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Checklist and key for the identification of fish fauna of the Uberaba River, Upper Paraná River system, Brazil
expand article infoDouglas de Castro Ribeiro, Jumma Miranda Araújo Chagas§, Mariana Ribeiro Thereza, Francisco Langeani
‡ Universidade Estadual Paulista, Rua Cristóvão Colombo, Brazil
§ Universidade Estadual Paulista, Ilha Solteira, Brazil
Open Access

Abstract

The Uberaba River is an important right-bank tributary to the Grande River, in the Upper Paraná River system, Brazil, and the main water source for the public supply of the Uberaba city, Minas Gerais state. An inventory, an identification key, and photographs of the fish species of the Uberaba River are provided, based on samples made between 2012 and 2014 at 14 sampling sites in the river system. A total of 73 species was recorded from six orders, 20 families, and 49 genera. Characiformes and Siluriformes are the most speciose orders and Characidae and Loricariidae are the most commonly recorded families. Most species are autochthonous, nine are considered allochthonous, and two species are exotic. The Uberaba River has a diverse and heterogeneous ichthyofauna, typical of rheophilic environments, with endemic species and few non-native species.

Keywords

Brazilian Cerrado, freshwater fish, Neotropical Region, rheophilic environment, threatened species

Introduction

Approximately 34,797 species of fish have been formally described worldwide (Fricke et al. 2018), and recent estimates suggest that ca. 13,000 species are partially or exclusively freshwater (Nelson 2016). The Neotropical region has a unique and diverse freshwater fish fauna (Albert and Reis 2011), with 9,100 species exclusively distributed in South America (Reis et al. 2016), an impressive number when compared to the global estimates. Approximately 43% of the Neotropical fish diversity occurs in Brazil (Buckup et al. 2007), and the Amazon and La Plata river drainages bear the largest fish diversity in South America (Langeani et al. 2007).

With geological origin dating from the Mesozoic (Neocretaceous), the La Plata River has an estimated drainage area of ca. 3 million km² across five countries, Bolivia, Brazil, Paraguay, Argentina, and Uruguay, and is the second largest drainage in South America, with the main drainages the Paraná-Paraguay drainage and Uruguay River (Albert and Reis 2011). The Upper Paraná River system is a catchment above the Sete Quedas Falls, currently flooded by the Itaipu hydroelectric dam, located at the border between Brazil, Paraguay, and Argentina. In the Brazilian portion, the Upper Paraná River system drains the states of Goiás, Minas Gerais, São Paulo, Mato Grosso do Sul, and Paraná, comprising the subsystems of the Grande, Paranaíba, Tietê, and Paranapanema rivers (Souza-Filho and Stevaux 1997; Langeani et al. 2007).

The Upper Paraná River, according to Langeani et al. (2007), harbors approximately 360 of fish species. Subsequently, Fagundes et al. (2015) provide 46 new records for this system. Additionally, at least 28 new species have been described since the last twenty years (e.g., Silveira et al. 2008; Martins and Langeani 2011a, 2011b; Carvalho and Langeani 2013; Serra and Langeani 2015). The increased number of species recorded in the Upper Paraná River in the last decade reflects intense sampling carried out in the region. Some authors (e.g., Langeani et al. 2007; Oyakawa and Menezes 2011) report that the Upper Paraná River is among the most well-sampled Brazilian regions, especially the São Paulo state (Oyakawa and Menezes 2011), and is one of the most impacted by dams, which considerably altered the hydrological regime and natural environments, affecting the dynamics and recruitment in fish populations (Agostinho et al. 2004). Fagundes et al. (2015) carried out intense samplings in tributaries of the Paranaíba, Araguari, and Grande rivers in the state of Minas Gerais, northwest, east, and southeast of the Triângulo Mineiro region, contributing significantly to the knowledge on local fish faunas. However, despite the recent contributions to the Upper Paraná River system, some areas were poorly sampled (e.g., south and southwest of the Triângulo Mineiro region, northeast and south parts of the Minas Gerais state, most of the Mato Grosso do Sul and Goiás states) and information on fish fauna composition and distribution is still missing.

The Uberaba River is a right-bank tributary of the Grande River, in the Upper Paraná River system, Brazil, and it is the main water source for Uberaba city in Minas Gerais state. In the driest period, the water level of the Uberaba River is very low and it is not able to be the only source of public water supply to the Uberaba city. This problem becomes worse with the intensive anthropogenic impact on the environment which results in modifications of hydrological dynamics and associated biotic structures (Candido et al. 2010; Cruz 2003; Valera et al. 2016). A dam located in the middle section of the Uberaba River, designed to capture and treat water for human consumption, significantly altered the natural characteristics and self-depuration capacity of the river (Sousa et al. 2016), even more aggravated by the high loads of raw sewage released into some river sections (Cruz 2003).

The fish fauna of the Uberaba River is only partially known, with only few sections sampled and no seasonal investigations (see SEMEA 2004; Souza et al. 2016). In this paper, we present an inventory of the fish fauna of the Uberaba River based on samples from several sections of the river system. In addition, an identification key and photographs of some species are presented.

Materials and methods

Study area

The Uberaba River catchment area is located in the southeastern region of Minas Gerais state, Brazil, center-south of the Triângulo Mineiro region, 19°30'37"S – 20°07'40"S; 47°39'2"W – 48°34'34"W (Figure 1). The Uberaba River system covers an area of approximately 2, 428.73 km² and is subordinated to the “Comitê da Bacia Hidrográfica dos Afluentes Mineiros do Baixo Rio Grande (CBH-GD)”. The Uberaba River extends for 184.90 km, with a gap of approximately 554 m, and is supplied by 86 tributaries of diverse orders along its course. Its headwaters are located east of the municipality of Uberaba-MG, a hydromorphic field along the BR-262 road, at 1,014 m of altitude. The Uberaba River discharges in the right side of the Grande River in the municipality of Planura, Minas Gerais state, at 460 m of altitude (CODAU 2005). Along its route, the Uberaba River crosses five municipalities, Uberaba (1,198.75 km²), followed by Conceição das Alagoas (643.19 km²), Veríssimo (568.65 km²), Planura (33.39 km²), and Campo Florido (4.59 km²) (IGAM 2010).

Figure 1. 

Map of the Uberaba River drainage. A Upper Paraná River system highlighted in the Neotropical region B location of the Uberaba River drainage in the Upper Paraná River system C red triangles showing the sampling sites in the Uberaba River.

