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550
Accepted by J. Armbruster: 19 Jul. 2017; published: 13 Sept. 2017
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2017 Magnolia Press
Zootaxa 4319 (3): 550
–
560
http://www.mapress.com/j/zt/
Article
https://doi.org/10.11646/zootaxa.4319.3.7
http://zoobank.org/urn:lsid:zoobank.org:pub:5ED288D1-BB13-4DEB-9B97-9E5C4902D5B8
New species of Trichomycterus (Siluriformes: Trichomycteridae) lacking pelvic
fins from Paranapanema basin, southeastern Brazil
LUZ E. OCHOA
1
, GABRIEL S. C. SILVA
1
, GUILHERME J. COSTA E SILVA
1
, CLAUDIO OLIVEIRA
1
&
ALESSIO DATOVO
2
1
Departamento de Morfologia, Instituto de Biociências, Universidade Estadual Paulista – UNESP, R Prof. Dr. Antonio C. W. Zanin, s/
n, Rubião Jr, 18618–689, Botucatu SP, Brazil
2
Museu de Zoologia da Universidade de São Paulo (MZUSP), Laboratório de Ictiologia; Av. Nazaré, 481, 04263-000, São Paulo, SP,
Brazil
Abstract
A new species of trichomycterid catfish, Trichomycterus pascuali, is described from Paranapanema basin and is distin-
guished from all congeners by the possession of five pectoral-fin rays and the absence of pelvic fin, girdle, and muscles.
Additional features further differentiate the new species from the other congeners lacking pelvic fins, T. candidus, T. cat-
amarcensis, and T. tropeiro. The identification of T. pascuali is additionally corroborated by genetic divergence based on
DNA-barcode analysis. Osteological and myological data unequivocally support the inclusion of the new species in the
Trichomycterinae and molecular analyses justify its allocation to the genus Trichomycterus rather than Eremophilus, a
trichomycterine taxon traditionally diagnosed by the lack of pelvic fins. Our genetic analysis further indicates that pelvic
fins were independently lost in E. mutisii, T. candidus, and T. pascuali.
Key words: Upper Paraná Basin, Freshwater catfish, pelvic-fin loss, taxonomy.
Introduction
Trichomycteridae is one of the most species-rich groups of freshwater fishes (de Pinna 1998) in the Neotropics
with approximately 290 valid species (Eschmeyer et al. 2017) distributed into 41 genera and eight subfamilies. The
highest species diversity of the family is concentrated in the Trichomycterinae, especially in the genus
Trichomycterus, which currently includes about 190 valid species (Eschmeyer et al. 2017), most of which are
characterized by restricted geographic distributions and high levels of endemism (de Pinna 1992; Malabarba et al.
2009).
In the last decades, studies on the diversity of genus Trichomycterus increased with the description of several
new species and the identification of smaller putative monophyletic subunits (de Pinna 1989; Costa 1992; Wosiacki
2002; Barbosa & Costa 2003a; Lima & Costa 2004; Wosiacki & de Pinna 2008, Dutra et al. 2012; Barbosa & Costa
2010; Fernández & Osinaga 2006; Datovo et al. 2012). Nevertheless, the non-monophyletic status of the genus and
the often poorly informative characters used to describe species are considered some of the main obstacles to the
understanding of the phylogenetic relationships of the Trichomycteridae (de Pinna 1998; Datovo & Bockmann
2010). New and old species descriptions of Trichomycterus are frequently characterized by ambiguous diagnoses,
vague descriptions and limited taxonomic comparison with little concern about similar species previously
described (Garcia-Melo et al. 2016). Additionally, the intra- and interspecific phenotypic variability are commonly
overlapping among possibly related species, which demand innovative approximations such as the use of
molecular tools for the identification of biological species (Hebert et al. 2003).
In a collection performed in a small tributary of the Paranapanema basin, Upper Paraná drainage, we collected
a trichomycterid species lacking pelvic fins. Morphological survey and genetic analysis based on DNA barcode
support the allocation of these specimens in the genus Trichomycterus, which is herein described.
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Material and methods
Sample collection and preservation. All specimens belonging to the new species (Table S1) were collected at the
same location. After capture, the individuals were anesthetized and euthanized using a solution of 1% benzocaine.
Specimens were preserved in 95% ethanol for molecular studies, and vouchers were fixed in 10% formaldehyde for
morphological studies. All samples used in morphological and molecular analysis were cataloged in the the
Laboratório de Biologia e Genética de Peixes, Universidade Estadual Paulista, Botucatu, Brazil (LBP) and
remaining types were deposited in the Museu de Zoologia, Universidade de São Paulo, São Paulo, Brazil
(MZUSP).
