Tomás Maiztegui1,2 
, 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 |
|
|
CPUEn |
CPUEw |
|||||
|
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. |
||||