The average annual precipitation in the region ranges between 1,300 mm and 1,700 mm, characterized by a rainy period of six to seven months (October to March) and the driest period (April to September) with less than 60 mm. The thermal regime is defined by an average annual temperature ranging from 20 to 24° Celsius, with a minimum of 18° C in colder months (June/July). These climatic factors characterize two major seasons in the region, one, cold and dry, between autumn and winter, and the other, hot and rainy, between spring and summer (Gomes et al. 1982).

Data

The collections were carried out between 2012 and 2014 in 14 sampling sites (Figures 1, 2; Table 1) along the entire system. Permission for collecting was provided by IEF / DPBIO / GPFF No.44551-1156-2011. The samplings were performed both during the daytime and nighttime, using gill nets (2.5 to 120 mm mesh), dip nets (0.5 mm mesh), seines (1.5 mm mesh), and cast nets (2.5 to 100 mm mesh sizes). After sampling, the specimens were anesthetized in a solution containing 100 mg of eugenol by L-1 previously dissolved in 100% ethanol in proportion of 1:1 v/v, fixed in 10% formalin buffered with sodium phosphate (pH 7.0 and 0.2 Mol) for 24 to 72 hours, and then transferred to 70o G.L. ethanol.

Figure 2. 

Sampling sites in the Uberaba River, Upper Paraná River system, Brazil. Detailed description of sites in Table 1.

Table 1.

Description of sampling sites (S1 to S14) of the Uberaba River, Upper Paraná River system, Brazil.

Site Locality Coordinates Elevation Characteristics
S1 Serra do Grotão, headspring of the Uberaba River, on the margins of BR 262, Ponte Alta, MG 19.40575S, 47.405430W 1015 Lentic environment; organic sediment and sand as substrate; clear and warm water, 1 m deep; abundant aquatic plants
S2 Small stream (no name), unpaved road at BR262, tributary of Veríssimo River, Veríssimo, MG 19.39538S; 48.181390W 622 Lotic environment, medium flow; clay as substrate; shallow water, less than 80 cm deep; few marginal plants
S3 Small stream (no name), unpaved road at Mula Preta farm, tributary of the Lageado River, Uberaba, MG 19.45312S; 47.484494W 715 Medium flow stream; sand and clay as substrate; turbid water; less than 1.5 m deep; riparian vegetation and open areas
S4 Small stream (no name), into APP Vale encantado, tributary of the Saudade stream, Uberaba, MG 19.33573S; 47.534852W 901 Lentic environment; organic sediment and sand as substrate; clear and warm water; 0.5 m depth; few aquatic plants
S5 Alegria stream, unpaved road at Alegria farm, tributary of the Uberaba River, Uberaba, MG 19.40224S; 47.522022W 803 Lotic environment, medium flow; clay soil as a substrate; shallow and turbid water, 1 m depth; dense riparian forest and pasture area
S6 Small stream (no name), Rocinha farm, unpaved road at Pará Pereira Gomes road, tributary of the Lageado stream, Uberaba, MG 19.41135S; 47.542032W 778 Lotic environment, medium flow; sand and leaves as substrate; shallow and crystalline waters, 30 cm deep; riparian forest sparse
S7 Uberaba River, below of the PCH Monjolo, Veríssimo, MG 19.41466S; 48.113035W 632 Lotic environment, fast flowing, several rapids and small backwaters, basaltic rocks and sand as substrate, riparian vegetation well preserved.
S8 Uberaba River, Conceição das Alagoas, MG 19.54288S; 48.23155W 495 Lotic environment, fast flowing, several rapids and small backwaters, basaltic rocks and sand as substrate, riparian vegetation well preserved, urban effluent present.
S9 Ribeirão das Alagoas stream (or Eliezer stream), Eliezer farm, unpaved road at MG427, Conceição das Alagoas, MG 19.58451S; 48.274545W 495 Medium-flow lotic environment; sand and clay as substrate; turbid waters, 1.5 m deep; degraded area
S10 Small stream (no name), unpaved road at a sanitary landfill, tributary of the Uberaba River, Conceição das Alagoas, MG 19.55268S; 48.233689W 507 Lotic environment, low flow, clay soil as a substrate, very shallow water, less than 30 cm deep; few marginal plants, very degraded area
S11 Small stream (no name), 0.7 km at IFTM campus, affluent of the Uberaba River, Uberaba, MG 19.67431S; 47.978456W 779 Medium flow stream, gravel and basaltic rocks as substrate; crystalline waters, 1 m deep, dense riparian vegetation
S12 Uberaba River, Carijó farm, 4.5 km upstream from Gorfo waterfall, Conceição das Alagoas, MG 19.92382S; 48.404833W 490 Lotic environment, fast flow, several rapids, basaltic rocks and gravel as a substrate, well preserved riparian vegetation, urban effluent present.
S13 Ribeirão das Alagoas stream (or Eliezer stream), near the confluence with the Uberaba River, Conceição das Alagoas, MG 19.97009S; 48.384722W 506 Lotic environment, medium flow, sand and clay as substrate, large basaltic rocks, turbid water, 1 m depth, degraded riparian vegetation
S14 Small stream (no name), unpaved road at Conceição das Alagoas city, tributary of the Uberaba River, Conceição das Alagoas, MG 19.91363S; 48.375123W 516 Lotic environment, low flow, loam and sand as substrate; shallow water, 70 cm deep; many marginal grasses, degraded area

Specimens were identified using appropriate literature sources (e.g., Langeani et al. 2007; Langeani and Rêgo 2014; Castro et al. 2004; Ota et al. 2018) or by direct comparisons with specimens in museum collections. Vouchers are in the DZSJRP fish collection of the Departamento de Zoologia e Botânica do Instituto de Biociências, Letras e Ciências Exatas, Universidade Estadual Paulista 'Júlio de Mesquita Filho', São José do Rio Preto, SP, Brazil. Some groups are in need of a taxonomic revision, consequently the particle aff. (meaning “not the referred species, but very similar”) is used. The morphometric measurements were taken on the left side of the body, using a digital caliper with an accuracy of 0.01 mm. Lower-level taxonomy and species names follow Fricke et al. (2018) and suprageneric taxonomic groups are those listed in Betancur et al. (2017), except for Cynolebiidae and Bryconidae that follow van der Laan (2016). Allochthonous species are those with their origins from any other hydrographic system in South America outside the Upper Paraná River as defined above. Exotic species are those with origins from any other continent.

