neotropicales en su límite de distribución sur.

Palabras clave: Biodiversidad, Conservación, Ictiofauna, Neotrópico, Riqueza.

Introduction

Freshwater ecosystems such as rivers, ponds, lakes, and streams are essential for human survival. Yet, they are among the most degraded habitats of the world (Dudgeon et al., 2006), making freshwater species the most threatened on Earth. Alarmingly, loss of freshwater biodiversity is mounting fast (Kumar, 2000; Sala et al., 2000) due to anthropogenic activity such as habitat alteration, water pollution, introduction of exotic species, and overexploitation of natural resources (Dudgeon et al., 2006; Arthington et al., 2010; Liermann et al., 2012; Pelicice et al., 2021).

Vitule et al. (2016) stated that loss of freshwater fish biodiversity cannot be measured simply by the number of threatened species or of species that became extinct, arguing that a decrease in freshwater fish populations, threatened or not, may drastically alter the ecosystem functioning. Moreover, research into the biological diversity of freshwater ecosystems is yet limited (Stiassny, 2002). As inventories allow assessing the presence of species or infer their absence within an area (Morrison et al., 2008), greater insights are required to develop the basis for measuring the biodiversity of species. Green et al. (2009) claimed that inventories made from compendiums of information previously collected from an area show all the inherent limitations associated with non-quantitative methodologies. From studies of the vulnerability of freshwater fish diversity (Tedesco et al., 2017) derived the need to generate updated quali-quantitative ichthyological research used as a basis to evaluate species conservation and extinction risk, particularly in South America (Baigún et al., 2012). For instance, Jaureguizar et al. (2015) on the basis of the abundance and relative frequency of each species, typified the components of fish assemblage on the Argentinean estuarine-marine coast. Such approach had led to a thorough understanding to improve or develop integral conservation policies, strategic management decisions, and/or accurate environmental impact assessments.

The Río de la Plata (RdlP) is the collector of the La Plata River basin, the second largest watershed in South America, draining a surface of almost three million km² (Pasquini, Depetris, 2007) and representing the third most diverse basin of the continent with more than 900 fish taxa (Reis et al., 2016). Although ichthyological studies have been conducted on freshwater RdlP and neighboring streams scattered on the space and time scales, most are related to governmental reports centered on the relevant species for local fisheries (CARP-INIDEP-INAPE, 1990; CARU, 1992; CARU-CARP, 2012; 2016); hence, fish assemblages are still being poorly understood (Llompart et al., 2012). According to Baigún et al. (2012), 26% of the fish found in the lower La Plata River basin are classified as Data Deficient (IUCN), due to lack of information on their population and distribution. RdlP is the meridional edge of the Lower Paraná Ecoregion (Abell et al., 2008), which constitutes the distribution boundary for the most representatives of freshwater Neotropical fish (Ringuelet, 1975).

Bibliographic compilations describing the composition of the freshwater fish of RdlP were provided by Nion (1998), López et al. (2003), and Volpedo et al. (2010), recording approximately 170 species in the area. However, one of the main issues arising from these inventories is that compilers used the exhaustive work of Ringuelet et al. (1967) as a reference. This pioneer study represents the first Argentinean fish species list based on specimens collected in field samplings and on bibliographic compilations. Over the last twenty years, extensive reviews have been made for the freshwater ichthyofauna of Argentina and Uruguay, and several species have been synonymized or disregarded as part of the fish fauna of those countries (Litz, Koerber, 2014; Koerber et al., 2016a,b; 2020; Mirande, Koerber, 2020). However, the validity of fish records, the actual number of species, their frequency of occurrence, and abundance in the area have not been reviewed nor have they been contrasted with current surveys. This indicates a need to review and updated lists of current freshwater fish living in the RdlP, based on a quantitative approach.

In addition, freshwater RdlP is subject to environmental pollution due to the large urban and industrial centers located on its coast (AA-AGOSBA-OSN-SHN, 1992; FREPLATA, 2004), threatening the fish biodiversity inhabiting this ecosystem. Thus, the revision and characterization of the composition, distribution and community of fish species present in this area and their hierarchical structure within the assemblage will provide the foundation for assessing the conservation status of species, assemblages, and fisheries.

Material and methods

Study area. The RdlP system is located on the western South American Atlantic coast (35°S 58°W) and can be defined as a funnel coastal plain tidal river with a semi-closed shelf at the mouth (Baigún et al., 2016). This extensive environment is divided into two main areas by the submersed shoal known as Barra del Indio that crosses the river transversally (Fig. 1): the riverine zone (< 5 Practical Salinity Units: PSU) and the estuarine zone (> 5 PSU) (Guerrero et al., 1997; Baigún et al., 2016). The riverine zone is a shallow area not exceeding 5 m deep, characterized by a silty sand bottom that gradually turns to silt into the estuarine zone (Wells, Daborn, 1998). The main tributaries of the riverine RdlP are the Paraná and Uruguay rivers, providing 97% of the freshwater inflow (Guerrero et al., 1997), together with several streams draining its coasts. Despite its main fluvial regime, this area also exhibits tidal influence (Balay, 1961), allowing a dynamic interaction between freshwater-brackish waters that can be highly affected by the wind action (Simionato et al., 2007).

From an ichthyogeographical approach, Ringuelet (1975) argued that La Plata city (Fig. 1), and its surroundings, should be considered as the boundary where most freshwater fish fauna of La Plata River basin lives, due to a pauperization of taxa related to the saline gradient towards the estuary. However, López et al. (2002, 2008) extended the area of freshwater fish downstream, closer to the estuarine zone. For the present work, the study area comprised the Río de la Plata stretch from the mouth of Paraná (downstream the “Delta”) and Uruguay rivers (downstream the locality of “Punta Gorda”) to the imaginary line from La Balandra (Argentina) to Juan Lacaze (Uruguay) in opposite banks, hereinafter referred to as “freshwater RdlP” (Fig. 1). Such places were defined as limit because the last two stable freshwater fisheries of RdlP are placed in their banks, in the middle of the limits proposed by Ringuelet (1975) and López et al. (2002; 2008).

FIGURE 1| Division of the Río de la Plata (RdlP) by the Barra del Indio shoal, in a riverine zone and an estuarine zone (FREPLATA, 2005), showing the sampled sites (black circles) and the limits of the area of freshwater RdlP considered for this study.

Literature review. Based on the list of freshwater fishes performed by López et al. (2003), species mentioned for the area in scientific literature, and our own fieldwork, a review of the species recorded for the freshwater RdlP was performed. The mention of each species was revised through the geographical distribution database of freshwater fishes from Argentina (Liotta, 2020). This online data source compile georeferenced information published in around 690 scientific publications on the distribution of inland fishes of the country. Furthermore, we conducted a systematic search in the Web of Science (WOS – in all fields) and Google Scholar database on Jan-2022, using the keywords: “scientific name” of each species and “rio de la Plata” (i.e., Prochilodus lineatus Río de la Plata). After these searches, around 1000 ichthyologycal documents were obtained including journal papers, governmental technical reports, books, conference presentations, and PhD theses. Then, we reviewed titles, abstracts and in many cases the full text of each document retaining only the surveys with actual fieldwork and capture location inside the delineated study area, obtaining 119 documents. The species were sorted in a list of fish species mentioned for the freshwater RdlP, following the classification of Nelson et al. (2016) and changes in taxonomy were reviewed and updated following Fricke et al. (2022). Each species was analyzed to confirm the number of records, defined as the number of times that the species were registered in field studies in the literature. Also, to this figure we added the species registered in our fieldwork as one more record. To solve old metadata replication from Ringuelet et al. (1967), species registered once with collection date before this work or older, and taxa without first record traceable were considered as “not confirmed”. Such timeframe in the history of ichthyological studies of RdlP was defined considering the publication date of the study of the distribution of fish species in Argentina conducted by Ringuelet et al. (1967). The review of the list also included the analysis of species included in the last lists of freshwater fish species from Argentina (Mirande, Koerber, 2015; Koerber et al., 2016a, 2020) and Uruguay (Teixeira de Mello et al., 2011; Litz, Koerber, 2014; Koerber, Litz, 2016b; Koerber et al., 2020, 2021). The species previously mentioned for the study area, but not included in these lists, were categorized as “absent” for the freshwater RdlP and excluded from further analysis. In addition, species currently considered junior synonyms of other species those documented in the study area were referred to as “synonymized species”. The remaining taxa, cited for Argentina and Uruguay within the study area, were categorized based on the absence or existence of specimens in ichthyological collections as “supported by literature” or “confirmed”, respectively. The non-native species were grouped as “established exotic species” and “non-established exotic species”, following Maiztegui et al. (2016), García Romero et al. (1998), and Liotta, Giacosa (2017).

