The fish fauna of the upper Piraí drainage, a transposed mountain river system in southeastern, Brazil

Abstract

An annotated checklist of the ichthyofauna from the upper Piraí river drainage is provided. The Piraí river was a major right-bank tributary of the Paraíba do Sul river, but it has been artificially diverted to the coastal Guandu river system in southeastern Brazil to generate electric power and water for the metropolitan area of Rio de Janeiro. Based on our field sampling of 23 sites, 32 species belonging to 24 genera and 12 families were collected in 6 headwater tributaries of the Piraí river between 2009 and 2016. Phalloceros harpagos (Lucinda, 2008), Astyanax intermedius (Eigenmann, 1908), and Neoplecostomus microps (Steindachner, 1877b) were the most abundant and most widely distributed species in the samples. The sampled ichthyofauna is mostly composed by species from the Paraíba do Sul basin. Eight species are reported for the first time in the upper Piraí drainage, showing the importance of continuous ichthyofaunal surveys of fish in remaining areas of Atlantic Forest.

Full text

The sh fauna of the upper Piraí drainage, a transposed mountain
river system in southeastern, Brazil
Victor de Brito, Paulo Andreas Buckup
Departamento de Vertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, Rio de Janeiro, RJ, CEP 20.940-040,
Brazil.
Corresponding author: Victor de Brito, victordebrito.nf@gmail.com
Abstract
An annotated checklist of the ichthyofauna from the upper Piraí river drainage is provided. The Piraí river was a major
right-bank tributary of the Paraíba do Sul river, but it has been articially diverted to the coastal Guandu river system in
southeastern Brazil to generate electric power and water for the metropolitan area of Rio de Janeiro. Based on our eld
sampling of 23 sites, 32 species belonging to 24 genera and 12 families were collected in 6 headwater tributaries of the
Piraí river between 2009 and 2016. Phalloceros harpagos (Lucinda, 2008), Astyanax intermedius (Eigenmann, 1908),
and Neoplecostomus microps (Steindachner, 1877b) were the most abundant and most widely distributed species in the
samples. The sampled ichthyofauna is mostly composed by species from the Paraíba do Sul basin. Eight species are
reported for the rst time in the upper Piraí drainage, showing the importance of continuous ichthyofaunal surveys of
sh in remaining areas of Atlantic Forest.
Key words
Atlantic Forest, coastal streams, ichthyological survey, Neotropical shes, Paraíba do Sul.
Academic editor: Mariangeles Arce H. | Received 19 September 2018 | Accepted 8 January 2019 | Published 22 February 2019
Citation: de Brito V, Buckup PA (2019) The sh fauna of the upper Piraí drainage, a transposed mountain river system in southeastern, Brazil.
Check List 15 (1): 235–247. https://doi.org/10.15560/15.1.235
Introduction
The Brazilian Atlantic Forest is the second largest biome
in South America, with one of the highest rates of spe-
cies richness and endemism on the planet (Galindo-Leal
and Câmara 2003, Ribeiro et al. 2009). Due to environ-
mental impacts originating from human activities, only
16% of the original forest is preserved in reduced and
isolated patches (Ribeiro et al. 2009, 2011). Freshwater
streams in southeastern Brazil are directly aected by
the degradation of the Atlantic Forest, as their headwa-
ters are located in these few remaining patches of for-
est, and the reduction of the riparian vegetation is one
of the main sources of environmental impact in water
bodies (Cetra and Ferreira 2016). Systematic surveys of
sh communities that inhabit these streams are neces-
sary to reduce knowledge gaps about biodiversity and to
understand the state of conservation of the Atlantic For-
est (Buckup et al. 2014).
The upper Piraí river drainage is a former tributary
of the Paraíba do Sul river basin that has been articially
diverted to the Guandu river system through a complex
system of tunnels and pumps to generate electric power
and water for the metropolitan area of Rio de Janeiro.
The Piraí river drains the Biodiversity Corridor Tinguá-
Bocaina (BCTB) (Paiva and Coelho 2015). The BCTB
Check List 15 (1): 235–247
https://doi.org/10.15560/15.1.235
1
15
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ANNOTATED LIST OF SPECIES

236 Check List 15 (1)
connects Atlantic Forest fragments delimited by the Tin-
guá Biologic Reserve, in the central region of Rio de
Janeiro, and the National Park of Serra da Bocaina, on
the southern coast of the state. The BCTB is one of the
most important priority areas for the preservation of the
Atlantic Forest biodiversity due to its location at the most
critical point of fragmentation of the largest continuous
area of this biome in the Serra do Mar (Vilar et al. 2012).
In the present study, we surveyed the ichthyofauna from
Upper Piraí drainage in order to provide an annotated
checklist of freshwater sh species occurring in the area
of study. This study is part of a long-term program to
monitor the eects of forest restoration in the Guandu
river system partly published by Vilar et al. (2012) and
Castello Branco (2015). It includes a larger area and an
updated inventory of sh species presented in previous
studies (e.g. Buckup et al. 2014).
Methods
Study area. The Piraí river drainage (Fig. 1) is located
within the boundaries of the municipalities of Rio Claro
and Barra do Piraí, Rio de Janeiro state, southeastern Bra-
zil. The Piraí river was a major tributary of the Paraíba
do Sul river. Since the beginning of the 20th century,
it has been diverted to the coastal Guandu river basin,
in the Rio de Janeiro Metropolitan Region. The head-
waters of the Piraí drainage are composed by streams
that ow from the northern face of the mountain system
of Serra do Mar. This mountain chain has an extension
of 1,000 km between Rio de Janeiro state and north of
Santa Catarina state (Almeida and Carneiro 1998).