Results

In total, 2,722 specimens were collected and assigned to 49 genera and 73 species. The identified taxa are listed in Table 2. Most of the species in the Uberaba River are autochthonous (80.0%). Nine species (12.3%) have been recognized as allochthonous (Galeocharax gulo (Cope), Metynnis lippincottianus (Cope), Knodus aff. moenkhausii (Eigenmann & Kennedy), Hoplerythrinus unitaeniatus (Spix & Agassiz), Gymnotus inaequilabiatus (Valenciennes), Trichomycterus brasiliensis Lütken, Megalechis thoracata (Valenciennes), Poecilia reticulata Peters, and Cichla piquiti Kullander & Ferreira), and only two (2.7%) species are exotic (Coptodon rendalli (Boulenger) and Oreochromis niloticus (Linnaeus)). Six orders were recognized, of which Characiformes and Siluriformes were the most representative (90.3%), with eight families and 33 species for the former and five families and 27 species for the latter. Gymnotiformes (two families and three spp.), Cichliformes (one family and seven spp.), Cyprinodontiformes (two families and three spp.), and Synbranchiformes (one sp.) together represent 9.7% of the groups collected (Figure 3). Characidae (48.8%) and Loricariidae (16.8%) correspond to the most abundant families (Figure 4) and occur in the entire river system. The species richness suggested a longitudinal gradient, with more species in the lower reaches whereas in the upper reaches the richness does not exceed ten species (Figure 5 and Table 3). The loricariids are mainly represented by Hypostomus species, up to 92% of the total loricariid number. The most abundant species is Knodus aff. moenkhausii with 507 collected specimens comprising 38% of all characiform species. All other species have already been recorded in the Upper Paraná River.

Figure 3. 

Species richness for each fish order collected in Uberaba River, Upper Paraná River system, Brazil.

Figure 4. 

Species richness of each fish family collected in Uberaba River, Upper Paraná River system, Brazil.

Figure 5. 

Species richness along longitudinal gradient in Uberaba River, Upper Paraná River system, Brazil. Circle diameter corresponds to species richness.

Table 2.

List of fish species from the Uberaba River, Upper Paraná River system, Brazil. Vouchers and origin/status are provided.

Taxa Voucher Origin
CHARACIFORMES
Anostomidae
1 Leporinus amblyrhynchus Garavello & Britski, 1987 DZSJRP15809 Autochthonous
2 Leporinus friderici (Bloch, 1794) uncataloged Autochthonous
3 Leporinus octofasciatus Steindachner, 1915 DZSJRP16097 Autochthonous
4 Leporinus striatus Kner, 1858 DZSJRP21396 Autochthonous
5 Schizodon nasutus Kner, 1858 DZSJRP21388 Autochthonous
Bryconidae
6 Brycon nattereri Günther, 1864 DZSJRP17489 Autochthonous/VU
Characidae
7 Astyanax bockmanni Vari & Castro, 2007 DZSJRP15819 Autochthonous
8 Astyanax aff. fasciatus (Cuvier, 1819) DZSJRP15818 Autochthonous
9 Astyanax lacustris (Lütken, 1875) DZSJRP21399 Autochthonous
10 Astyanax aff. paranae Eigenmann, 1914 DZSJRP17486 Autochthonous
11 Astyanax paranae Eigenmann, 1914 DZSJRP15823 Autochthonous
12 Bryconamericus turiuba Langeani et al., 2005 DZSJRP05533 Autochthonous
13 Galeocharax gulo (Cope, 1870) DZSJRP16096 Allochthonous
14 Hasemania uberaba Serra & Langeani, 2015 DZSJRP18781 Autochthonous
15 Hyphessobrycon uaiso Carvalho & Langeani, 2013 DZSJRP18783 Autochthonous
16 Knodus aff. moenkhausii (Eigenmann & Kennedy, 1903) DZSJRP15825 Allochthonous
17 Oligosarcus pintoi Campos, 1945 DZSJRP05553 Autochthonous
18 Piabarchus stramineus (Eigenmann, 1908) DZSJRP21383 Autochthonous
19 Piabina argentea Reinhardt, 1867 DZSJRP17487 Autochthonous
Serrasalmidae
20 Metynnis lippincottianus (Cope, 1870) DZSJRP21397 Allochthonous
21 Myloplus tiete (Eigenmann & Norris, 1900) DZSJRP21398 Autochthonous/EN
22 Serrasalmus maculatus Kner, 1858 DZSJRP21386 Autochthonous
Curimatidae
23 Steindachnerina insculpta (Fernández-Yépez, 1948) DZSJRP15812 Autochthonous
Erythrinidae
24 Hoplerythrinus unitaeniatus (Spix & Agassiz, 1829) DZSJRP21402 Allochthonous
25 Hoplias intermedius (Günther, 1864) DZSJRP21389 Autochthonous
26 Hoplias aff. malabaricus (Bloch, 1794) DZSJRP10546 Autochthonous
Parodontidae
27 Apareiodon affinis (Steindachner, 1879) DZSJRP21391 Autochthonous
28 Apareiodon ibitiensis Campos, 1944 DZSJRP15813 Autochthonous
29 Apareiodon piracicabae (Eigenmann, 1907) DZSJRP16100 Autochthonous
30 Parodon nasus Kner, 1859 DZSJRP21400 Autochthonous
Crenuchidae
31 Characidium aff. zebra Eigenmann, 1909 DZSJRP17484 Autochthonous
32 Crenuchidae (undescribed genus and species) DZSJRP15806 Autochthonous
Prochilodontidae
33 Prochilodus lineatus (Valenciennes, 1837) DZSJRP21385 Autochthonous
GYMNOTIFORMES
Sternopygidae
34 Eigenmannia trilineata López & Castello, 1966 DZSJRP21392 Autochthonous
Gymnotidae
35 Gymnotus inaequilabiatus (Valenciennes, 1839) uncataloged Allochthonous
36 Gymnotus sylvius Albert & Fernandes-Matioli, 1999 DZSJRP16101 Autochthonous
SILURIFORMES
Callichthyidae
37 Aspidoras fuscoguttatus Nijssen & Isbrücker, 1976 DZSJRP18785 Autochthonous
38 Corydoras difluviatilis Britto & Castro, 2002 DZSJRP15824 Autochthonous
39 Megalechis thoracata (Valenciennes, 1840) DZSJRP21106 Allochthonous
Heptapteridae
40 Imparfinis borodini Mees & Cala, 1989 DZSJRP17488 Autochthonous
41 Pimelodella avanhandavae Eigenmann, 1917 DZSJRP21105 Autochthonous
42 Rhamdia quelen (Quoy & Gaimard, 1824) DZSJRP16799 Autochthonous
43 Rhamdiopsis sp. DZSJRP15817 Autochthonous
Loricariidae
44 Curculionichthys insperatus (Britski & Garavello, 2003) DZSJRP21120 Autochthonous
45 Hypostomus albopunctatus (Regan, 1908) DZSJRP21390 Autochthonous
46 Hypostomus ancistroides (Ihering, 1911) DZSJRP15810 Autochthonous
47 Hypostomus butantanis (Ihering, 1911) DZSJRP16098 Autochthonous
48 Hypostomus fluviatilis (Schubart, 1964) DZSJRP21114 Autochthonous
49 Hypostomus aff. hermanni (Ihering, 1905) DZSJRP21107 Autochthonous
50 Hypostomus margaritifer (Regan, 1908) DZSJRP02107 Autochthonous
51 Hypostomus nigromaculatus (Schubart, 1964) DZSJRP16103 Autochthonous
52 Hypostomus aff. paulinus (Ihering, 1905) DZSJRP21108 Autochthonous
53 Hypostomus regani (Ihering, 1905) DZSJRP21124 Autochthonous
54 Hypostomus strigaticeps (Regan, 1908) DZSJRP21125 Autochthonous
55 Hypostomus topavae (Godoy, 1969) DZSJRP21098 Autochthonous
56 Loricaria lentiginosa Isbrücker, 1979 uncataloged Autochthonous
57 Microlepdogaster dimorpha Martins & Langeani, 2012 DZSJRP18784 Autochthonous
58 Proloricaria prolixa (Isbrücker & Nijssen, 1978) DZSJRP16102 Autochthonous
59 Rineloricaria latirostris (Boulenger, 1900) DZSJRP15811 Autochthonous
Trichomycteridae
60 Trichomycterus brasiliensis Lütken, 1874 DZSJRP21116 Allochthonous
61 Trichomycterus candidus (Miranda-Ribeiro, 1949) DZSJRP15820 Autochthonous
Auchenipteridae
62 Tatia neivai (Ihering, 1930) DZSJRP21111 Autochthonous
CYPRINODONTIFORMES
Cynolebiidae
63 Melanorivulus giarettai Costa, 2008 DZSJRP18782 Autochthonous
Poeciliidae
64 Phalloceros harpagos Lucinda, 2008 DZSJRP17485 Autochthonous
65 Poecillia reticulata Peters, 1859 DZSJRP17483 Allochthonous
CICHLIFORMES
Cichlidae
66 Cichla piquiti Kullander & Ferreira, 2006 DZSJRP21401 Allochthonous
67 Cichlasoma paranaense Kullander, 1983 DZSJRP21394 Autochthonous
68 Coptodon rendalli (Boulenger, 1897) DZSJRP05549 Exotic
69 Crenicichla britskii Kullander, 1982 DZSJRP21393 Autochthonous
70 Crenicichla jaguarensis Haseman, 1911 DZSJRP21387 Autochthonous
71 Geophagus brasiliensis (Quoy & Gaimard, 1824) DZSJRP21395 Autochthonous
73 Oreochromis niloticus (Linnaeus, 1758) uncataloged Exotic
SYNBRANCHIFORMES
Synbranchidae
73 Synbranchus marmoratus Bloch, 1795 DZSJRP21384 Autochthonous
Table 3.