Fieldwork. A fish sampling was carried out in freshwater RdlP at six sampling sites, along 35 km of the coastline between 34°46’56.0”S 58°00’39.9”W and 34°55’43.2”S 57°42’45.6”W (Fig. 1). Four seasonal fish samplings were performed from April 2016 to May 2017, using two fyke net with a rectangular mesh 10 x 5 mm (Colautti, 1998) and two trammel nets at each site, 25 m length and 1.9 m height each, with an inner mesh of 19 mm and 21 mm bar for one net, and 27 mm and 30 mm bar for the other net. The external mesh was 130 mm bar for both trammel nets. Fishing gears were arranged for one night, with a standardized effort at each site (16h). Fish were identified, counted, and weighed (g). During fieldwork, some fish were identified and released alive; others were euthanized with an overdose of benzocaine solution, fixed in 10% formaldehyde, and stored in 70% ethanol for laboratory identification. Fish were identified following Azpelicueta, Braga (1991), López, Miquelarena (1991), Braga (1993, 1994), Aquino (1997), Casciotta et al. (2005), Miquelarena, Menni (2005), Miquelarena et al. (2008), and Rosso et al. (2018). Changes in taxonomy were reviewed and updated following Fricke et al. (2022). Voucher species were deposited in the ichthyology collection of the Fundación de Historia Natural Félix de Azara, Buenos Aires, Argentina (CFA-IC). In addition, species collected for the first time in the area were indicated as “new records”.

Taking into account the categories supported by literature, confirmed, established exotic species, and new records showed in the list of fish species mentioned for the freshwater RdlP, we obtained the current fish assemblage of the freshwater RdlP with their respective number of records. Meanwhile, the taxa categorized as not confirmed, absent, and synonymized species were considered as not current components.

Literature and fieldwork. Comparison and representativeness. The relative distribution of species richness within each taxonomic order was compared between the current fish assemblage from the literature and that from our fieldwork using the test of goodness fit of Kolmogorov-Smirnov (p < 0.05). In turn, we calculated the frequency distribution of species and ratio between number of species captured in our fieldwork and number of species mentioned in the literature as function of number of records.

Hierarchical structure of fish assemblage. To describe the hierarchical structure of fish species, an Olmstead-Tukey (OT) diagram (Sokal, Rohlf, 1981) was drawn from the fieldwork sampled area, categorizing each species as “dominant”, “frequent”, “occasional” or “rare”. Such classification was made according to the arrangement of species in a two-dimensional space defined by the percentage frequency of occurrence of species (occurrence %) and their catch per unit effort in fish number (CPUEn) and in fish weight (CPUEw). In both cases, values were standardized to 16h of fishing and, log transformed (ln X+1). The category obtained for each species was indicated in the list of fish species mentioned for the freshwater RdlP.

Results

From the literature, 206 native and four exotic fish species were mentioned in freshwater RdlP (Tab. 1). From the analysis of each species, the following number by categories were defined: 48 not confirmed, 13 absent, five synonymized species, 29 supported by literature, 107 confirmed, one established exotic species, and three non-established exotic species.

A total of 28,147 individuals that weighted 912,723 g, encompassing 72 species distributed in 11 orders and 28 families were captured during fieldwork (Tab. 1). This figure represents half of the fish assemblage species reported in the literature, including four new records. These taxa, together with supported by literature, confirmed, and established exotic species categories found in the literature, define a list of 141 species that should be considered as current fish assemblage of the freshwater RdlP (Tab. 1). The remaining categories: not confirmed, absent, synonymized species, and non-established exotic species, accounted 66 taxa and were considered as not current components.

TABLE 1 | List of fish species mentioned for the freshwater Río de la Plata (RdlP) with their taxonomic position, discriminating between species categorized as current and not current component of the fish assemblage. Number of records (N°Rec), references, and category provided after literature review (Cat.: NC, not confirmed; Ab, absent; SS, synonymized species; SL: supported by literature; Co, confirmed; NR, new records; EEx, established exotic species; NEx, non-established exotic species). The species captured in the sampling were classified according to Olmstead-Tukey (OT CPUEn and CPUEw: D, dominant; F, frequent; R, rare; O, occasional) and their voucher number (VN) in ichthyology collections. Museo de La Plata, Instituto de Limnologia, La Plata (MLP); Museo Argenino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires (MACN); Laboratorio de Evolución of Facultad de Ciencias, Universidad de la República Uruguay, Montevideo (EVP); Museu de Ciências e Tecnologia, Pontifícia Universidade Católica do Rio Grande do Sul, Porto Alegre (MCP); Museo Nacional de Historia Natural y Antropologia, Zoologia, Montevideo (MHNM).

Classification	N°Rec	References	Cat.	OT		VN
Classification	N°Rec	References	Cat.	CPUEn	CPUEw	VN
Current components of the fish assemblage
CHONDRICHTHYES
MYLIOBATIFORMES
Potamotrygonidae
Potamotrygon brachyura (Günther, 1880)	3	CARU (2012); Llompart et al. (2012); Lucifora et al. (2015).	SL
OSTEICHTHYES