Sampling. Fish were collected in 23 sampling sites along
6 streams, rio Coutinhos, rio Papudos, rio Parado, rio
Passa Quatro, and Rio das Pedras, and the main channel
of the upper rio Piraí (Table 1, Fig. 1) in 4 expeditions
between the years of 2009 and 2016, under collecting
permit 12129, issued by the Instituto Chico Mendes de
Conservação da Biodiversidade. The sampling sites
were selected to cover a wide range of habitats, from the
alluvial plain of the streams to the high-mountain rap-
ids. Each site was surveyed during 1 h of exhaustive
sampling activity by a team of 5 ichthyologists. Fishing
equipment included beach seines (2, 3, 5 and 15 m long,
4 mm nylon mesh), dip nets with metal handles (40 ×
90 cm basket area, 2 mm plastic mesh), and casts nets
(12 mm mesh size). An acoustic amplier of electric sig-
nals was used to detect specimens of the order Gymno-
tiformes hidden in marginal vegetation and rocks. The
collected specimens were preserved in 10% formalin
solution and transferred to 70% ethanol after 48 h. Part
of the specimens were preserved in anhydrous ethanol
Figure 1. Map of the upper Piraí river drainage, indicating the collecting sites along rio Coutinhos (C), rio Papudos (P), rio Parado (A), rio
Passa Quatro (Q), Rio das Pedr as (R), an d the main channel of rio Piraí (I). Geogr aphic coordinates and altitude of th e sites are listed in Table 1.

de Brito and Buckup | Ichthyofauna from Upper Piraí 237
for DNA-sequencing studies. The examined specimens
were deposited in the ichthyological collection of the
Museu Nacional, Universidade Federal do Rio de Janeiro
(MNRJ) (Appendix, Table A1). Standard length, abbre-
viated as SL, was measured to the base of the middle
caudal-n rays.
Results
We collected 7,860 sh specimens of 32 species belong-
ing to 24 genera and 12 families in the upper Piraí river
drainage (Table 2). Among the river sections that com-
prise the sampled drainage, the main channel of the Piraí
river had the higher species richness, with 22 species.
The number of species captured in the other 5 streams
varied between 18 in the Coutinhos, Passa Quatro, and
Rio das Pedras, and 17 in the Papudos and Parado. Cat-
shes were the most diverse taxonomic group, including
16 species of Siluriformes (50%), followed by 9 Char-
aciformes (28%), 4 Cichliformes (13%), 2 Cyprinodon-
tiformes (6%), and 1 Gymnotiformes (3%). The most
diverse families were Loricariidae with 9 species (28%),
Characidae and Cichlidae, with 4 species (13%) each,
and Trichomycteridae and Heptapteridae, with 3 species
(9%) each.
Phalloceros harpagos was the most abundant spe-
cies, with 2,437 specimens (31.1%), followed by Astyanax
intermedius with 1,597 specimens (20.4%), Neople-
costomus microps with 676 specimens (8.6%), Pareio-
rhina rudolphi with 553 specimens (7.1%), Geophagus
brasiliensis with 497 specimens (6.4%), and Characid-
ium lauroi with 444 specimens (5.7%). These 6 species
combined represented 79.3% of all specimens collected
in all sampled sites. Some species were notably rare in
the samples, including Astyanax sp. a. scabripinnis,
Hemipsilichthys gobio, and Hypostomus luetkeni, which
were represented by 1 specimen each. The 3 most abun-
dant species were also the most widely distributed along
the Upper Piraí drainage, with P. harpagos absent only
in 1 site (C2), N. microps absent in 2 sites (A1, A2), and
A. intermedius absent in 3 sites (A1, A2, R1). Trichomyc-
terus nigroauratus, G. brasiliensis, and T. macrophthal-
mus were also widely distributed, having been present in
18, 16 and 14 sites, respectively.
The geographic distribution of the taxa collected in
the study area and the diagnosis of the sympatric con-
generic species are commented below. Although species
listed here occur in other drainages, diagnostic char-
acters described next are limited to identify the set of
species that occur in the upper Piraí drainage. These
diagnoses might be useful for ichthyological studies in
other drainages of the Paraíba do Sul river basin, but in
that case, the occurrence of other species not listed in the
upper Piraí have to be considered.
Order Characiformes
Family Anostomidae
Hypomasticus mormyrops (Steindachner, 1875)
Figure 2B
Geographic distribution: Coastal rivers between
Mucuri and Paraíba do Sul river basins (Vieira et al.
2015).
Family Crenuchidae
Characidium lauroi Travassos, 1949
Figure 2F
Geographic distribution: Upper reaches of Paraíba do
Sul tributaries.
Diagnosis: Characidium lauroi diers from C. vidali
by the presence of small rounded dark marks below the
lateral stripe, which are separated from the dorsal-lateral
bars, and pigmentation bars on the caudal n are usually
absent or inconspicuous (Melo 2001b, Buckup et al. 2014).
Characidium vidali Travassos, 1967
Figure 2H
Geographic distribution: Coastal rivers between
Guanabara bay and Paraíba do Sul river basins.
Diagnosis: Characidium vidali can be distinguished
from C. lauroi by the presence of vertical polyhedric
dark marks along the medium and inferior portion of
the body and by having pigmentation bars on caudal n
(Buckup et al. 2014).
Family Bryconidae
Brycon opalinus (Cuvier, 1819)
Figure 2G
Geographic distribution: Paraíba do Sul and Doce
river basins.
Tab le 1. Geographic coordinates from the sampling sites in the
Upper Piraí river drainage, Rio de Janeiro state, Brazil.
Stream Site Lat itude (S) Longitude (W) Alti tude (m)
Coutinhos C1 22°50’37” 4 4° 13’45” 610
C2 22°50’37” 44°13’36” 582
C3 22°50’10” 44°12’33” 537
C4 22°49’28” 44°12’39” 572
Papudos P1 22°55’27” 4 4° 13’27” 1002
P2 22°54’20” 44°13’38” 951
P3 22°52’42” 44°13’38” 571
P4 22°52 ’24” 44°12’39” 563
Parado A1 22°51’51’’ 4 4°08’ 21” 1050
A2 22° 51’41’’ 44°09’07’’ 976
A3 22°50’59’ ’ 44 °10’4 6’’ 561
A4 22°50’03’’ 4 4°11’17 ’ ’ 545
Passa Quatro Q1 22°49’ 31’’ 4 4°08’16’’ 712
Q2 22°49’22’’ 44°09’32’ ’ 622
Q3 22°49’05’’ 4 4 ° 11’ 13 ’’ 513
Piraí I1 22 °51’18 ” 4 4°11’5 5” 555
I2 22°50’56” 4 4 °11’4 5” 548
I3 22°50’24” 4 4° 12’ 09” 535
I4 22°49’39” 4 4°11’43 ” 519
Rio das Pedras R1 22°54’11” 44° 11’10 ” 842
R2 22°53’33” 44°11’30” 786
R3 22°52’30” 4 4 °11 ’4 8 ” 576
R4 22° 51’59” 44 °12 ’0 7” 554

238 Check List 15 (1)
Tab le 2. List of the freshwater shes of upper Piraí river drainage.