Species collected (X) in each site (S1 to S14) of the Uberaba River, Upper Paraná River system, Brazil.

Species Sites
S1 S2 S3 S4 S5 S6 S7 S8 S9 S10 S11 S12 S13 S14
Apareiodon affinis X X X X X
Apareiodon ibitiensis X X X X X
Apareiodon piracicabae X X X X
Aspidoras fuscoguttatus X
Astyanax bockmanni X X X X
Astyanax aff. fasciatus X X X X X
Astyanax lacustris X X X X X X X
Astyanax paranae X X X X
Astyanax aff. paranae X
Brycon nattereri X X
Bryconamericus turiuba X X
Characidium aff. zebra X X
Cichla piquiti X
Cichlasoma paranaense X X X X
Coptodon rendalli X X
Corydoras difluviatilis X X
Crenicichla britskii X
Crenicichla jaguarensis X
Crenuchidae (undescribed genus and species) X
Curculionichthys insperatus X
Eigenmannia trilineata X
Galeocharax gulo X
Geophagus brasiliensis X X X X
Gymnotus inaequilabiatus X
Gymnotus sylvius X X X
Hasemania uberaba X
Hoplerythrinus unitaeniatus X X
Hoplias intermedius X X
Hoplias aff. malabaricus X X X X
Hyphessobrycon uaiso X X X
Hypostomus albopunctatus X X X
Hypostomus ancistroides X X X X X
Hypostomus butantanis X X
Hypostomus fluviatilis X
Hypostomus aff. hermani X X
Hypostomus margaritifer X X
Hypostomus nigromaculatus X X X X X
Hypostomus aff. paulinus X X X
Hypostomus regani X X
Hypostomus strigaticeps X X X
Hypostomus topavae X X X X X
Imparfinis borodini X
Knodus aff. moenkhausii X X X
Leporinus amblyrhynchus X X X
Leporinus friderici X X X X
Leporinus octofasciatus X X X
Leporinus striatus X
Loricaria lentiginosa X X
Megalechis thoracata X X
Melanorivulus giarettai X X
Metynnis lippincottianus X
Microlepdogaster dimorpha X
Myloplus tiete X X
Oligosarcus pintoi X
Oreochromis niloticus X X X
Parodon nasus X X X X X
Phalloceros harpagos X
Piabarchus stramineus X X
Piabina argentea X X
Poecillia reticulata X X X
Prochilodus lineatus X
Proloricaria prolixa X X
Rhamdia quelen X X X X
Rhamdiopsis sp. X
Rineloricaria latirostris X X
Schizodon nasutus X
Serrasalmus maculatus X
Steindachnerina insculpta X X X X X
Synbranchus marmoratus X X X
Tatia neivai X X
Trichomycterus brasiliensis X
Trichomycterus candidus X
Species richness 4 2 2 3 4 4 24 53 15 5 2 31 24 2