Sciaenidae
Pachyurus bonariensis Steindachner, 1879	8	Almirón (1989); Almirón et al. (2000); Llompart et al. (2012); CARU (2012, 2016); Paracampo (2013); Paracampo et al. (2020); This study.	Co	F		CFA8064
Plagioscion ternetzi Boulenger, 1895	8	Almirón (1989); Almirón et al. (2000); CARU (2012, 2016); Llompart et al. (2012); Paracampo (2013); Paracampo et al. (2020); This study.	Co	R		CFA8049
ATHERINIFORMES Atherinopsidae
Odontesthes argentinensis (Valenciennes, 1835)	1	D´Anatro et al. (2020)	Co			EVP150
Odontesthes humensis de Buen, 1953	1	Bogan et al. (2015).	Co			CFA4800
Odontesthes retropinnis (de Buen, 1953)	1	Cuello (2020).	Co			MLP9670
Odontesthes bonariensis (Valenciennes, 1835)	14	Almirón (1989); Almirón et al. (2000); CARU-CARP (2012, 2016); Paracampo (2013); Avigliano, Volpedo (2013a,b); Avigliano et al. (2015a); Valdés et al. (2015); Valencia et al. (2017); Villanova et al. (2018); Colautti et al. (2019); Scarabotti et al. (2021), This study.	Co	F
Odontesthes perugiae Evermann & Kendall, 1906	1	This study.	NR	R		CFA8087
CICHLIFORMES
Cichlidae
Australoheros facetus (Jenyns, 1842)	8	Ringuelet et al. (1967); Ringuelet et al. (1978); Almirón, García (1992); Almirón et al. (2000); López et al. (2009); Llamazares Vegh et al. (2012); Paracampo (2013); Paracampo et al. (2020).	Co			MLP12-XII-32-18
Cichlasoma dimerus (Heckel, 1840)	1	CARU-CARP (2016).	SL
Crenicichla lepidota Heckel, 1840	1	Llompart et al. (2012).	SL
Crenicichla scottii (Eigenmann, 1907)	2	Llamazares Vegh et al. (2012); Paracampo (2013).	Co			CFA2191
Gymnogeophagus meridionalis Reis & Malabarba, 1988	8	Escalante (1984); Almirón, García (1992); Almirón et al. (2000); Llamazares Vegh et al. (2012); Llompart et al. (2012); Paracampo (2013); Yorojo Moreno et al. (2017); This study.	Co	R		CFA8038
CLUPEIFORMES
Pristigasteridae
Pellona flavipinnis (Valenciennes, 1837)	4	Ringuelet et al. (1967); Remes Lenicov, Colautti (2000); CARU-CARP (2012, 2016).	Co			MLP2-VII-54-11
Engraulidae
Lycengraulis grossidens (Spix & Agassiz, 1829)	7	Ringuelet et al. (1967); Remes Lenicov, Colautti (2000); CARU-CARP(2012, 2016); Llompart et al. (2012); Scarabotti et al. (2021); This study.	Co	R		CFA8089
Platanichthys platana (Regan, 1917)	1	CARU-CARP (2012).	SL
Ramnogaster melanostoma (Eigenmann, 1907)	5	Ringuelet et al. (1967); Almirón (1989); Llompart et al. (2012); Paracampo (2013); This study.	Co	F		CFA8088
CYPRINIFORMES
Cyprinidae
Cyprinus carpio Linnaeus, 1758	13	Mac Donagh (1945); Candia (1989); Fabiano et al. (1992); Colautti (1997); Colombo et al. (2000); CARU-CARP (2012, 2016); Llamazares Vegh et al. (2012); Llompart et al. (2012); Paracampo (2013); Guerrero et al. (2017); Paracampo et al. (2020); This study.	EEx	F	D	CFA8096
CYPRINODONTIFORMES
Anablepidae
Jenynsia lineata (Fowler, 1940)	10	Ringuelet et al. (1967); Ringuelet et al. (1978); Escalante (1983); Almirón, García (1992); Almirón et al. (2000); López et al. (2009); Llamazares Vegh et al. (2012); Llompart et al. (2012); Paracampo (2013); Paracampo et al. (2020).	Co
Poeciliidae
Cnesterodon decemmaculatus (Jenyns, 1842)	9	Escalante (1983); Almirón, García (1992); Almirón et al. (2000); Remes Lenicov et al. (2005); López et al. (2009); Llamazares Vegh et al. (2012); Llompart et al. (2012); Paracampo (2013); Paracampo et al. (2020).	Co			CFA2163
Phalloceros caudimaculatus (Hensel, 1868)	4	Almirón (1989); Almirón et al. (2000); Callicó Fortunato et al. (2010); Llompart et al. (2012).	SL
Rivulidae
Austrolebias bellottii (Steindachner, 1881)	6	Escalante (1984); Almirón, García (1992); Almirón et al. (2000); Calviño (2007); Llompart et al. (2012); Calviño (2016).	Co			MLP6847
Austrolebias elongatus (Steindachner, 1881)	2	Ringuelet et al. (1967); Calviño (2016).	Co			MLP6408
Austrolebias nigripinnis (Regan, 1912)	4	Ringuelet et al. (1967); Calviño (2007); Calviño (2016) Llompart et al. (2012).	Co			MLP7889
CHARACIFORMES
Acestrorhynchidae
Acestrorhynchus pantaneiro Menezes, 1992	7	Almirón, García (1992); Almirón (1989); Almirón et al. (2000); Remes Lenicov, Colautti (2000); CARU-CARP (2012, 2016) ; This study.	Co	R		CFA8060
Anostomidae
Abramites hypselonotus (Günther, 1868)	1	Braga (1993).	Co			MLP17-XII-32-12
Leporinus acutidens (Valenciennes, 1837)	1	Almirón (1989)	SL
Megaleporinus obtusidens (Valenciennes, 1837)	11	Candia (1989); Almirón (1989); Braga (1993); Penchaszadeh et al. (2000); Remes Lenicov, Colautti (2000); García, Protogino (2005); CARU-CARP (2012, 2016); Paracampo (2013); Rojo et al. (2021); This study.	Co	F	D	CFA8082
Pseudanos trimaculatus (Kner, 1858)	1	Braga (1993).	Co			MACN6207
Schizodon borellii (Boulenger, 1900)	2	CARU-CARP (2012, 2016).	SL
Schizodon platae (Garman, 1890)	6	Almirón (1989); Almirón et al. (2000); Remes Lenicov, Colautti (2000); Llompart et al. (2012); Paracampo (2013); This study.	Co	F		CFA8086
Characidae
Aphyocharax dentatus Eigenmann & Kennedy, 1903	1	López et al. (1984).	SL
Astyanax abramis (Jenyns, 1842)	5	Almirón (1989); CARU-CARP (2012, 2016); Paracampo (2013); This study.	Co	R		CFA8099
Astyanax alleni (Eigenmann & McAtee, 1907)	3	Almirón (1984); Llompart et al. (2012); Paracampo (2013).	SL
Astyanax lacustris (Lütken, 1875)	4	Remes Lenicov, Colautti (2000); Llompart et al. (2012); CARU-CARP (2016); This study.	Co	F		CFA8072
Brycon orbignyanus (Valenciennes, 1850)	5	Ringuelet et al. (1967); CARU-CARP (2012, 2016); Guerrero et al. (2017); This study.	Co	R		MLP30-V-33-43
Bryconamericus exodon Eigenmann, 1907	1	Llompart et al. (2012).	SL
Bryconamericus iheringii (Boulenger, 1887)	7	Almirón (1989); Almirón et al. (2000); López et al. (2009); Llamazares Vegh et al. (2012); Paracampo (2013); Paracampo et al. (2020); This study.	Co	R		CFA8044
Bryconamericus rubropictus (Berg, 1901)	1	Almirón et al. (2000).	SL
Charax stenopterus (Cope, 1894)	7	Escalante (1984); Almirón, García (1992); Almirón et al. (2000); López et al. (2009); Llompart et al. (2012); Paracampo (2013); Paracampo et al. (2020).	Co			CFA2168
Cheirodon interruptus (Jenyns, 1842)	11	Ringuelet et al. (1967); Miquelarena et al. (1981); Almirón, García (1992); Almirón et al. (2000); Remes Lenicov et al. (2005); López et al. (2009); Llamazares Vegh et al. (2012); Llompart et al. (2012); Paracampo (2013); Paracampo et al. (2020); This study.	Co	R		CFA8043
Cynopotamus argenteus (Valenciennes, 1837)	7	Almirón (1989); Remes Lenicov, Colautti (2000); CARU-CARP (2012, 2016); Llompart et al. (2012); Paracampo (2013); This study.	Co	F		CFA8062
Cynopotamus kincaidi (Schultz, 1950)	1	This study.	NR	R
Deuterodon luetkenii (Boulenger, 1887)	1	Paracampo (2013).	SL
Diapoma terofali (Géry, 1964)	3	Miquelarena et al. (1981); Almirón et al. (2000); Paracampo (2013).	Co			MLP19-XII-79-1
Galeocharax humeralis (Valenciennes, 1834)	3	CARU-CARP (2012); Llompart et al. (2012); This study.	Co	R
Heterocheirodon yatai (Casciotta, Miquelarena & Protogino, 1992)	1	Almirón et al. (2000).	SL
Hyphessobrycon meridionalis Ringuelet, Miquelarena & Menni, 1978	5	Ringuelet et al. (1978); Almirón, García (1992); Almirón et al. (2000); López et al. (2009); Paracampo (2013).	Co			CFA2181
Hyphessobrycon togoi Miquelarena & López, 2006	1	Miquelarena, López (2006).	Co			CFA1246
Moenkhausia intermedia Eigenmann, 1908	1	This study.	NR	R		CFA8052
Odontostilbe pequira (Steindachner, 1882)	2	Almirón (1984); This study.	Co	R		CFA8069
Oligosarcus jenynsii (Günther, 1864)	8	Almirón, García (1992); Almirón et al. (2000); López et al. (2009); CARU-CARP (2012); Llamazares Vegh et al. (2012); Paracampo (2013); Paracampo et al. (2020); This study.	Co	R		CFA8090
Oligosarcus oligolepis (Steindachner, 1867)	7	Escalante (1984); Almirón et al. (2000); Llompart et al. (2012); Paracampo (2013); CARU-CARP (2016); Paracampo et al. (2020); This study.	Co	F		CFA8055
Piabarchus stramineus (Eigenmann, 1908)	1	Azpelicueta, Braga (1980).	SL
Psalidodon anisitsi (Eigenmann, 1907)	3	Ringuelet et al. (1978); Almirón (1989); Paracampo (2013).	Co			CFA2186
Psalidodon eigenmanniorum (Cope, 1894)	7	Escalante (1982); Almirón, García (1992); Almirón et al. (2000); López et al. (2009); Llompart et al. (2012); Paracampo (2013); Paracampo et al. (2020).	Co			CFA2175
Psalidodon erythropterus (Holmberg, 1891)	2	Llompart et al. (2012); This study.	Co	R		CFA8066
Psalidodon rutilus (Jenyns, 1842)	13	Ringuelet et al. (1967); Escalante (1984); Almirón, García (1992); Almirón et al. (2000); Remes Lenicov, Colautti (2000); Remes Lenicov et al. (2005); López et al. (2009); CARU-CARP (2012, 2016); Llompart et al. (2012); Paracampo (2013); Paracampo et al. (2020); This study.	Co	D	F	CFA8070
Pseudocorynopoma doriae Perugia, 1891	7	Ringuelet et al. (1967); Almirón, García (1992); Almirón et al. (2000); Remes Lenicov et al. (2005); López et al. (2009); Paracampo (2013); Paracampo et al. (2020).	Co			MLP22-XI-34-9
Roeboides descalvadensis Fowler, 1932	1	Lucena (2007).	Co			MCP10148
Roeboides microlepis (Reinhardt, 1851)	6	Ringuelet et al. (1967); Almirón (1989); Remes Lenicov, Colautti 2000; Llompart et al. (2012); CARU-CARP (2016); This study.	Co	R		CFA8050
Salminus brasiliensis (Cuvier, 1816)	14	Ringuelet et al. (1967); Bonetto et al. (1971); Escalante (1984); Sverlij, Espinach Ros (1986); Candia (1989); Almirón et al. (2000); Remes Lenicov, Colautti (2000); CARU-CARP (2012, 2016); Paracampo (2013); Guerrero et al. (2017); Rojo et al. (2021); Scarabotti et al. (2021); This study.	Co	F	D	CFA8085
Crenuchidae