Tax on
Sites
C1 C2 C3 C4 P1 P2 P3 P4 A1 A2 A3 A4 Q1 Q2 Q3 I1 I2 I3 I4 R1 R2 R3 R4
CHARACIFORMES
Anostomidae
Hypomasticus mormyro ps X X X
Crenuchidae
Characidium lauroi X X X X X X X X X
Characidium vidali X X X X X X X X X X
Bryconidae
Brycon opali nus X X
Characidae
Astyanax giton XXXX X
Astyanax intermedius XXXXXXXX XXXXXXXXX XXX
Astyanax sp. a. scabripinnis X
Oligosarcus hepsetus X X X X X X X X X X X X
Erythrinidae
Hoplias malabaricus X X X X
SILURIFORMES
Trichomycteridae
Trichomycterus
macrophthalmus
X X XX XXXXXXXX XX
Trichomycterus mariamole X X X X X
Trichomycterus nigroauratus XXXXXXXXXXXX XX XXXX
Callichthyidae
Scleromystax barbatus X
Loricariidae
Harttia carvalhoi X X X X X X X X X X X X
Harttia loricariformis X X X X X X X
Hemipsilichthys gobio X
Hemipsilichthys papillatus X X
Hypostomus anis X X X X X X X X
Hypostomus luetkeni X
Neoplecostomus microps XXXXXXXX XXXXXXXXXXXXX
Pareiorhina rudolphi X X X X X
Rineloricaria sp. cf. R . lima X X X X X X X X X X X X
Heptapteridae
Imparnis minutus X X X X X X X X X X X
Pimelodella lateristriga X X X X X X X X X
Rhamdia quelen X X X X X X X X X
GYMNOTIFORMES
Gymnotidae
Gymnotus pantherinus XXX XX XX XXXXX
CYPRINODONTIFORMES
Poeciliidae
Phalloceros harpagos X XXXXXXXXXXXXXXXXXXXXX
Poecilia reticulata X X
CICHLIFORMES
Cichlidae
Australoheros sp. X X
Crenicichla lepidota X X
Geophagus brasiliensis XX XX XXXXXXXXXX XX
Oreochromis niloticus X X
Family Characidae
Astyanax giton Eigenmann, 1908
Figure 2A
Geographic distribution: Coastal rivers between
Paraíba do Sul and Itabapoana river basins.
Diagnosis: Astyanax giton is distinguished from its
congeners in the Piraí drainage by presenting the posterior
portion of pelvic n surpassing the urogenital opening. A
gradual variation of size in dentary teeth size and higher
body height (approximately 41% SL) are also distinguish-
able features of this species in relation to Astyanax inter-
medius and Astyanax sp. a. scabripinnis (Melo 2001a).
Astyanax intermedius Eigenmann, 1908
Figure 2C
Geographic distribution: Paraíba do Sul and Doce
river basins and coastal rivers of Rio de Janeiro state

de Brito and Buckup | Ichthyofauna from Upper Piraí 239
(Lezama et al. 2011).
Diagnosis: Astyanax intermedius diers from A.
giton by body depth less than 41% of SL and abrupt
reduction of tooth size posterior to the fth dentary tooth
(Melo 2001a). Also, in contrast with A. giton, the poste-
rior portion of pelvic n in A. intermedius does not sur-
pass the urogenital opening in this species. Astyanax
intermedius can be distinguished from Astyanax sp. a.
scabripinnis, by the presence of 5 cusps in most dentary
teeth and regular sized adipose n.
Astyanax sp. a. scabripinnis (Jenyns, 1842)
Figure 2E
Geographic distribution: Unknown.
Diagnosis: This species was represented by a single
individual and diers from A. intermedius by having 7
cusps in most dentary teeth (vs. usually 5 cusps in A.
intermedius) and by having a smaller adipose n. Asty-
anax sp. a. scabripinnis diers from A. giton by the
following characters: body depth smaller than 41% of SL
and abrupt size reduction posterior to the fth dentary
teeth (Melo 2001a)
Oligosarcus hepsetus (Cuvier, 1829)
Figure 2I
Geographic distribution: Coastal rivers from Santa
Catarina to Rio de Janeiro state.
Family Erythrinidae
Hoplias malabaricus ( Blo ch, 179 4)
Figure 2D
Geographic distribution: Widespread across South
and Central America (Vieira et al. 2015).
Order Siluriformes
Family Trichomycteridae
Trichomycterus macrophthalmus Barbosa & Costa, 2012
Figure 3A
Geographic distribution: Piraí river drainage.
Figure 2. Fish species collected in upper Piraí river drainage. A. Astyanax giton, MNRJ 37986, 63.6 mm SL. B. Hypomasticus mormyrops,
MNRJ 46 698, 128.6 mm SL . C. Astyanax intermedius, MNR J 43805, 77.2 mm SL. D. Hopl ias malabaricus, MNRJ 43830, 70.8 mm SL . E. Ast yanax
sp. a. scabripinnis, MNRJ 36427, 71.8 mm SL. F. Characidium lauroi, MNRJ 43823, 57.3 mm SL. G. Brycon opalinus, MNRJ 47259, 116.6 mm SL.