Key to fish species of the Uberaba River drainage

1 Single mid-ventral gill opening; eel-shaped body Synbranchus marmoratus
Two laterally located gill openings; not eel-shaped body 2
2 Dorsal and pelvic fins absent; anal-fin rays more than 100 3
Dorsal fin present; pelvic fin commonly present; anal-fin rays up to 50 5
3 Body uniformly clear with relatively inconspicuous longitudinal stripes; anal fin not reaching the tail end; terminal mouth, both jaws approximately equal Eigenmannia trilineata
Body dark with clear transverse bands; anal fin extending to the tail end; prognathous, lower jaw longer than upper jaw 4
4 Obliquely-oriented dark transversal bars fragmented, forming a pattern of irregular spots; anal-fin posterior membrane striped Gymnotus inaequilabiatus
Obliquely-oriented dark transversal bars not fragmented; anal-fin posterior region darkly pigmented or translucent Gymnotus sylvius
5 Body naked or covered by bony plates 6
Body covered by scales 30
6 Body covered by bony plates, at least partially 7
Body covered by thick skin; bony plates absent 24
7 Mouth forming a ventral oral disk; bony plates rows on flanks 3–5 8
Mouth not forming ventral oral disk, with terminal or subterminal opening; bony plates rows on flank 2 22
8 Adipose fin absent 9
Adipose fin present 12
9 Caudal peduncle very elongate and depressed 10
Caudal peduncle rounded or elliptical in cross-section 13
10 Lips with small papillae, occasionally with short, thick, non-filamentous projections Rineloricaria latirostris
Lips fringed, with filamentous projections 11
11 Head with dark brown spots, much smaller than the eye diameter Loricaria lentiginosa
Head light brown without spots Proloricaria prolixa
12 Scapular bridge fully exposed; well-developed and pointed odontodes on the anterior portion of the snout Curculionichthys insperatus
Scapular bridge exposed only laterally; small and spatulate odontodes on the anterior portion of the snout Microlepidogaster dimorpha
13 Body light with dark spots 14
Body dark with light spots or vermiculations 17
14 Lateral keels on body present (three rows), with hypertrophied odontodes Hypostomus ancistroides
Lateral keels on body absent 15
15 Pectoral-fin spine claviform, with well-developed odontodes on distal portion; eyes small, 6–6.5 × in head length Hypostomus nigromaculatus
Pectoral-fin spine not claviform, with subequal odontodes along entire spine; eyes large, 3.5–5 × in head length Hypostomus fluviatilis
16 Abdomen completely covered by plates; dentary angle more than 60°; bony plates between dorsal and adipose fins 5 pairs Hypostomus topavae
Abdomen without plates on pelvic-fin region; dentary angle approximately 45°; bony plates between dorsal and adipose fins 4 pairs Hypostomu s aff. hermani
17 Pectoral-fin spine equal to or shorter than pelvic-fin spine Hypostomus albopunctatus
Pectoral-fin spine longer than pelvic-fin spine 18
18 Premaxillary and dentary with short and sturdy teeth (18–32), arranged in obtuse angle 19
Premaxillary and dentary with long and thin teeth (more than 35), arranged in acute angle 20
19 Body and fins with light spots, aligned longitudinally, but not forming continuous line Hypostomus margaritifer
Head and fins with light vermiculations, with four longitudinal yellow lines on flank, from dorsal fin to caudal-fin base Hypostomus butantanis
20 Pectoral girdle covered with large plates; bony plates between anal and caudal fins 10 or 11; dentary teeth more than 140 Hypostomus aff. paulinus
Pectoral girdle covered with very small plates or skin; bony plates between anal and caudal fins 12 or 13; dentary teeth up to 130 21
21 Mid-lateral plates series 28 or 29; snout-operculum distance greater than the width of the lips; dorsal fin large, reaching adipose fin; premaxillary and dentary teeth more than 65 Hypostomus regani
Mid-lateral plates series 25 or 26; snout-operculum distance equal to width of the lips; dorsal fin of moderate size, distant from adipose fin; premaxillary and dentary teeth up to 60 Hypostomus strigaticeps
22 Mental barbels absent; jaws teeth present; nuchal plate covered by skin; caudal fin truncated Megalechis thoracata
Mental barbels present; jaws teeth absent; nuchal plate exposed; caudal fin forked 23
23 Supraoccipital long and reaching the nuchal plate; pectoral-fin rays anterior portion without posterior bone lamellae Corydoras difluviatilis
Supraoccipital short, not reaching the nuchal plate; pectoral-fin rays anterior portion with posterior bone lamellae (more evident in the first rays) Aspidoras fuscoguttatus
24 Operculum and preoperculum with odontodes; dorsal-fin origin situated posterior the middle of the body 25
Operculum and preoperculum without odontodes; dorsal-fin origin situated approximately at the middle of the body 26
25 Pelvic fin present Trichomycterus brasiliensis
Pelvic fin absent Trichomycterus candidus
26 Adipose fin short, shorter than anal fin length; nuchal plate reaching the posterior portion of head Tatia neivai
Adipose fin long, approximately 2 × anal fin length; nuchal plate not reaching the posterior portion of head 27
27 Body very elongate, depth contained 8.0 × in standard length; 4 dark brown dorsal transverse bands (first at vertical passing at pectoral fin, second at vertical passing anterior portion of dorsal-fin base, third at vertical passing at last third of dorsal-fin base, and the last one at vertical passing at adipose-fin origin); eyes dorsally placed Imparfinis borodini
Body short, depth contained up to 6.0 × in standard length; dark brown dorsal transverse bands absent; eyes laterally placed 28
28 Body uniformly clear; longitudinal black stripe on flank present; maxillary barbels long, and reaching or surpassing the anal-fin origin Pimelodella avanhadavae
Body with small dark spots or irregular vermiculations; longitudinal black stripe on flank absent; maxillary barbels short, never reaching the anal-fin origin 29
29 Anal-fin rays up to 12; eyes large, approximately 5 × head length Rhamdia quellen
Anal-fin rays more than 15; eyes small, more than 7.5 × head length Rhamdiopsis sp.
30 Dorsal and anal fins anterior rays modified into spines; pelvic fin in thoracic position, below of pectoral fin; lateral line divided into 2 branches, 1 anterior, near the base of the dorsal fin and another posterior, along the middle portion of the body and caudal peduncle; ctenoid scales 31
Dorsal and anal fins anterior rays not modified into spines; pelvic fin posteriorly located, close to anal fin; lateral line not divided into 2 branches; cycloid or spinoid scales 37
31 Dorsal-fin spines separate from soft rays by notch Cichla piquiti
Dorsal-fin spines not separate from soft rays by notch 32
32 Body elongate (fusiform), 3.6–5.2 × in standard length; preoperculum posterior margin serrated 33
Body deep, more than 3.5 × in standard length; preoperculum posterior margin smooth 34
33 Scales in longitudinal series 33–40; flank with black transverse bands; dorsal fin with XVI + 14 or 15 rays; anal fin with III + 9 or 10 rays; black humeral blotch present Crenicichla britskii
Scales in longitudinal series 41–50; flank without black transverse bands (crossing the longitudinal stripe); dorsal fin with XIX–XXI + 10–12 rays; anal fin with III + 7 or 8 rays; black humeral blotch absent Crenicichla jaguarensis
34 Anterior lateral line with 19 or fewer scales; scales in longitudinal series 22–27; black lateral spot present 35
Anterior lateral line with 20 or more scales; scales in longitudinal series 28–35; black lateral spot absent 36
35 Posterior lateral line with 10–14 scales; scales in longitudinal series 24–27; dorsal fin with XV or XVI + 10–13 rays; black lateral spot on flank larger than the eye diameter Geophagus brasiliensis