Characidium rachovii Regan, 1913	6	Ringuelet et al. (1978); López et al. (1980); Almirón et al. (2000); Remes Lenicov et al. (2005); López et al. (2009); Paracampo (2013).	Co			MLP24-IV-79-2
Curimatidae
Cyphocharax platanus (Günther, 1880)	7	Almirón (1989); Remes Lenicov, Colautti (2000); CARU-CARP (2012, 2016); Llompart et al. (2012); Paracampo (2013); This study.	Co	D	F	CFA8053
Cyphocharax saladensis (Meinken, 1933)	3	Almirón (1990); Llompart et al. (2012); This study.	Co	R
Cyphocharax spilotus (Vari, 1987)	5	Almirón (1989); Azpelicueta, Almirón (1991); Llompart et al. (2012); CARU-CARP (2016); This study.	Co	R
Cyphocharax voga (Hensel, 1870)	9	Almirón, García (1992); Almirón et al. (2000); Remes Lenicov et al. (2005); López et al. (2009); Llompart et al. (2012); Paracampo (2013); CARU-CARP (2016); Paracampo et al. (2020); This study.	Co	D		CFA8054
Potamorhina squamoralevis (Braga & Azpelicueta, 1983)	2	CARU-CARP (2016); This study.	Co	R
Psectrogaster curviventris Eigenmann & Kennedy, 1903	2	CARU-CARP (2016); This study.	Co	R
Steindachnerina biornata (Braga & Azpelicueta, 1987)	5	Braga, Azpelicueta (1987); Almirón et al. (2000); López et al. (2009); Paracampo (2013); This study.	Co	R		CFA8078
Cynodontidae
Rhaphiodon vulpinus Spix & Agassiz, 1829	7	Ringuelet et al. (1967); Almirón (1989); Almirón et al. (2000); Remes Lenicov, Colautti (2000); Llompart et al. (2012); Paracampo (2013); This study.	Co	R		CFA8057
Erythrinidae
Hoplias argentinensis Rosso, González-Castro, Bogan, Cardoso, Mabragaña, Delpiani & Díaz de Astarloa, 2018	12	Ringuelet et al. (1967); Escalante (1984); Almirón, García (1992); Almirón et al. (2000); Remes Lenicov et al. (2005); López et al. (2009); CARU-CARP (2012, 2016); Llamazares Vegh et al. (2012); Paracampo (2013); Paracampo et al. (2020); This study.	Co	F	D	CFA8042
Parodontidae
Apareiodon affinis (Steindachner, 1879)	3	Almirón (1989); Paracampo (2013); This study.	Co	R		CFA8068
Prochilodontidae
Prochilodus lineatus (Valenciennes, 1837)	32	Ringuelet et al. (1967); Almirón (1989); Candia (1989); Almirón, García (1992); Almirón et al. (2000); Colombo et al. (2000, 2011); Remes Lenicov, Colautti (2000); Villar et al. (2001); Remes Lenicov et al. (2005); Colombo et al. (2007); Speranza, Colombo (2009); Lombardi et al. (2010a,b); CARU-CARP (2012, 2016); Llamazares Vegh et al. (2012); Llompart et al. (2012); Speranza et al. (2012, 2013, 2016, 2020); Paracampo (2013); Cappelletti et al. (2015); Avigliano et al. (2017b, 2019b,c, 2020); Paracampo et al. (2020); Rojo et al. (2021); Scarabotti et al. (2021); This study.	Co	F	D	CFA8063
Serrasalmidae
Metynnis otuquensis Ahl, 1923	1	Braga (1994).	Co			MACN5700
Mylossoma duriventre (Cuvier, 1818)	3	Ringuelet et al. (1967); Remes Lenicov, Colautti (2000); CARU-CARP (2016).	Co			MLP7-XII-32-40
Pygocentrus nattereri Kner, 1858	4	Almirón (1989); Remes Lenicov, Colautti (2000); CARU-CARP (2012, 2016).	SL
Serrasalmus maculatus Kner, 1858	5	Almirón (1989); Remes Lenicov, Colautti (2000); CARU-CARP (2012, 2016); Llompart et al. (2012).	SL
Serrasalmus marginatus Valenciennes, 1837	1	Remes Lenicov, Colautti (2000).	SL
Triportheidae
Triportheus nematurus (Kner, 1858)	3	Ringuelet et al. (1967); Remes Lenicov, Colautti (2000); CARU-CARP (2016).	Co			MLP5-VI-35-15
GYMNOTIFORMES
Apteronotidae
Apteronotus albifrons (Linnaeus, 1766)	2	Ringuelet et al. (1967); CARU-CARP (2016).	Co			MLP20-X-32-32
Gymnotidae
Gymnotus inaequilabiatus (Valenciennes, 1839)	1	Ringuelet et al. (1967).	Co			MLP4-IV-61-48
Rhamphichthyidae
Rhamphichthys hahni (Meinken, 1937)	3	Ringuelet et al. (1967); CARU-CARP (2016); This study.	Co	R
Sternopygidae
Eigenmannia trilineata López & Castello, 1966	4	López, Castello (1966); Llompart et al. (2012); CARU-CARP (2016); This study.	Co	F		CFA8056
Eigenmannia virescens (Valenciennes, 1836)	5	Ringuelet et al. (1967); Remes Lenicov, Colautti (2000); CARU-CARP (2012, 2016); This study.	Co	R
MUGILIFORMES
Mugilidae
Mugil liza Valenciennes, 1836	7	Ringuelet et al. (1967); Almirón (1989); Almirón et al. (2000); Colombo et al. (2000); Paracampo (2013); CARU-CARP (2016); This study.	Co	R		CFA8059
SILURIFORMES
Ariidae
Genidens barbus (Lacepède, 1803)	5	CARU-CARP (2012); Avigliano et al. (2015b, 2017a, 2019a); Maciel et al. (2021).	SL
Aspredinidae
Bunocephalus doriae Boulenger, 1902	1	Paracampo (2013).	Co			CFA2162
Pseudobunocephalus iheringii (Boulenger, 1891)	2	Almirón et al. (2000); This study.	Co	R		CFA8079
Auchenipteridae
Ageneiosus inermis (Linnaeus, 1766)	3	Ringuelet et al. (1967); Volpedo, Fuchs (2010); CARU-CARP (2012).	Co			MLP1-III-60-1
Ageneiosus militaris Valenciennes, 1835	8	Ringuelet et al. (1967); Almirón et al. (2000); Llompart et al. (2012); CARU-CARP (2012, 2016); Paracampo (2013); Guerrero et al. (2017); This study.	Co	R		CFA8041
Auchenipterus osteomystax (Miranda Ribeiro, 1918)	5	Ringuelet et al. (1967); Almirón et al. (2000); CARU-CARP (2012, 2016); Llompart et al. (2012).	Co			MLP1-V-41-19
Trachelyopterus albicrux (Berg, 1901)	1	Remes Lenicov, Colautti (2000).	SL
Trachelyopterus cf. galeatus (Linnaeus, 1766)	5	Ringuelet et al. (1967); Almirón et al. (2000); Llompart et al. (2012); Guerrero et al. (2017); This study.	Co	F		CFA8058
Trachelyopterus lucenai Bertoletti, da Silva & Pereira, 1995	1	CARU-CARP (2016).	SL
Callichthydae
Callichthys callichthys (Linnaeus, 1758)	3	Ringuelet et al. (1967) Remes Lenicov et al. (2005); Llompart et al. (2012).	Co			MLP2-III-59-4
Corydoras paleatus (Jenyns, 1842)	9	Ringuelet et al. (1967); Escalante (1983); Almirón et al. (2000); Remes Lenicov et al. (2005); López et al. (2009); Llompart et al. (2012); Paracampo (2013); Paracampo et al. (2020); This study.	Co	F		CFA8039
Doraridae
Oxydoras kneri Bleeker, 1862	3	Ringuelet et al. (1967); Llompart et al. (2012); Scarabotti et al. (2021).	Co			MLP30-V-33-2
Pterodoras granulosus (Valenciennes, 1821)	12	Candia (1989); Amestoy, Fabiano (1992); Remes Lenicov, Colautti (2000); Ferriz et al. (2000); Villar et al. (2001); Cataldo (2002); Garcia, Protogino (2005); Volpedo, Fuchs (2010); CARU-CARP (2012, 2016); Scarabotti et al. (2021); This study.	Co	R		CFA8076
Rhinodoras dorbignyi (Kner, 1855)	6	Ringuelet et al. (1967); Garcia, Protogino (2005); CARU-CARP (2012, 2016); Llompart et al. (2012); This study.	Co	R		CFA8074
Heptapteridae
Heptapterus mustelinus (Valenciennes, 1835)	2	Ringuelet et al. (1967); Fuster (2017).	Co			MLP10-VIII-32-24
Pimelodella gracilis (Valenciennes, 1835)	6	Almirón (1989); Almirón, García (1992); Almirón et al. (2000); CARU-CARP (2012, 2016); This study.	Co	R		CFA8075
Pimelodella laticeps Eigenmann, 1917	8	Almirón et al. (2000); Almirón, García (1992); Llompart et al. (2012); López et al. (2009); Paracampo (2013); García et al. (2017); Paracampo et al. (2020); This study.	Co	R		CFA8081
Rhamdia aff. quelen (Quoy & Gaimard, 1824)	8	Almirón, García (1992); Almirón et al. (2000); Llamazares Vegh et al. (2012); Llompart et al. (2012); Paracampo (2013); CARU-CARP (2012); Paracampo et al. (2020); This study.	Co	R		CFA8067
Loricariidae
Ancistrus cirrhosus (Valenciennes, 1836)	3	Ringuelet et al. (1967); Almirón et al. (2000); This study.	Co	R		CFA8061
Brochiloricaria chauliodon Isbrücker, 1979	2	Almirón et al. (2000); Garcia, Protogino (2005).	SL
Hisonotus maculipinnis (Regan, 1912)	1	Almirón et al. (2000).	SL
Hoplosternum littorale (Hancock, 1828)	3	Ringuelet et al. (1967); Llompart et al. (2012); This study.	Co	R		CFA8091
Hypostomus aspilogaster (Cope, 1894)	3	Cardoso et al. (2011); CARU-CARP (2016); This study.	Co	R		CFA8093
Hypostomus borellii (Boulenger, 1897)	1	CARU-CARP (2012).	SL
Hypostomus commersoni Valenciennes, 1836	12	Ringuelet et al. (1967); Almirón, García (1992); Almirón et al. (2000); Remes Lenicov et al. (2005); Volpedo, Fuchs (2010); Llamazares Vegh et al. (2012); Llompart et al. (2012); Paracampo (2013); CARU-CARP (2012, 2016); Paracampo et al. (2020); This study.	Co	F	D	CFA8040
Hypostomus laplatae (Eigenmann, 1907)	3	Ringuelet et al. (1967); Cardoso et al. (2019); This study.	Co	R		CFA8097
Hypostomus microstomus Weber, 1987	1	López, Miquelarena (1991).	Co			MHNM1524
Hypostomus roseopunctatus Reis, Weber & Malabarba, 1990	1	CARU-CARP (2016).	SL
Hypostomus uruguayensis Reis, Weber & Malabarba, 1990	1	This study.	NR	R		CFA8094
Loricaria simillima Regan, 1904	1	CARU-CARP (2016).	Co
Loricaria apeltogaster Boulenger, 1895	2	López (1970); CARU-CARP (2016).	SL
Loricariichthys anus (Valenciennes, 1835)	10	López (1970); Almirón, García (1992); Almirón et al. (2000); Volpedo, Fuchs (2010); CARU-CARP (2012, 2016); Llompart et al. (2012); Paracampo (2013); Paracampo et al. (2020); This study.	Co	F	D	CFA8083
Loricariichthys melanocheilus Reis & Pereira, 2000	2	Rodríguez (2003); CARU-CARP (2012).	Co			MLP755
Otocinclus arnoldi Regan, 1909	7	Almirón, García (1992); Schaefer (1997); Almirón et al. (2000); López et al. (2009); Llompart et al. (2012); Paracampo (2013); Paracampo et al. (2020).	Co			CFA2170
Otocinclus vittatus Regan, 1904	1	Almirón et al. (2000).	SL
Paraloricaria vetula (Valenciennes, 1835)	7	López (1970); Rodríguez (2003); Garcia, Protogino (2005); Volpedo, Fuchs (2010); CARU-CARP (2012, 2016); This study.	Co	R		CFA8077
Pterygoplichthys ambrosettii (Holmberg 1893)	2	CARU-CARP (2016); This study.	Co	R		CFA8036
Rhinelepis strigosa Valenciennes, 1840	3	CARU-CARP (2012, 2016); This study.	Co	F		CFA8095