H. Characidium vidali, MNRJ 43807, 66.6 mm SL. I. Oligosarcus hepsetus, MNRJ 43812, 63.1 mm SL.

240 Check List 15 (1)
Diagnosis: Trichomycterus macrophthalmus is char-
acterized by the colour pattern consisting of transverse
dark bars crossing the dorsum, which can be fused with
lateral marks in this species (Barbosa and Costa 2012).
Trichomycterus macrophthalmus is distinguished from
T. mariamole and T. nigroauratus by the presence of 9
pairs of ribs and eye diameter 13.2% to 14.6% the size of
head length (HL) (Barbosa and Costa 2012).
Trichomycterus mariamole Barbosa & Costa, 2010
Figure 3C
Geographic distribution: Piraí river drainage and
upper reaches of Alambari and Barreiro rivers, Paraíba
do Sul river basin.
Diagnosis: Trichomycterus mariamole has light yel-
low background coloration with small circular brown
marks on anks, diameter of the eyes 9% to 11% of HL,
opercular region of odontoids reduced and laterally posi-
tioned, and 7 pectoral n rays (Barbosa and Costa 2010).
Trichomycterus mariamole diers from T. claudiae,
a species that occurs in the middle portion of the Piraí
drainage, by the short ill-dened stripe restricted to the
anterior portion of the ank (vs. well dened stripe along
the whole an k) and fewer interopercular odontodes (27–
34 vs. 41–46).
Trichomycterus nigroauratus Barbosa & Costa, 2008
Figure 3E
Geographic distribution: Upper Piraí and Barreiro
river drainages in the Paraíba do Sul river basin.
Diagnosis: Trichomycterus nigroauratus diers mor-
phologically from T. mariamole and T. macrophthalmus
by the presence of golden spots on the snout and body,
and broad (wider than long) metapterygoid bone (Bar-
bosa and Costa 2008). Trichomycterus nigroauratus
presents ontogenetic variation of coloration, with juve-
niles exhibiting a black stripe along the lateral midline,
and adults presenting stripes with irregular borders and
dark stains that can cover the entire body in a homoge-
neous pattern (Buckup et al. 2014).
Family Callichthyidae
Scleromystax barbatus (Quoy & Gaimard, 1824)
Figure 3G, H
Geographic distribution: Coastal rivers from Santa
Catarina to Rio de Janeiro state.
Family Loricariidae
Harttia carvalhoi Miranda Ribeiro, 1939
Figure 4A
Geographic distribution: Paraíba do Sul river basin.
Diagnosis: The 2 species of Harttia that occur in the
area of study can be distinguished by the distribution of
plates in front of the anus: Harttia carvalhoi is distin-
guished from H. loricariiformis by the absence of pre-
anal plates (Langeani et al. 2001).
Figure 3. Fish species collected in upper Piraí river drainage. A. Trichomycterus macrophthalmus, MNRJ 43760, 52.4 mm SL. B. Imparnis
minutus, MNRJ 43770, 83.4 mm SL. C. Trichomycterus mariamole, MNRJ 36525, 58.72 mm SL. D. Pimelodella lateristriga, MNRJ 43888, 39.7
mm SL. E. Trichomycterus nigroauratus, MNRJ 38003, 48.8 mm SL. F. Rhamdia quelen, MNRJ 43889, 98.6 mm SL. G. Scleromystax barbatus,
MNRJ 46722, 66.1 mm SL, male. H. Scleromystax barbatus, MNRJ 46722, 57.62 mm SL, female.

de Brito and Buckup | Ichthyofauna from Upper Piraí 241
Harttia loricariformis Steindachner, 1877a
Figure 4B
Geographic distribution: Paraíba do Sul river basin
and coastal rivers from Rio de Janeiro to Espírito Santo.
Diagnosis: Dierently from its congener, Harttia lori-
cariformis presents 2 large and trapezoid preanal plates
touching the small anterior plates (Langeani et al. 2001).
Hemipsilichthys gobio (Lütken, 1874a)
Figure 4C
Geographic distribution: Paraíba do Sul river basin.
Diagnosis: Hemipsilichthys gobio diers from H. papil-
latus by its rectangular or oval-shaped dorsal-n spinelet,
and by the larger orbital diameter (12.0–14.7% HL, Reis et
al. 2006).
Hemipsilichthys papillatus Pereira et al. 2000
Figure 4D
Geographic distribution: Paraíba do Sul river basin.
Diagnosis: Hemipsilichthys papillatus diers from
H. gobio by not having plates in the dorsal midline
between the base of the dorsal n and the adipose n,
and a smaller orbital diameter (8.6–11.8% HL, Reis et
al. 2006).
Hypostomus anis (Steindachner, 1877a)
Figure 4E
Geographic distribution: Paraíba do Sul and Doce
river basins and coastal rivers of southeastern Brazil.
Diagnosis: Individuals of Hypostomus anis can be
distinguished H. luetkeni by the presence of 1 main post-
supraoccipital plate and 4 lateral ridges on anks (Maz-
zoni et al. 1994). The prevailing pigmentation pattern
in adults of H. anis comprises numerous black spots,
whereas H. luetkeni has a more uniform dark pigmenta-
tion pattern.
Hypostomus luetkeni (Steindachner, 1877b)
Figure 4F
Geographic distribution: Paraíba do Sul river basin.
Diagnosis: Hypostomus luetkeni ca n be distinguished
from H. anis by the presence of 2 or 3 plates in the
main post-supraoccipital series and the absence of ridges
on the anks of the body (Mazzoni et al. 1994). Dark
spots in the body, if present, are less conspicuous than
in H. anis.
Neoplecostomus microps (Steindachner, 1877b)
Figure 4G
Figure 4. Fish species collected in upper Piraí river drainage. A. Harttia carvalhoi, MNRJ 43816, 78.1 mm SL. B. Hart tia loricariformis, MNRJ
43773, 42.1 mm SL. C. Hemipsilichthys gobio, MNRJ 43868, 75.3 mm SL (Adult individual collected in the rio do Braço, 22° 47’ 09”S 44° 10’
44”W, a tributary of the Piraí river. The exemplar of H. gobio, MNRJ 36482, collected in this study is a small juvenile.). D. Hemipsilichthys
papillatus, MNRJ 46657, 53.7 mm SL. E. Hypostomus anis, MNRJ 43824, 62.6 mm SL. F. Hypostomus luetkeni, MNR J 47007, 112.8 mm SL.