Posterior lateral line with 5–8 scales; scales in longitudinal series 22 or 23; dorsal fin with XIII or XV + 10–15 rays; black lateral spot approximately equal than the eye diameter Cichlasoma paranaense
36 Scales in transverse series above the lateral line 3 or 3½; gill rakers in inferior branch of the first branchial arch 18 or more Oreochromis niloticus
Scales in transverse series above the lateral line 2 or 2½; gill rakers in inferior branch of the first branchial arch 15 or fewer Coptodon rendalli
37 Top of head covered by scales; upper jaw protractile 38
Top of head not covered by scales; upper jaw non-protractile 40
38 Dorsal fin closer to caudal fin than to middle of body; gonopodium absent Melanorivulus giarettai
Dorsal fin at middle of body; gonopodium present 39
39 Males with intense colored spots in life, black when preserved; females without spots; gonopodium with moderate size (3.2–3.6 × in standard length), with terminal portion almost straight Poecilia reticulata
Males and females with vertically elongate black spot on medium portion of flank; gonopodium long (2.6–3.1 × in standard length), with terminal portion trifid and ventrally oriented Phalloceros harpagos
40 Teeth absent in adults Steindachnerina insculpta
Teeth present in all life stages 41
41 Teeth small, numerous and depressibly implanted in the lips Prochilodus lineatus
Teeth well-developed, non-depressibly implanted in the jaw bones 42
42 Body fusiform or moderately compressed laterally; abdominal serrae absent 43
Body very compressed laterally; abdominal serrae present 70
43 Teeth incisiform (rabbit-like), truncated or cuspidate, premaxillary and dentary with 3 teeth each, premaxillary with 3 and dentary with 3 or 4 teeth, or premaxillary and dentary with 4 teeth each 44
Teeth conical or multicuspid, no incisiform; teeth number variable, but not as above 48
44 Teeth cuspidate; flank silver in life, spots or bands absent; a conspicuous, horizontally elongate black spot at end of caudal peduncle extending to the median caudal-fin rays Schizodon nasutus
Teeth truncated; body with large black spots or longitudinal stripes; horizontally elongate black spot on end of caudal peduncle absent 45
45 Premaxillary and dentary with 4 teeth each; 3 large black spots on flank (first bellow dorsal fin, second above the anal-fin base and third at the end of caudal peduncle Leporinus friderici
Premaxillary with 3 teeth; dentary with 3 or 4 teeth, body with longitudinal black stripes or transverse bars, large black spots on flank absent 46
46 Premaxillary and dentary with 3 teeth each; black longitudinal stripe on flank present; dorsal dark transverse bars (but not reaching the longitudinal stripe) 10 or more; subterminal mouth; prominent snout Leporinus amblyrhynchus
Premaxillary with 3 teeth; dentary with 4 teeth; black dorsal transverse bars absent; terminal or subterminal mouth; non-prominent snout 47
47 Body elongate, depth 4.1 × in standard length; four longitudinal black stripes on flank; fins usually hyaline or slightly red Leporinus striatus
Body deep, depth 3.2 × in standard length; eight black transverse bars on flank; fins yellow, orange or red in life Leporinus octofasciatus
48 Premaxillary teeth in 1 row 49
Premaxillary teeth in 2 or more rows 57
49 Adipose fin absent; posterodorsal portion of head with straight margin; caudal fin rounded or truncate 50
Adipose fin usually present; posterodorsal portion of head convex or with a posterior projection; caudal fin forked or emarginate 52
50 Dorsal-fin rays up to 11; pectoral, pelvic and anal fins without dark brown stripes; teeth canine on maxillary absent Hoplerythrinus unitaeniatus
Dorsal-fin rays more than 12; pectoral, pelvic and anal fins with dark brown stripes; teeth canine on maxillary present 51
51 Medial margin of dentary bones parallel in ventral view; denticles on tongue absent Hoplias intermedius
Medial margin of dentary bones converging towards the symphysis in ventral view; denticles on tongue present Hoplias aff. malabaricus
52 Teeth on anterior portion of dentary absent; lower jaw anterior portion straight 53
Teeth on anterior portion of dentary present; lower jaw anterior portion rounded 56
53 Dentary teeth present Parodon nasus
Dentary teeth absent 54
54 Black lateral stripe with broad projections above and below, giving a zig-zag appearance; body greenish in life Apareiodon ibitiensis
Black lateral stripe without broad projections above and below; 6–8 transverse, rectangular or triangular black thin bars above; body silver in life 55
55 Scales in pre-anal series 29 or fewer; premaxillary teeth cusps up to 12 Apareiodon piracicabae
Scales in pre-anal series 29½ or more; premaxillary teeth cusps 12–15 Apareiodon affinis
56 Adipose fin absent; pectoral-fin unbranched rays 10–13; principal caudal-fin rays 16 Crenuchidae (undescribed genus and species)
Adipose fin present; pectoral-fin unbranched rays 3; principal caudal-fin rays 18 or 19 Characidium aff. zebra
57 Premaxillary teeth in three rows; teeth conical in the symphysis region present Brycon nattereri
Premaxillary teeth in two rows; teeth conical in the symphysis region absent 58
58 Teeth on the palate present Oligosarcus pintoi
Teeth on the palate absent 59
59 Anal-fin branched rays more than 30; spinoid scales Galeocharax gulo
Anal-fin branched rays up to 29; cycloid scales 60
60 Lateral line incomplete 61
Lateral line complete 62
61 Adipose fin present Hyphessobrycon uaiso
Adipose fin absent Hasemania uberaba
62 Internal series of premaxillary with 4 teeth; body relatively elongate, depth 3.0–4.2 × in standard length 63
Internal series of premaxillary with 5 teeth; body relatively deep, depth 1.8–3.6 × in standard length 66
63 Upper jaw projecting anteriorly; premaxillary teeth misaligned Piabina argentea
Upper and lower jaws of equal size; premaxillary teeth aligned 64
64 Supraorbital groove present; caudal-fin lobes covered by small scales Knodus aff. moenkhausii
Supraorbital groove absent; scales only at the caudal-fin base 65
65 Dorsal stripe broad, extending from the supraoccipital crest to the caudal-fin base, with a gap at the region of the adipose fin; humeral spot conspicuous Bryconamericus turiuba
Dorsal stripe narrow, continuous, extending from the supraoccipital crest to the caudal-fin base; humeral spot inconspicuous or absent Piabarchus stramineus
66 Maxillary teeth absent; humeral spot clearly defined, horizontally elongate associated with two diffuse vertical black stripes; fins yellow in life Astyanax lacustris
Maxillary teeth present; humeral spot absent or inconspicuous; fins orange or red in life 67
67 Flank with a silvery longitudinal stripe; scales on abdomen without chromatophores on distal portion Astyanax aff. fasciatus
Flank without silvery longitudinal stripe; scales on abdomen with black chromatophores on distal portion 68
68 Body relatively deep, up to 3.0 × in standard length; anal-fin rays 22 or more Astyanax bockmanni
Body relatively elongate, more than 3.1 × in standard length; anal-fin rays 20 or fewer 69
69 Eye with light iris, silver in life; pelvic-fin tip reaching anal fin Astyanax aff. paranae
Eye with dark iris, gold or brown in life; pelvic-fin tip not reaching anal fin Astyanax paranae
70 Teeth tricuspid present; premaxilla and dentary teeth in 1 row Serrasalmus maculatus
Teeth tricuspid absent; premaxilla and dentary teeth in 2 rows (the inner dentary row represented by 2 small conical teeth 71
71 Adipose-fin base longer than taller; dorsal-fin rays 20 or fewer; pre-dorsal spine present Metynnis lippincottianus
Adipose-fin base taller than longer; dorsal-fin rays 20 or more; pre-dorsal spine absent Myloplus tiete