Ricola macrops (Regan, 1904)	5	Ringuelet et al. (1967); López (1970); Rodríguez (2003); Garcia, Protogino (2005); CARU-CARP (2016).	Co
Rineloricaria catamarcensis (Berg, 1895)	1	Rodríguez (2003).	Co			CFA1680
Rineloricaria lima (Kner, 1853)	2	López (1970); Almirón, García (1992).	SL
Pimelodidae
Hypophthalmus oremaculatus Nani & Fuster de Plaza, 1947	1	López et al. (1980).	Co			MLP2-VIII-73-13
Iheringichthys labrosus (Lütken, 1874)	7	Volpedo, Fuchs (2010); CARU-CARP (2012, 2016); Llompart et al. (2012); Paracampo (2013); Paracampo et al. (2020); This study.	Co	F		CFA8073
Luciopimelodus pati (Valenciennes, 1835)	11	Ringuelet et al. (1967), Sverlij, Espinach Ros (1986); Candia (1989); Almirón et al. (2000); Remes Lenicov, Colautti (2000); Volpedo, Fuchs (2010); CARU-CARP (2012, 2016); Llompart et al. (2012); Scarabotti et al. (2021); This study.	Co	F		CFA8084
Pinirampus argentina (MacDonagh, 1938)	1	Ringuelet et al. (1967).	Co			MLP30-V-33-3
Pimelodus maculatus Lacepède, 1803	17	Ringuelet et al. (1967); Candia (1989); Almirón, García (1992); Gutiérrez, Martorelli (1999); Almirón et al. (2000); Remes Lenicov, Colautti (2000); Garcia, Protogino (2005); Remes Lenicov et al. (2005); López et al. (2009); Volpedo, Fuchs (2010); CARU-CARP (2012, 2016); Llamazares Vegh et al. (2012); Llompart et al. (2012); Paracampo (2013); Paracampo et al. (2020); Rojo et al. (2021); This study.	Co	D		CFA8071
Parapimelodus valenciennis (Lütken, 1874)	13	Ringuelet et al. (1967); Almirón (1989); Almirón et al. (2000); Remes Lenicov, Colautti (2000); Remes Lenicov et al. (2005); Volpedo, Fuchs (2010); CARU-CARP (2012, 2016); Llompart et al. (2012); Paracampo (2013); Paracampo et al. (2020); Scarabotti et al. (2021); This study.	Co	D		CFA8047
Pimelodus albicans (Valenciennes, 1840)	13	Ringuelet et al. (1967); Almirón (1989); Candia (1989); Remes Lenicov, Colautti (2000); Gutiérrez, 2001; Remes Lenicov et al. (2005); Vergara et al. (2008); Volpedo, Fuchs (2010); CARU-CARP (2012, 2016); Llamazares Vegh et al. (2012); Llompart et al. (2012); Paracampo, 2013; This study.	Co	F	D	CFA8098
Pimelodus argenteus Perugia, 1891	3	Volpedo, Fuchs (2010); Llompart et al. (2012); This study.	Co	R		CFA8048
Pseudoplatystoma corruscans (Spix & Agassiz, 1829)	6	Ringuelet et al. (1967); Remes Lenicov, Colautti (2000); CARU-CARP (2012, 2016); Paracampo (2013); This study.	Co	F		CFA8035
Sorubim lima (Bloch & Schneider, 1801)	5	Almirón (1989); Remes Lenicov, Colautti (2000); Volpedo, Fuchs (2010); Llompart et al. (2012); This study.	Co	R		CFA8092
Zungaro jahu (Ihering, 1898)	3	Llompart et al. (2012); CARU-CARP (2016); Guerrero et al. (2017).	SL
Pseudopimelodidae
Microglanis malabarbai Bertaco & Cardoso, 2005	5	Castello (1971); Almirón et al. (2000); Llompart et al. (2012); Paracampo (2013); This study.	Co	R		CFA8080
SYNBRANCHIFORMES
Synbranchidae
Synbranchus marmoratus Bloch, 1795	11	Ringuelet et al. (1967); Almirón, García (1992); Almirón et al. (2000); Remes Lenicov et al. (2005); López et al. (2009); Llamazares Vegh et al. (2012); Llompart et al. (2012); Paracampo (2013); Guerrero et al. (2017); Paracampo et al. (2020); This study.	Co	R		CFA8046
PLEURONECTIFORMES
Achiridae
Catathyridium jenynsii (Günther, 1862)	4	Candia (1989); Lunaschi (2003); Llompart et al. (2012); This study.	Co	R		CFA8065
Not current components of the fish assemblage
PETROMYZONTIDA
PETROMYZONTIFORMES
Geotriidae
Geotria macrostoma (Burmeister, 1868)	1	Burmeister (1868).	NC
CHONDRICHTHYES
MYLIOBATIFORMES
Potamotrygonidae
Potamotrygon histrix (Müller & Henle, 1839)	1	Pozzi (1945).	NC
Potamotrygon motoro (Müller & Henle, 1841)	1	Pozzi (1945).	NC
OSTEICHTHYES
ACIPENSERIFORMES
Acipenseridae
Acipenser baerii Brandt, 1869	2	Azpelicueta, Almirón (1999); Liotta, Giacosa (2017).	NEx
Acipenser gueldenstaedtii Brandt & Ratzeburg, 1833	1	Demonte et al. (2017).	NEx
ANGUILLIFORMES
Muraenidae
Gymnothorax ocellatus Agassiz, 1831	1	Marelli (1924).	NC
ACANTHURIFORMES
Sciaenidae
Plagioscion macdonaghi Daneri, 1954		Junior synonim of Plagioscion ternetzi Boulenger, 1895 by Casatti (2003).	SS
CLUPEIFORMES
Clupeidae
Brevoortia aurea (Spix & Agassiz, 1829)	1	Lahille (1895).	NC
Brevoortia pectinata (Jenyns, 1842)	1	Berg (1895).	NC
Ramnogaster arcuata (Jenyns, 1842)	1	De Buen (1950) collected outside the study area (Montevideo, Uruguay)	NC
CHARACIFORMES
Anostomidae
Leporinus striatus Kner, 1858	1	Devincenzi, Teague (1942) collected outside the study area (middle Uruguay River).	NC
Characidae
Aphyocharax pusillus Günther, 1868	1	Pozzi (1945).	NC
Astyanax orbignyanus (Valenciennes, 1850)	1	Cuvier, Valenciennes (1850).	NC
Diapoma speculiferum Cope, 1894	1	Myers (1942) collected outside the study area (Río Cebollatí, Uruguay).	NC
Hyphessobrycon bifasciatus Ellis, 1911	1	Messner (1962).	NC
Markiana nigripinnis (Perugia, 1891)	1	Perugia (1891).	NC
Odontostilbe microcephala Eigenmann, 1907	1	Ringuelet (1975) without traceable record.	NC
Poptella paraguayensis (Eigenmann, 1907)	1	Günther (1880).	NC
Roeboides affinis (Günther, 1868)	1	Ringuelet, Arámburu (1957).	NC
Serrapinnus piaba (Lütken, 1875)	1	Nion (1998) without traceable record.	NC
Tetragonopterus argenteus Cuvier, 1816	1	Lahille (1895).	NC
Curimatidae
Steindachnerina brevipinna (Eigenmann & Eigenmann, 1889)	1	Berg (1897).	NC
Gasteropelecidae
Thoracocharax stellatus (Kner, 1858)	1	Pozzi, Bordalé (1936).	NC
Lesbiasinidae
Pyrrhulina australis Eigenmann & Kennedy, 1903	1	Pozzi (1945).	NC
Parodontidae
Parodon suborbitalis Valenciennes, 1850	1	De Buen (1950).	NC
Serrasalmidae
Mylossoma paraguayensis Norman, 1929		Junior synonim of Mylossoma duriventre (Cuvier, 1818) Jégu (2003)	SS
Piaractus mesopotamicus (Holmberg, 1887)	1	Pozzi (1945).	NC
Serrasalmus spilopleura Kner, 1858	1	Norman (1929).	Ab
Trichomycteridae
Pseudostegophilus maculatus (Steindachner, 1879)	1	Stigchel (1947).	NC
Scleronema minutum (Boulenger, 1891)	1	Devincenzi, Teague (1942) collected outside the study area (middle Uruguay River).	NC
CICHLIFORMES
Cichlidae
Crenicichla lacustris (Castelnau, 1855)	1	Marelli (1924).	Ab
Gymnogeophagus australis (Eigenmann, 1907)	1	Pozzi (1945).	NC
Gymnogeophagus labiatus (Hensel, 1870)	1	Pereyra, García (2008) without traceable record.	NC
Gymnogeophagus rhabdotus (Hensel, 1870)	1	Nion (1998) without traceable record.	NC
CYPRINIFORMES
Cyprinidae
Hypophthalmichthys molitrix (Valenciennes, 1844)	1	García Romero et al. (1998).	NEx
CYPRINODONTIFORMES
Poeciliidae
Phalloptychus januarius (Hensel, 1868)	1	Lahille (1923).	Ab
Poecilia vivipara Bloch & Schneider, 1801	1	Ihering (1931).	Ab
GYMNOTIFORMES
Rhamphichthydae
Rhamphichthys rostratus (Linnaeus, 1766)	1	Lahille (1910).	Ab
Hypopomidae
Hypopomus artedi (Kaup, 1856)	1	Lahille (1921).	Ab
MUGILIFORMES
Mugilidae
Mugil platanus Günther, 1880		Junior synonim of Mugil liza Valenciennes, 1836 by Menezes et al. (2010).	SS
SILURIFORMES
Ageneiosidae
Ageneiosus dentatus Kner, 1858	1	Marelli (1924).	NC
Auchenipteridae
Trachelyopterus ceratophysus (Kner, 1858)	1	Perugia (1891).	NC
Trachelyopterus striatulus (Steindachner, 1877)	1	Mac Donagh, Cabrera (1937).	NC
Callichthydae
Corydoras undulatus Regan, 1912	1	Nijssen, Isbrücker (1980) collected by Wolterstorff in 1912.	NC
Corydoras aeneus (Gill, 1858)	1	Pozzi (1945).	NC
Doraridae
Megalodoras laevigatulus (Berg, 1901)		Junior synonim of Pterodoras granulosus (Valenciennes, 1821) by Sabaj (2003).	SS
Loricariidae
Ancistrus gymnorhynchus Kner, 1854	1	Stigchel (1947) misidentified this species which is distributed in Venezuela (Fricke et al., 2022).	Ab
Farlowella hahni Meinken, 1937	1	Almirón et al. (1992) without traceable record.	NC
Hypostomus punctatus Valenciennes, 1840	1	Angelescu, Gneri (1949).	NC
Hypostomus robinii Valenciennes, 1840	1	Lopez et al. (1987) misidentified this species which is distributed in Colombia (Fricke et al., 2022).	Ab
Loricariichthys platymetopon Isbrücker & Nijssen, 1979	1	Roule (1826).	NC
Otocinclus affinis Steindachner, 1877	1	Schaefer (1997).	Ab
Paraloricaria commersonoides (Devincenzi, 1943)	1	De Vincenzi (1943) collected outside the study area (middle Uruguay River).	NC
Rineloricaria felipponei (Fowler, 1943)	1	Fowler (1943).	NC
Rineloricaria pareiacantha (Fowler, 1943)	1	Fowler (1943).	NC
Rineloricaria thrissoceps (Fowler, 1943)	1	Fowler (1943).	NC
Spatuloricaria nudiventris (Valenciennes, 1840)	1	Devincenzi, Teague (1942) collected outside the study area (middle Uruguay River).	NC
Heptapteridae
Myoglanis colletti (Steindachner, 1881)	1	Steindachner (1881).	NC
Rhamdella jenynsii (Günther, 1864)	1	Günther (1880).	NC
Pimelodidae
Bergiaria platana (Steindachner, 1908)	1	Steindachner (1908).	NC
Bergiaria westermanni (Lütken, 1874)	1	Lütken (1874).	NC
Megalonema platanum (Günther, 1880)	1	Günther (1880).	NC
Hemisorubim platyrhynchos (Valenciennes, 1840)	1	Nion (1998) without traceable record.	NC
Hypophthalmus edentatus Spix & Agassiz, 1829	1	Lopez et al. (1980) erroneous record.	Ab
Pimelodus brevis Marini, Nichols & LaMonte, 1933	1	Junior synonim of Pimelodus argenteus Perugia, 1891 by Mirande, Koerber (2015).	SS
Paulicea luetkeni (Steindachner, 1876)	1	Haseman (1911).	Ab
Pseudopimelodus mangurus (Valenciennes, 1835)	1	Marelli (1924).	Ab
Pseudoplatystoma reticulatum Eigenmann & Eigenmann, 1889	1	De Buen (1950) without traceable record.	NC
PLEURONECTIFORMES
Achiridae
Achirus lineatus (Linnaeus, 1758)	1	Pozzi (1945) misidentified this tropical species which is distributed up to northern Argentina (Fricke et al., 2022).	Ab