G. Neoplecostomus microps, MNRJ 43808, 87 mm SL. H. Pareiorhina rudolphi, MNRJ 46835, 65.9 mm SL. I. Rineloricaria sp. cf. R. lima, MNRJ
43826, 76.3 mm SL.

242 Check List 15 (1)
Geographic distribution: Paraíba do Sul river basin
and coastal rivers from Rio de Janeiro state.
Pareiorhina rudolphi (Miranda Ribeiro, 1911)
Figure 4H
Geographic distribution: Paraíba do Sul river basin.
Rineloricaria sp. cf. R. lima (Kner, 1853)
Figure 4I
Geographic distribution: Unknown.
Diagnosis: In this study we refer this species as
Rineloriaria sp. cf. R. lima due to similarity of the spec-
imens from the Piraí drainage with specimens of this
genus collected in the Paraíba do Sul previously identi-
ed as Rineloricaria cf. lima by other authors (e.g. Fich-
berg 2008). The taxonomic status of R. lima, however, is
uncertain because the type material is lost, and the type
locality is unknown.
Family Heptapteridae
Imparnis minutus (Lütken, 1874b)
Figure 3B
Geographic distribution: Upper São Francisco, upper
Paraná , Paraíba do Sul and Ribeira de Igu ape river basi ns.
Pimelodella lateristriga (Lichtenstein, 1823)
Figure 3D
Geographic distribution: Coastal rivers between
Jequitinhonha and Paraíba do Sul river basins.
Rhamdia quelen (Quoy & Gaimard, 1824)
Figure 3F
Geographic distribution: Widespread from Mexico
to Argentina.
Order Gymnotiformes
Family Gymnotidae
Gymnotus pantherinus (Steindachner, 1908) (Fig. 5A)
Geographic distribution: Coastal rivers of Southeast-
ern Brazil.
Order Cyprinodontiformes
Family Poeciliidae
Phalloceros harpagos Lucinda, 2008
Figure 5C, E
Geographic distribution: Paraná and Paraguai river
basins and coastal drainages from Santa Catarina to
Espírito Santo states.
Poecilia reticulata Peters, 1859
Figure 5G, I
Geographic distribution: Introduced species, natural
from northern South America.
Figure 5. Fish species collected in the upper Piraí river drainage. A. Gymnotus pantherinus, MNRJ 43827, 172.5 mm SL. B. Australoheros
sp., MNRJ 43780, 50.3 mm SL. C. Phalloceros harpagos, MNRJ 43828, 34.2 mm SL, female. D. Crenicichla lepidota, MNRJ 43779, 85.9 mm SL.
E.Phalloceros harpagos, MNRJ 43828, 25.3 mm SL, male. F. Geophagus brasiliensis, MNRJ 43778, 64.2 mm SL. G. Poecilia reticulata, MNRJ
43763, 36.8 mm SL, female. H. Oreochromis niloticus, MNRJ 37964, 96.56 mm SL. I. Poecilia reticulata, MNRJ 43763, 19.2 mm SL, male.

de Brito and Buckup | Ichthyofauna from Upper Piraí 243
Order Cichliformes
Family Cichlidae
Australoheros sp. Říčan & Kullander, 2006
Figure 5B
Geographic distribution: Unkown.
Crenicichla lepidota Heckel, 1840
Figure 5D
Geographic distribution: Widely distributed in South
America.
Geophagus brasiliensis (Quoy & Gaimard, 1824)
Figure 5F
Geographic distribution: Coastal rivers of northeast-
ern Brazil to Uruguay.
Oreochromis niloticus (Linnaeus, 1758)
Figure 5H
Geographic distribution: Introduced species, origi-
nally from Northern and eastern Africa.
Discussion
In this study, the prevalence of Siluriformes and Chara-
ciformes in the composition of the sh assemblage fol-
lows the observed composition pattern of Atlantic Forest
streams (e.g. Sarmento-Soares et al. 2007, Ferreira and
Petrere 2009, Camilo et al. 2015) and most Neotropical
freshwater systems (Lowe-McConnell 1999, Abilhoa et
al. 2011). The ichthyofauna of the upper Piraí is com-
posed mostly by species that are widely distributed in the
Paraíba do Sul basin. Scleromystax barbatus is a wide-
spread species in coastal rivers; however, this species
was collected only once, near the city of Lídice, despite
our extensive sampling eort in the Piraí basin. The neo-
type of S. barbatus was collected in a nearby coastal
river, and the species is quite abundant in the adjacent
coastal plain. The city of Lidice is located less than 10
km from a recreational area in a coastal lowland river.
This ichthyofaunal composition indicates that endemic
species from Guandu do not live in the study area, proba-
bly because the complex system of tunnels, pumps, dams
and hydropower turbines, as well as the steep altitudi-
nal gradient across the transposition system serve as an
eective barrier for upstream sh migration.
As expected, the species diversity was higher in the
lowest stretches of the upper Piraí drainage than in its
headwaters. According to the River Continuum Concept
(Vannote et al. 1980), the composition and structure of
ichthyofauna in freshwater streams is directly related
to altitude. Thus, high altitude stretches with compara-
tively lower temperature, high dissolved oxygen and fast
owing waters tend to have lower species richness than
downstream sites with decreasing slope, increasing tem-
perature and high availability of pools.
Among the widespread species in the study area, the
high number of individuals of Phalloceros harpagos
present in our samples is probably correlated with the
ability of poecilids to colonize a wide range of habitats
and tolerate dierent levels of environmental quality
(Araújo et al. 2009). The second most abundant spe-
cies in upper Piraí drainage, Astyanax intermedius, is an
opportunistic strategist, characterized by the capacity
of producing rapid population turnover and colonisation
(Souza et al. 2015). Regarding taxonomic representation,
Astyanax and Trichomycterus were the most diverse
genera in the study area, represented by 3 species each.