Discussion

The diversity recorded in the Uberaba River (73) is slightly greater than in similar tributaries of the Grande River in São Paulo state, in which 64 species have been recorded in the tributaries of the Pardo, Turvo, and Sapucaí rivers (Castro et al. 2004). Our data increase the number of species previously recorded for the Uberaba River by 44, which corresponds to an increase of 150% of the species referred so far in the region (see more details in SEMEA 2004; Souza et al. 2016). However, these figures may reflect the differences in sampling methods used by us and the previous authors as well as a larger area investigated in this study. Estimates of species richness and diversity considerably depend on methods used as discussed by Oliveira et al. (2014).

The number of species (73) recorded in the Uberaba River comprises ca. 19% of the total species number known in the Upper Paraná River system when compared to the data in Langeani et al. (2007). The ichthyofauna of the Uberaba River is composed mainly of autochthonous species, few allochthonous species and only two exotic species. The autochthonous origin of some of these species in the Upper Parana River still needs further research. For example, the scarcity of data on the origin or taxonomic status of some putative species such as Knodus aff. moenkhausii, Trichomycterus brasiliensis or Megalechis thoracata, does not allow to reasonably hypothesize on their origin.

Some species recorded in the Uberaba River potentially correspond to new species and some considerations are provided. Astyanax fasciatus (Cuvier) is described for the São Francisco River basin and it is widely distributed in the Paraná-Paraguay drainage and coastal drainages of eastern of Brazil. However, based on the definitions by Eigenmann (1921) it is possible to infer the existence of a "A. fasciatus species complex" in the Paraná-Paraguay and other coastal drainages. Thus, the name A. fasciatus should be used strictly for the São Francisco River lineage (Melo and Buckup 2006). In the La Plata drainage, the Hoplias malabaricus species group is constantly corroborated by morphological, cytogenetic and molecular evidence, and a recognition and taxonomic delineating of new entities is currently in progress (Rosso et al. 2018). Additionally, the nominal species name Hoplias malabaricus (Bloch) should be applied exclusively to the Guiana shield lineage (Rosso et al. 2018). Similarly, some authors (see Buckup 1992) suggest that populations of Characidium zebra Eigenmann throughout South America represent more than one species. Characidium zebra was described in tributaries of the Branco River (Negro River system) in the Amazon. Recent evidence suggests that C. zebra populations in the San Francisco and Paraná rivers correspond to the same species (Serrano et al. 2018) distinct from the C. zebra populations of the Amazon drainage.

Figure 6. 

Characiformes collected in the Uberaba River. 1 Leporinus amblyrhynchus 2 Steindachnerina insculpta 3 Leporinus striatus 4 Leporinus friderici (uncataloged) 5 Leporinus octofasciatus 6 Hoplerythrinus unitaeniatus 7 Schizodon nasutus 8 Prochilodus lineatus 9 Hoplias intermedius 10 Hoplias aff. malabaricus 11 Brycon nattereri 12 Apareiodon piracicabae 13 Galeocharax gulo 14 Apareiodon affinis 15 Apareiodon ibitiensis 16 Astyanax lacustris 17 Astyanax paranae 18 Astyanax aff. paranae 19 Parodon nasus 20 Metynnis lippincottianus 21 Astyanax aff. fasciatus 22 Astyanax bockmanni 23 Bryconamericus turiuba 24 Myloplus tiete 25 Hasemania uberaba 26 Hyphessobrycon uaiso 27 Oligosarcus pintoi 28 Knodus aff. moenkhausii 29 Piabarchus stramineus 30 Piabina argentea 31 Serrasalmus maculatus 32 Crenuchidae (undescribed genus and species) and 33 Characidium aff. zebra. Photographs are of specimens presented in Table 2. Scale bar: 10 mm.

Figure 7. 

Siluriformes (Loricariidae absent), Gymnotiformes, Cichliformes, Cyprinodontiformes, and Synbrachiformes collected in the Uberaba River. 1 Megalechis thoracata 2 Tatia neivai 3 Trichomycterus candidus 4 Trichomycterus brasiliensis 5 Imparfinis borodini 6 Rhamdiopsis sp. 7 Corydoras difluviatilis 8 Aspidoras fuscoguttatus 9 Rhamdia quelen 10 Pimelodella avanhadavae 11 Eigenmannia trilineata 12 Gymnotus sylvius 13 Gymnotus inaequilabiatus (uncataloged) 14 Crenicichla jaguarensis 15 Crenicichla britskii 16 Cichla piquiti 17 Geophagus brasiliensis 18 Coptodon rendalli 19 Oreochromis niloticus (uncataloged) 20 Cichlasoma paranaense 21 Phalloceros harpagos (female above and male below) 22 Poecillia reticulata (female above and male below) 23 Melanorivulus giarettai (male above and female below) and 24 Synbranchus marmoratus. Photographs are of specimens presented in Table 2. Scale bar: 10 mm.