The comparison between literature and sampling taxa composition showed no differences in the relative number distribution of species by order (Kolmogorov Smirnov, D = 0.38; p = 0.29). It also showed that Siluriformes and Characiformes were equally prevalent, representing around 80% of the taxa (Fig. 2), followed by Gymnotiformes, Atheriniformes, and Clupeiformes with 4–3%. Cichliformes, Acanthuriformes, Cypriniformes, Mugiliformes, Pleuronectiformes, and Synbranchiformes ranged between 4–1%. Cyprinodontiformes and Myliobatiformes were found only in the literature, with 4% or less. Considering families, comparison evidenced that Characidae, Loricariidae, and Pimelodidae, in descending order, were the richest taxa.

The category supported by literature, reached the highest value for taxa with a single record, followed by confirmed taxa and new records. Based on the 141 taxa that should be considered as current fish assemblage, the number of species by number of records value (Tab.1) showed that half of these species showed three records or less in the study area, and that the remaining half were recorded up to 32 (Fig. 3). As records increased, the ratio between number of species caught in fieldwork and number of species mentioned in the literature increased, suggesting that the species most frequently cited were also those most commonly captured in the study area.

FIGURE 2| Percentage number of species by order, literature review (black bar) differing from fieldwork (grey bar) in the freshwater Río de la Plata (RdlP).