Oddly, individuals of Trichomycterus claudiae (Barbosa
and Costa 2010) were not registered in the upper Piraí
drainage. This species was expected to occur in the sam-
pled drainage due to the proximity of its type locality.
Two species collected in the upper Piraí drainage
are included in the list of Brazilian fauna threatened by
extinction (ICMBio 2016): Brycon opalinus, present in
the lower reaches of Rio das Pedras stream, and Hemip-
silichthys gobio, collected in the main channel of rio
Piraí. Brycon opalinus, known as “pirapitinga”, is Vul-
nerable due to commercial exploration and environment
degradation, and its restricted distribution to preserved
tributaries of the Paraíba do Sul and Doce headwaters
(Gomiero and Braga 2007, ICMBio 2016). Hemipsilich-
thys gobio is Endangered due to habitat reduction and
fragmentation (ICMBio 2016). The distribution of this
species is limited to highly oxygenated cold waters of
high-elevation, fast owing tributaries of the Paraíba do
Sul river basin. The presence of these 2 species indicates
that preserved stretches in the upper Piraí drainage serve
as natural refuge areas for threatened sh fauna.
Two non-native species were recorded in 4 sampling
sites along the study area: Poecilia reticulata was cap-
tured in the lowest stretches of the Passa Quatro stream,
and Oreochromis niloticus occurred in 2 sites of the
main channel of upper Piraí. Poecilia reticulata is an
exotic poecilid species from Venezuela, widely intro-
duced in aquatic environments of the Americas due to
its predation on mosquito larvae (Santos 1958). Oreo-
chromis niloticus, known as Nile Tilapia, is an invasive
cichlid species, originally from Africa, that is widely
cultivated in tanks and net cages in Brazil, and was prob-
ably introduced by escapes from pisciculture (Orsi and
Agostinho 1999). Regarding environmental conditions,
invasive species are usually more successful in nega-
tively aected systems or where native species popula-
tion are depleted (Leidy et al. 2011). The introduction
and population establishment of these invasive species
in the mid-lower reach of the Passa Quatro stream and
the main channel of upper Piraí is largely associated with
areas of intense human occupation and land-use.
Among the 32 species reported in this study, the fol-
lowing 8 were not listed in the rst survey of the Piraí
headwaters (Buckup et al. 2014): Hypomasticus mor-
myrops, Hoplias malabaricus, Scleromystax barbatus,
Hemipsilichthys gobio, Hypostomus luetkeni, Poecilia
reticulata, Australoheros sp., and Crenicichla lepidota.
The absence of Hypostomus luetkeni, Poecilia reticu-
lata, Australoheros sp., and C. lepidota in the sites sur-
veyed by Buckup et al. (2014) has been conrmed in our

244 Check List 15 (1)
current survey, as these species have only been captured
outside of the area of that earlier study. The single speci-
men of Hemipsilichthys gobio (MNRJ 36482) was mis-
identied by Buckup et al. (2014) as H. papillatus. In
addition to the slight increase of diversity, some species
were captured in signicant numbers only in the recent
survey (e.g. Brycon opalinus). The increase in species
richness and abundance of the ichthyofauna in the upper
Piraí river drainage recorded over our 7-year survey pos-
sibly correlates with the consolidation of recent conser-
vation programs implemented in the study area to recover
riparian forest (Castello-Branco 2015), although other
environmental factors cannot be ruled out. Thus, the
record of additional species not previously captured in
the area, despite the use of standardized sampling eort,
demonstrates the importance of continuous surveys in
freshwater systems, in order to increase the knowledge
on the distribution and impact of human activity on sh
assemblages present in Atlantic Forest remnants.
Acknowledgements
We thank The Nature Conser vancy, represented by Paulo
Petry and Anita Diderichsen, and the Associação Pró-
Gestão das Águas da Bacia Hidrográca do Rio Paraíba
do Sul (Termo de Concessão No. 03.007.001.17) for the
nancial support of this project. We also thank our col-
leagues from Museu Nacional, Decio Ferreira de Moraes
Junior, Emanuel Neuhaus, Gabriel Beltrão, Igor Caval-
canti Santos, Leandro Vila-Verde, Marcelo Ribeiro de
Britto, Mar ia Clara N. Chaves, and Fernando Kilesse Sal-
gado for help with the eld work. VB and PAB research
are supported by the Conselho Nacional de Desenvolvi-
mento Cientíco e Tecnológico (processes 134423/2016-
0, 476822/2012-2, 312801/2017-3) and the Coordenação
de Aperfeiçoamento do Pessoal de Nível Superior.
Authors’ Contributions
PAB conceived and coordinated the project. VB and PAB
performed eldwork and analyzed the data. VB identi-
ed the specimens, took the photographs and wrote the
manuscript. PAB revised and corrected the manuscript.
This study is part of a M.Sc. dissertation study con-
ducted by VB.
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Appendix
Table A1. Catalog numbers of voucher specimens.