Astyanax aff. paranae Eigenmann collected from the Uberaba River may represent a distinct species in the complex “Astyanax scabripinnis species complex” sensu Moreira-Filho and Bertollo (1991), a group with an underestimated diversity (Bertaco and Malabarba 2001) as it differs by a number of features (e.g., eye coloration and some measurements). Knodus moenkhausii (Eigenmann & Kennedy) was described from the Arroyo Trementina in the Paraguay River system. The specimens from the Upper Paraná River and identified so far as K. moenkhausii apparently represents an undescribed species (F. R. Carvalho pers. comm.).

The taxonomic boundaries of the Hypostomus species are unclear. Some species of the genus Hypostomus are highly variable morphologically and widely distributed. In addition, some important diagnostic characters, such as color pattern, cannot be seen at present in type specimens collected more than 100 years ago, making identification of the species difficult (Zawadzki et al. 2004). For example, Hypostomus hermanni Ihering is widely distributed within the Upper Paraná River system. A comparison of the specimens collected in the Uberaba River with specimens from other locations revealed a discrepancy in some meristic and color traits. The Uberaba specimens are especially different from specimens from the Piracicaba River, the type locality of H. hermanni. It has been also shown that different populations of Hypostomus paulinus (Ihering) are effectively reproductively isolated and characterized by a high degree of inbreeding (Zawadzki et al. 2004).

The occurrence of Metynnis lippincottianus may be a result of accidental introduction (Ota 2015). Among the allochthonous species, Poecilia reticulata was introduced to control mosquito larvae (Ota et al. 2018). Cichla piquiti was probably introduced for sport fishing (Langeani et al. 2007; Ota et al. 2018), and Gymnotus inaequilabiatus originally from the Lower Paraná River, Paraguay and Uruguay rivers (Maxime and Albert 2014), colonized the upper reaches of the Paraná River after the construction of the Itaipu hydroelectric dam in the 1980s. Ota et al. (2018) suggested that the occurrence of Hoplerythrinus unitaeniatus in the Upper Paraná River can be associated with its introduction as a live bait or after inundation of the Sete Quedas Falls after the construction of the Itaipu dam. Galeocharax gulo is widely distributed in almost all Upper Amazon River systems, also in the Orinoco, Oyapok, Araguaia-Tocantins, and Paraná rivers (Giovannetti et al. 2017). The occurrence of this species in the Upper Paraná system may be a result of natural dispersion. Coptodon rendalli and O. niloticus probably represent results of escapes from fish farms (Langeani et al. 2007; Ota et al. 2018) and the populations of both species are probably established in the region as they have been regularly registered since long ago. Finally, Souza et al. (2016) report the occurrence of Cyphocharax nagelii (Steindachner) and Steindachnerina brevipinna (Eigenmann & Eigenmann) in the system, but we could not confirm these data and refrained from including them in the species list.

New taxa have been described from the Uberaba River system over the past decade, e.g., Hasemania uberaba (Serra and Langeani 2015), Hyphessobrycon uaiso (Carvalho and Langeani 2013), and Microlepidogaster dimorpha (Martins and Langeani 2011). These newly described species are only known from their type localities or from a few localities corroborating several examples of endemism in the Upper Paraná River, previously indicated by some authors (e.g., Langeani et al. 2007). This clearly demonstrates the importance of inventories and consequent conservation measures. Two species registered in the Uberaba River are definitely threatened: Brycon nattereri Günther and Myloplus tiete (Eigenmann & Norris) are assigned to “Vulnerable” (VU) and “Endangered” (EN) respectively, on the IBAMA Red List of Endangered Species (ICMBio 2015). The main threats to the local fauna are related to changes in hydrological cycles and the loss of riparian vegetation, as well as overexploitation of natural stocks (Lima et al. 2008; Lima et al. 2015). In addition, the presence of migratory rheophilic species such as Prochilodus lineatus (Valenciennes), Leporinus friderici (Bloch), B. nattereri, and M. tiete, is because these species use local resources, at least partially, to complete their life cycle, as suggested by Carolsfeld et al. (2003). Considering all the factors discussed above, the Uberaba River contains a diverse and heterogeneous fish fauna, with two endemic species, H. uberaba and an undescribed crenuchid (a description is in the process by Ribeiro et al.) and a low number of allochthonous and exotic species. The Uberaba River has undergone several anthropogenic actions over the last decades, such as the increase of the area destined to grazing, resulting in only 17.7% of native vegetation remains (Valle-Junior et al. 2010) and the reduction of the lotic environments due to damming. The impact of human-induced environmental change is dramatic on the structure and composition of the local fauna. Development of management plans on conservation areas such as the implementation of “Area de Proteção Ambiental Rio Uberaba – APA-Rio Uberaba” project (SEMEA 2004) is necessary to mitigate the effects and help the sustainable use of local natural resources.

Figure 8. 

Loricariidae, genus Hypostomus collected in the Uberaba River (dorsal, lateral, and ventral photographs). 1 Hypostomus ancistroides 2 Hypostomus albopunctatus 3 Hypostomus strigaticeps 4 Hypostomus margaritifer 5 Hypostomus butantanis 6 Hypostomus regani 7 Hypostomus aff. paulinus 8 Hypostomus topavae and 9 Hypostomus nigromaculatus, Photographs are of specimens presented in Table 2. Scale bar: 10 mm.

Figure 9. 

Another loricariids collected in the Uberaba River (dorsal, lateral, and ventral photographs). 1 Hypostomus aff. hermanni 2 Hypostomus fluviatilis 3 Rineloricaria latirostris 4 Proloricaria prolixa 5 Curculionichthys insperatus 6 Microlepdogaster dimorpha and 7 Loricaria lentiginosa (uncataloged). Photographs are of specimens presented in Table 2. Scale bar: 10 mm.

Acknowledgements

We thank Fernando Carvalho, Fernanda Martins, and Claudio Zawadzki for help with species identification; Felipe Montefeltro, Nivaldo Urzulin, and Cristiéle Ribeiro for proofreading; Cristiane Santos, Simony Santos, Ronaldo Santos, Flavienne Carvalho, Gustavo Carvalho, and Luciano Henrique for the help in the field work; José Noel Prata, Paulo César Macedo, Vander Neto, Mario Cesar, Eliana Miranda Araújo, and Pedro César Chagas for support in various activities throughout the work. We are grateful for the support given by the Military Police of Minas Gerais (16° Pel. and 5ª Cia PM Ind. MAmb. de Planura). This project was partially funded by Prefeitura Municipal de Uberaba. DCR and MRT were supported by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (Capes) and JMAC and FL by the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq). The manuscript benefitted from reviews by Carla Pavanelli, Nina Bogutskaya, Nathalie Yonow, and anonymous referees.

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