The Olmstead-Tukey quadrants for occurrence % – CPUEn (Fig. 4A) were limited by a mean frequency of occurrence of 30.67% and an average CPUE of 8 ind./16 h. Five species were dominant (6.94%), 22 frequent (30.6%) and 45 rare (62.5%) (Tab.1). Dominant species exhibited an occurrence higher than 88%, representing 90% of the total specimens collected (n = 25,311). Frequent species accounted for 8.5% of the total captured fish and rare species, representing 1.5% of the total captured specimens. While for occurrence % – CPUEw an average 264 g/16h was obtained, with 11 dominant (11.2%), 16 frequent (22.2%), and 45 rare species (62.5%) (Fig. 4B). Summed up, the group of five dominant species exhibited a CPUEw higher than 90% of the total weight collected (g = 828,531). Frequent species accounted for 6% and rare species 3.2% of the total captured biomass. Occurrence % – CPUEn and occurrence % – CPUEw analyses evidenced that Parapimelodus valenciennis (Lütken, 1874) and Pimelodus maculatus Lacepède, 1803 were dominant taxa, that no occasional species were found, and that the rare species were the same.

FIGURE 3| Number of records of each species of the current fish assemblage of the freshwater Río de la Plata (RdlP) (SL, supported by literature; NR, new records; EEx, established exotic species; Co, confirmed) with their respective number of species and ratio between number of species captured in fieldwork and number of species mentioned in the literature.

In turn, main differences between CPUEn and CPUEw were the hierarchy change from dominant to frequent of Psalidodon rutilus (Jenyns, 1842) and Cyphocharax platanus (Günther, 1880); and the category change from frequent to dominant of Prochilodus lineatus (Valenciennes, 1837), Salminus brasiliensis (Cuvier, 1816), Megaleporinus obtusidens (Valenciennes 1837), Cyprinus carpio Linnaeus, 1758, Hoplias argentinensis Rosso, González-Castro, Bogan, Cardoso, Mabragaña, Delpiani & Díaz de Astarloa, 2018, Loricariichthys anus (Valenciennes, 1835), Hypostomus commersoni Valenciennes, 1836, and Pimelodus albicans (Valenciennes, 1840).

FIGURE 4| Olmstead-Tukey diagram showing the relationship between percentage frequency of occurrence of species (occurrence %) and log transformed (ln X+1) fish number and weight captured and standardized to 16h of fishing A. CPUEn and B. CPUEw in the freshwater Río de la Plata (RdlP), respectively. Grey lines correspond to the mean frequency of occurrence % (vertical) and average CPUEn and CPUEw (horizontal); they are used to define dominant, frequent, occasional, and rare species (categories are defined in Tab. 1). 1. Acestrorhynchus pantaneiro; 2. Ageneiosus militaris;3. Ancistrus cirrhosis;4. Apareiodon affinis; 5. Astyanax abramis; 6. Psalidodon erythropterus; 7. Astyanax lacustris; 8. Psalidodon rutilus; 9. Bryconamericus iheringii; 10. Brycon orbignyanus;11. Catathyridium jenynsii; 12. Cheirodon interruptus; 13. Corydoras paleatus; 14. Cynopotamus argenteus; 15. Cyprinus carpio; 16. Cynopotamus kincaidi; 17. Cyphocharax platanus; 18. Cyphocharax saladensis; 19. Cyphocharax spilotus; 20. Cyhocharax voga; 21. Eigenmannia trilineata; 22. Eigenmannia virescens; 23. Galeocharax humeralis; 24. Gymnogeophagus meridionalis; 25. Hoplosternum littorale; 26. Hoplias argentinensis; 27. Hypostomus aspilogaster; 28. Hypostomus commersoni; 29. Hypostomus laplatae; 30. Hypostomus uruguayensis; 31. Iheringichthys labrosus; 32. Megaleporinus obtusidens; 33. Loricariichthys anus; 34. Luciopimelodus pati; 35. Lycengraulis grossidens; 36. Microglanis malabarbai; 37. Moenkhausia intermedia; 38. Mugil liza; 39. Odontesthes bonariensis; 40. Odontesthes perugiae; 41. Odontostilbe pequira; 42. Oligosarcus jenynsii; 43. Oligosarcus oligolepis; 44. Pachyurus bonariensis; 45. Parapimelodus valenciennis;46. Paraloricaria vetula; 47. Pimelodus albicans; 48. Pimelodus argenteus; 49. Pimelodella gracilis; 50.Pimelodella laticeps; 51. Pimelodus maculatus; 52. Plagioscion ternetzi; 53. Potamorhina squamoralaevis; 54. Prochilodus lineatus; 55. Pseudoplatystoma corruscans; 56. Psectrogaster curviventris; 57. Pseudobunocephalus iheringii; 58. Pterygoplichthys ambrosettii; 59. Pterodoras granulosus; 60. Ramnogaster melanostoma; 61. Rhinodoras dorbigyi; 62. Rhamphichthys hahni; 63. Rhamdia aff. quelen; 64. Rhinelepis strigosa; 65. Rhaphiodon vulpinus; 66. Roeboides microlepis; 67. Salminus brasiliensis; 68. Schizodon platae; 69. Sorubim lima; 70. Steindachnerina biornata; 71. Synbranchus marmoratus; 72. Trachelyopterus galeatus.

Discussion

The present study puts forward an updated list of fish species that inhabit the freshwater RdlP; this is also the first proposal of the hierarchical composition of its Argentinean coastal fish assemblage. From the 206 native fish mentioned in the freshwater RdlP, literature showed 66 taxa categorized as not confirmed, absent, or synonymized species. The absence of not confirmed species in fish samplings for at least five decades, when most samplings in the area were developed, does not allow confirming their current inclusion in the fish assemblage under study.Further studies on this topic are therefore needed, whose results can potentially reduce the number of species in this group. However, it is highly probable that the poor water quality and habitat conditions evidenced in freshwater RdlP due to anthropogenic activities (Topalián et al., 1990; Kurucz et al., 1998; Natale, 2005; FREPLATA, 2005; Cirelli, Ojeda, 2008) have been impacting negatively on the richness of the fish assemblage of the area for the last five decades or so. Indeed, the disappearance of certain species in the freshwater coastal surrounding areas of La Plata city has already been correlated with local human pollution in the 70s decade (Ringuelet, 1975). As example, the nonappearance of a relevant species for fisheries in the basin, like Piaractus mesopotamicus (Holmberg, 1887) might be associated with this situation, as suggested by Quirós (1990) who linked its abundance declination with human pollution. The species included in the absent and synonymized species categories, based on current taxonomic studies (Litz, Koerber, 2014; Koerber, Litz, 2014; Koerber et al., 2016a,b; 2020), must be excluded from the list of fish species for the area as some correspond to confirmed taxonomic misidentifications (i.e., Hypostomus robinii Valenciennes, 1840, Ancistrus gymnorhynchus Kner, 1854, and Achirus lineatus (Linnaeus, 1758)) or outdated labelling.

The 140 native species that should be considered as current fish fauna of the study area were grouped into supported by literature, confirmed, and new records, reducing the number of autochthonous species previously compiled (Nion, 1998; López et al., 2003; Volpedo et al., 2010). Despite the historical relevance of previous compilations, our study suggests that the richness of species in freshwater RdlP is overestimated due to outdated old metadata replication. The supported by literature category refers to species that might represent misidentifications or taxa that migrate exceptionally from upper portions of the basin but do not become stablished. Either way, the lacking voucher number, making impossible to confirm their presence. This assumption is reinforced by the fact that the species in the supported by literature category were restricted to taxa recorded in five or less events, being maximum for species recorded once. Meanwhile, the confirmed group reflects species that undoubtedly are currently present in the study area. Such conclusion was based not only on the fact that these taxa were recently captured with voucher number deposit, but also on the fact that most were collected more than once. Indeed, it is highly probable that the taxa obtained two or more times, in independent fish samplings, represent usual fish fauna of the study area. However, species recently found once, as in the new records category, if not re-captured, could be re-categorized as not confirmed in subsequent research. In general, species recently captured once in the freshwater RdlP are commonly found in the upper parts of the La Plata River basin (Ringuelet, 1975), possibly extending their distribution. Volpedo et al., (2010) reported an increase in the number of taxa of RdlP over the last two decades owing to climate change. The referred mechanism could function as a counterpart of the effects caused by pollution and habitat modification, influencing fish assemblage composition and structure.