Tax on Catalog number (number of exemplars)
CHARACIFORMES
Anostomidae
Hypomasticus mormyro ps MNRJ 46 698 (3), MNR J 47001 (2), MNRJ 47012 (1), MNR J 47256 (1)
Crenuchidae
Characidium lauroi MNRJ 36451 (3), MNRJ 36 458 (3), MNR J 36472 (32), MNRJ 36486 (1), MNRJ 36508 (50), MNRJ 36518 (6),
MNRJ 37977 (1), MNRJ 38019 (26), MNRJ 38032 (13), MN RJ 38039 (10), MNR J 38045 (25), MNRJ 43748 (39),
MNRJ 43759 (5), MNRJ 43806 (18), MNRJ 46650 (8), MNR J 46668 (73), MNRJ 46690 (25), MNRJ 46702 (20),
MNRJ 47260 (29)
Characidium vidali MNRJ 36457 (1), MNRJ 36471 (1), MNRJ 36497 (1), MNRJ 36507 (10), MNRJ 38018 (69), MNRJ 38031 (13),
MNRJ 43758 (37), MNRJ 43767 (1), MNRJ 43807 (7), MNRJ 43821 (3), MNRJ 46636 (2), MNRJ 46822 (3),
MNRJ 47023 (1), MNRJ 47261 (14)
Bryconidae
Brycon opali nus MNRJ 38022 (3), MNRJ 46701 (1), MNRJ 47259 (25)
Characidae
Astyanax giton MNR J 37834 (1), MNRJ 37835 (20), MNRJ 37974 (5), MNRJ 37986 (2), MNRJ 38021 (5), MNRJ 50819 (1),
MNRJ 50820 (2)
Astyanax intermedius MNR J 36459 (15), MNRJ 36473 (21), MNRJ 36487 (21), MNRJ 36498 (29), MNRJ 36510 (81), MNRJ 36519 (51),
MNRJ 36524 (22), MNRJ 37954 (5), MNRJ 37965 (28), MNR J 37978 (34), MNRJ 38 020 (51), MNRJ 38033 (42),
MNRJ 380 40 (19), MNRJ 38053 (10), MNR J 38087 (31), MNRJ 43749 (79), MNRJ 43755 (58), MNRJ 43766 (74),
MNRJ 43805 (16), MNRJ 43820 (15), MNRJ 46631 (20), MNRJ 466 43 (95), MNR J 46689 (23), MNRJ 46699 (10),
MNRJ 46713 (15), MNRJ 46791 (11), MNRJ 46813 (92), MNRJ 47002 (14), MNR J 47014 (6), MNRJ 47257 (91)
Astyanax sp. a. scabripinnis MNRJ 36427 (1)
Oligosarcus hepsetus MNRJ 364 60 (2), MNRJ 36474 (2), MNRJ 36488 (1), MNRJ 36499 (5), MNR J 36509 (3), MNRJ 37966 (2),
MNRJ 37979 (1), MNRJ 38023 (3), MNRJ 38034 (16), MNRJ 38052 (2), MNRJ 43768 (6), MNRJ 43812 (6),
MNRJ 46644 (1), MNRJ 46700 (5), MNRJ 46714 (13), MNR J 46792 (21), MNRJ 46814 (1), MNRJ 47003 (15),
MNRJ 47015 (9), MNRJ 47258 (2)
Erythrinidae
Hoplias malabaricus MNR J 38051 (1), MNRJ 43830 (1), MNRJ 46802 (1), MNRJ 47013 (1)
SILURIFORMES
Trichomycteridae
Trichomycterus macrophthalmus MNRJ 36467 (1), MNRJ 36478 (4), MNRJ 36494 (16), MNRJ 36504 (12), MNRJ 36513 (4), MNRJ 37957 (5),
MNRJ 43750 (22), MNRJ 43760 (27), MNRJ 43772 (13), MNRJ 43813 (2), MNRJ 43832 (1), MNRJ 4 6640 (8),
MNRJ 46795 (1), MNRJ 46825 (1)
Trichomycterus mariamole MNRJ 36452 (2), MNRJ 36520 (1), MNRJ 36525 (2), MNRJ 38047 (1), MNRJ 38088 (3), MNRJ 47024 (1)
Trichomycterus nigroauratus MNRJ 36453 (11), MNR J 36468 (3), MNRJ 36477 (3), MNRJ 36512 (2), MNRJ 37968 (5), MNRJ 38025 (3),
MNRJ 38035 (2), MNRJ 38041 (7), MNRJ 3804 6 (2), MNR J 38091 (1), MNRJ 43719 (5), MNRJ 43754 (1), MNR J
43814 (2), MNRJ 43833 (2), MNRJ 46645 (3), MNRJ 46669 (11), MNR J 46694 (1), MNR J 46712 (2), MNRJ 46796 (2),
MNRJ 46824 (1), MNRJ 46834 (6), MNRJ 47025 (1), MNRJ 47026 (1), MNRJ 47253 (13)
Callichthyidae
Scleromystax barbatus MNR J 46722 (4)
Loricariidae
Harttia carvalhoi MNRJ 36461 (3), MNRJ 36480 (9), MNRJ 36501 (3), MNRJ 36515 (2), MNRJ 37969 (5), MNRJ 38026 (15),
MNRJ 38036 (3), MNR J 38057 (1), MNRJ 43816 (2), MNRJ 43823 (4), MNRJ 466 41 (2), MNRJ 4 6826 (1),
MNRJ 47022 (2)
Harttia loricariformis MNRJ 36514 (1), MNR J 38037 (1), MNRJ 43773 (4), MNRJ 466 46 (2), MNRJ 46706 (4), MNRJ 46827 (1),
MNRJ 47262 (1)
Hemipsilichthys gobio MNRJ 36482 (1)
Hemipsilichthys papillatus MNRJ 36 448 (1), MNRJ 46657 (1)
Hypostomus anis MNR J 36502 (1), MNRJ 37958 (1), MNRJ 38062 (1), MNRJ 43781 (1), MNRJ 43824 (3), MNRJ 46632 (1),
MNRJ 46710 (1), MNRJ 46723 (4), MNRJ 46803 (1), MNRJ 46820 (1), MNRJ 46821 (1)
Hypostomus luetkeni MNRJ 47007 (1)

de Brito and Buckup | Ichthyofauna from Upper Piraí 247
Tax on Catalog number (number of exemplars)
Neoplecostomus microps MNRJ 36431 (23), MNRJ 36 442 (13), MNRJ 364 49 (8), MNR J 36454 (6), MNRJ 36462 (17), MNRJ 36481 (27),
MNRJ 364 89 (14), MNRJ 36516 (27), MNRJ 36521 (8), MNRJ 36526 (1), MNRJ 37960 (4), MNRJ 37971 (9),
MNRJ 37983 (8), MNRJ 37993 (34), MNR J 38008 (20), MNRJ 38013 (13), MNRJ 38017 (21), MNRJ 38028 (47),
MNRJ 38038 (15), MNRJ 38043 (17), MNRJ 38049 (9), MNRJ 38089 (6), MNRJ 43751 (21), MNR J 43761 (43),
MNRJ 43774 (10), MNRJ 43808 (38), MNRJ 43825 (118), MNRJ 46633 (11), MNRJ 46 647 (8), MNR J 46653 (20),
MNRJ 46662 (18), MNRJ 46691 (3), MNRJ 46708 (18), MNRJ 46785 (6), MNRJ 46797 (2), MNRJ 46816 (14),
MNRJ 47006 (4), MNRJ 47017 (5), MNRJ 47254 (10), MNRJ 47263 (12)
Pareiorhina rudolphi MNRJ 36455 (72), MNRJ 36522 (24), MNRJ 38012 (1), MNR J 38016 (7), MNR J 38042 (64), MNRJ 38048 (112),
MNRJ 46652 (34), MNRJ 46670 (141), MNRJ 46692 (58), MNRJ 46835 (40)
Rineloricaria sp. cf. R . lima MNRJ 36432 (5), MNRJ 36443 (1), MNRJ 36463 (7), MNR J 36479 (1), MNRJ 36490 (7), MNRJ 36503 (10),
MNRJ 37959 (12), MNRJ 37970 (2), MNRJ 37982 (4), MNRJ 37994 (8), MNRJ 38007 (2), MNR J 38027 (3),
MNRJ 38058 (10), MNRJ 43771 (84), MNRJ 43817 (2), MNRJ 43826 (23), MNRJ 46634 (10), MNRJ 46661 (2),
MNRJ 46707 (6), MNRJ 46717 (58), MNRJ 4 6784 (2), MNR J 46798 (16), MNRJ 46817 (4), MNR J 47005 (36)
Heptapteridae
Imparnis minutus MNRJ 364 64 (2), MNRJ 36476 (5), MNRJ 36491 (1), MNR J 36511 (1), MNR J 37956 (1), MNRJ 37967 (2),
MNRJ 37980 (2), MNRJ 37990 (1), MNR J 38024 (3), MNR J 38056 (2), MNRJ 43770 (5), MNRJ 43822 (3),
MNRJ 46637 (2), MNRJ 46703 (5), MNRJ 46721 (1), MNR J 46788 (1), MNRJ 46793 (6), MNRJ 47004 (3)
Pimelodella lateristriga MNRJ 36 434 (1), MNRJ 364 65 (1), MNRJ 36492 (1), MNRJ 38055 (4), MNRJ 43815 (1), MNR J 43831 (1),
MNRJ 46639 (1), MNR J 46704 (2), MNRJ 46715 (4), MNRJ 46823 (1)
Rhamdia quelen MNRJ 36433 (1), MNRJ 3646 6 (7), MNRJ 36475 (3), MNRJ 36493 (1), MNRJ 36500 (1), MNRJ 37955 (1),
MNRJ 37981 (4), MNRJ 37991 (1), MNRJ 43769 (2), MNRJ 46638 (2), MNRJ 46660 (4), MNRJ 46705 (2),
MNRJ 46716 (2), MNRJ 46789 (1), MNRJ 46794 (5), MNRJ 46815 (1), MNRJ 47016 (3)
GYMNOTIFORMES
Gymnotidae
Gymnotus pantherinus MNR J 37961 (1), MNRJ 38054 (9), MNRJ 43777 (2), MNRJ 43809 (11), MNRJ 43827 (4), MNRJ 46635 (6),
MNRJ 46648 (2), MNRJ 46 663 (4), MNR J 46718 (2), MNRJ 4 6786 (9), MNR J 46799 (8), MNRJ 46828 (2),
MNRJ 47008 (1), MNRJ 47019 (2)
CYPRINODONTIFORMES
Poeciliidae
Phalloceros harpagos MNRJ 36 437 (29), MNR J 36446 (1), MNRJ 36450 (2), MNRJ 36456 (84), MNRJ 36469 (3), MNRJ 36483 (11),
MNRJ 36495 (4), MNR J 36505 (15), MNRJ 36517 (5), MNRJ 36523 (70), MNRJ 36527 (84), MNRJ 37962 (55),
MNRJ 37972 (5), MNRJ 37984 (4), MNRJ 37997 (36), MNRJ 38010 (1), MNRJ 38029 (1), MNRJ 38044 (136),
MNRJ 38050 (43), MNRJ 38059 (16), MNR J 38090 (67), MNRJ 43722 (199), MNRJ 43752 (417), MNRJ 43762 (294),
MNRJ 43775 (40), MNRJ 43811 (34), MNRJ 43828 (241), MNRJ 4 6642 (4), MN RJ 46655 (157), MNRJ 46664 (6),
MNRJ 46671 (110), MNRJ 46693 (33), MNRJ 46711 (3), MNRJ 46719 (2), MNRJ 46800 (7), MNR J 46818 (7),
MNRJ 46836 (158), MNR J 47009 (3), MNRJ 47020 (29), MNRJ 47231 (8), MNRJ 47255 (13)
Poecilia reticulata MNRJ 43763 (266), MNRJ 43776 (1)
CICHLIFORMES
Cichlidae
Australoheros sp. MNRJ 3806 0 (2), MNR J 43780 (1)
Crenicichla lepidota MNRJ 43779 (5), MNRJ 47010 (6)
Geophagus brasiliensis MNR J 36470 (38), MNR J 36484 (7), MNRJ 36496 (3), MNRJ 36506 (3), MNRJ 37963 (14), MNRJ 37973 (16),
MNRJ 37985 (4), MNRJ 37998 (1), MNRJ 38009 (5), MNR J 38030 (5), MNRJ 38061 (10), MNRJ 43753 (11),
MNRJ 43764 (53), MNRJ 43778 (20), MNRJ 43810 (17), MNR J 43829 (57), MNR J 46649 (2), MNRJ 46667 (2),
MNRJ 46709 (40), MNR J 46720 (31), MNRJ 46787 (37), MNRJ 46 801 (67), MNR J 46819 (3), MNRJ 47011 (16),
MNRJ 47021 (30), MNRJ 47230 (1), MNRJ 47264 (4)
Oreochromis niloticus MNR J 36485 (3), MNRJ 37964 (3)