In the established exotic species category, the establishment and expansion of Cyprinus carpio are common in environments connected with the Río de la Plata system in Argentina and Uruguay (Teixeira de Melo et al., 2011; Maiztegui et al., 2016). This is reinforced by the Olmstead-Tukey analysis outcomes, indicating that this fish is frequently found in the study area. In contrast, non-established exotic species can occasionally appear in freshwater RdlP; yet, their presence is mostly due to their escape from aquaculture facilities within the basin (i.e., Acipenser baerii Brandt, 1869, Acipenser gueldenstaedtii Brandt & Ratzeburg, 1833, and Hypophthalmichthys molitrix (Valenciennes, 1844)).

The fish fauna of La Plata River basin is mostly characterized by Siluriformes and Characiformes (Reis, 2013; Reis et al., 2016), and by a decreasing gradient of species richness from the north to the south (Ringuelet, 1975; Cussac et al., 2009). Such ichthygeographical attributes agree with the results obtained in the study area, when the number of species proposed contrasts with environments located in the northern areas of the basin (Teixeira de Melo et al., 2011; Almirón et al., 2015; Bertaco et al., 2016).

Although no statistical differences were found when comparing reported data with samplings, lack of Cyprinodontiformes could result from the small body size (<5 cm) of some taxa of this group common for the area, namely Jenynsia lineata (Fowler, 1940), Cnesterodon decemmaculatus (Jenyns, 1842), and Phalloceros caudimaculatus (Hensel, 1868), not captured with the fishing nets employed. The taxa of Rivulidae family includes species that inhabit coastal temporary ponds (Alonso et al., 2018), environments not sampled in this study. The absence of Myliobatiformes in samplings can be linked to selectivity of fishing gears (Lucifora et al., 2016).

Knowledge of richness and abundance of fish species in a particular area is associated with collecting effort (Bertaco et al., 2016); freshwater RdlP has been sampled for more than five decades by several fishing procedures. Comparison of number of records results with captured/not captured species ratio, clearly evidenced that the more the species were cited in the literature the more they were found in the samplings. Indeed, species with higher values of records, such as P. lineatus (32), S. brasiliensis (14), and Odontesthes bonariensis (Valenciennes, 1835) (14), were linked not only to their occurrence and abundance in the freshwater RdlP, but also due to their importance to local and regional fisheries of the RdlP basin (Baigún et al., 2003; Colautti et al., 2009). Accordingly, these findings provide valuable insights into which species are, historically and at present, usual or unusual fish components of the study area.

The Olmstead-Tukey analysis evidenced, in agreement with the literature for the southern bank of the study area (Llompart et al., 2012; Paracampo et al., 2015, 2020), a predominance of P. valenciennis and P. maculatus. These findings were documentednot only in coastal environments and the neighboring streams of RdlP, but also in deeper environments of the riverine sector of the RdlP (García et al., 2010; CARU-CARP, 2012; 2016). It is important to note that, the target of coastal local artisanal and sport fisheries, P. lineatus, S. brasiliensis, M. obtusidens, Luciopimelodus pati (Valenciennes, 1835),and Pseudoplatystoma corruscans (Spix & Agassiz, 1829) (Baigún et al., 2003; Colautti et al., 2009) were frequent or dominant in the Olmstead-Tukey analyses. Previous research into these species showed that they displayed a seasonal migration circuit between the freshwater RdlP up to the upper main rivers of La Plata basin during southern warm and colder seasons, respectively (Baigún et al., 2003; Espinach Ros et al., 2011; Avigliano, et al., 2020). However, our results would support the hypothesis that certain individuals of these species do not migrate and remain in the area throughout the entire year cycle.

In the Rio de la Plata, detritus is the most important component in the food web of the system (Lercari et al., 2015) and detritivorous species P. lineatus plays a key role in the cycling of nutrients, organic matter, and transferring energy to higher levels (Baigún et al., 2003; Speranza et al., 2013). The CPUEw dominance of P. lineatus, in addition the presence of L. anus and H. commersoni which are strictly detritivorous-alguivorous taxa (Lujan et al., 2012), reinforce the major role of detritus and the mentioned species in the system functioning.

Among the rare species, we can mention Rhaphiodon vulpinus Spix & Agassiz, 1829 and Rhamphichthys hahni (Meinken, 1937) considered, according to Baigún et al., (2012), as “data deficiency” based on information of their population status. In this category, Cynopotamus kincaidi (Schultz, 1950), Hypostomus uruguayensis Reis, Weber & Malabarba, 1990, Hypostomus aspilogaster (Cope, 1894),Galeocharax humeralis (Valenciennes, 1834),and Pterodoras granulosus (Valenciennes, 1821) species were not even considered by those authors as species found in the RdlP.

In view of ecological features of the estuary might generate dissimilarities between the fish assemblage of the southern bank (Argentina) respect to the northern one (Uruguay) (Bessonart et al., 2021), the hierarchy structure proposed could exhibit geographical restrictions. Despite such restrains, significant gaps in freshwater species knowledge (Abell et al., 2008) and data deficiency for neotropical fishes (Baigún et al., 2012), the fish structure reported here represent a base line for future research.

Considering the negative impact experienced by South American basins as a result of the rapid anthropogenic changes introduced during the 21st century (Allan et al., 2005; Barletta et al., 2010; Baigún et al., 2012; Reis et al., 2013), the insights into the composition and structure of the fish assemblages of the freshwater RdlP provided in this study could be used as starting point to monitor and develop criteria for the management and conservation of neotropical fish species in their southernmost distribution boundary.

Acknowledgments

The authors would like to thank Roberto Jensen (ILPLA) for his assistance in data collection and field operations. The research was partially financed by PIP (CONICET, process number 0652), P-UE (22920160100049CO), and PICT 1251. This paper is a Scientific Contribution number 1196, Institute of Limnology “Dr. Raúl A. Ringuelet” (ILPLA, CCT-La Plata CONICET, UNLP).

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Authors

Tomás Maiztegui^1,2 , Ariel Hernán Paracampo², Jorge Liotta³, Eva Cabanellas², Carlos Bonetto² and Darío César Colautti²

^[1] Comisión de Investigaciones Científicas de la provincia de Buenos Aires (CIC), La Plata, Buenos Aires, Argentina. (TM) maiztegui@ilpla.edu.ar (corresponding author).

^[2] Instituto de Limnología Dr. R. Ringuelet, CONICET, Universidad Nacional de La Plata (ILPLA), Boulevard 120 y 62, CP 1900 –CC712, La Plata, Buenos Aires, Argentina. (AHP) arielp@ilpla.edu.ar, (EC) evacabanellas.89@gmail.com, (CB) bonetto@ilpla.edu.ar, (DC) colautti@ilpla.edu.ar.

^[3] Museo de Ciencias Naturales “P. A. Scasso”, San Nicolás, Buenos Aires. Dirección de Pesca Continental, Ministerio deAgroindustria, C1107AQA Ciudad Autónoma de Buenos Aires, Buenos Aires, Argentina. (JL) jorgerliotta@gmail.com.

Authors’ Contribution

Tomás Maiztegui: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Writing-original draft, Writing-review and editing.

Ariel Hernán Paracampo: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Validation, Visualization, Writing-original draft, Writing-review and editing.

Jorge Liotta: Data curation, Methodology, Supervision, Validation.

Eva Cabanellas: Methodology, Writing-original draft.

Carlos Bonetto: Writing-review and editing.

Darío César Colautti: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Writing-original draft, Writing-review and editing.

Ethical Statement

Care during collection and handling of fish for this study complied with the Buenos Aires Province (Argentina) Wildlife and Fisheries Authority guidelines and policies (Law 11,477).

Competing Interests

How to cite this article

Maiztegui T, Paracampo AH, Liotta J, Cabanellas E, Bonetto C, Colautti DC. Freshwater fishes of the Río de la Plata: current assemblage structure. Neotrop Ichthyol. 2022; 20(3):e210159. https://doi.org/10.1590/1982-0224-2021-0159

Copyright

This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

Distributed under

Creative Commons CC-BY 4.0

Diversity and Distributions Published by SBI

Accepted July 12, 2022 by Franco Teixeira de Mello

Submitted November 11, 2021

Epub October 14, 2022

Abstract​

Introduction​

Material and methods

Results​

Discussion​

Acknowledgments​

References​