Morphological analysis. Measurements and counts were taken from the left side of the specimens.
Measurements were made from point to point, taken with a digital caliper to the nearest 0.1 mm and following
Tchernavin (1944) for the barbel lengths, Wosiacki and de Pinna (2008) for the peduncle-caudal length and depth
and Costa (1992) for the remaining measurements. Morphometrics are given as percentages of standard length
(SL), except for subunits of the head region that are expressed as percentages of head length (HL). Specimens were
cleared and double stained (c&s) according to anatomical preparations in Datovo and Bockmann (2010), which
follow the protocol of Taylor and Van Dyke (1985) with some modifications. Vertebral counts do not include those
in the Weberian apparatus and the compound caudal centrum (PU1+U1; = urostyle) is counted as a single element.
Nomenclature for laterosensory system and associated pores followed Bockmann et al. (2004).
DNA Extraction and Sequencing. Total DNA was extracted from ethanol preserved muscle samples with the
DNeasy Tissue Kit (Qiagen), following manufacturer’s instructions. Partial sequences of the gene cytochrome c
oxidase subunit I (COI) and the 16S genes were amplified using polymerase chain reaction (PCR) using the
primers FishF1 (5’-TCAACCAACCACAAA GACATTGGCAC-3’) and FishR1 (5’-
TAGACTTCTGGGTGGCCAAAGAATCA-3’) to COI and 16Sa-L 5’-ACGCCTGTTTATCAAAAACAT-3’ and
16Sb-H 5’-CCGGTCTGAACTCAGATCACGT-3’ (Palumbi 1996). Amplifications were performed in a total
volume of 12.5 μl with 1.25 μl of 10X buffer (10 mM Tris-HCl+15 mM MgCl2), 0.5 μl dNTPs (200 nM of each),
0.5 μl each 5 mM primer, 0.05 μl Platinum® Taq Polymerase (Invitrogen), 1 μl template DNA (12 ng), and 8.7 μl
ddH2O. The PCR reactions consisted of intial denaturation (4 min at 95ºC) followed by 30 cycles with a chain
denaturation (30 s at 95°C), primer hybridization (30-60 s at 54°C) and nucleotide extension 30-60 s at 72°C. All
PCR products were first visually identified on a 1% agarose gel and then purified using ExoSap-IT® (USB
Corporation) following instructions of the manufacturer. The purified PCR products were sequenced using the “Big
DyeTM Terminator v 3.1 Cycle Sequencing Ready Reaction Kit” (Applied Biosystems), purified again by ethanol
precipitation and loaded on an automatic sequencer 3130-Genetic Analyzer (Applied Biosystems) in the Instituto
de Biociências, Universidade Estadual Paulista, Botucatu, Brazil.
All individual sequences for each species were initially analyzed using the program Geneious v7.1.7
(Biomatters Ltd., Auckland, New Zealand) and the consensus sequences were obtained. All sequences were
aligned using MUSCLE (Edgar, 2004) under default parameters and the alignment was inspected by eye for any
obvious misalignments such as sequencing errors due to contamination, paralogy or pseudogenes.
Nucleotide variation, substitution patterns, and genetic distances were examined using MEGA 6.0 (Tamura et
al. 2013). To evaluate the occurrence of substitution saturation in the sequences, we estimated the index of
substitution saturation (Iss) described by Xia et al. (2003) and the rate of transitions/transversions (Xia and Lemey
2009) in DAMBE 5.2.31 (Xia 2013). The genetic divergence among specimens was estimated using COI matrix
and Kimura two parameter (K2P) distance model (Kimura 1980). To infer the congeneric relationships of the new
species, we used a concatenated matrix including two loci (cytochrome oxidase I and 16S, 1222 pb) to estimate a
maximum likelihood tree using RAxML (Stamatakis 2006) via CIPRES web portal (Miller et al. 2010). The
number of alternative runs was 100 and the analyses were performed under the model GTR+G.
Trichomycterus pascuali, new species
(Figs. 1–3, Table 1)
Holotype. MZUSP 121681, 48.8 mm SL; Brazil: São Paulo State, Municipality of Itatinga; unnamed tributary of
Tamanduá river, Paranapanema Basin, 23°13’27.06”S 48°31’45.34”W, G.S.C. Silva, G.J. Costa e Silva, L.E.
Ochoa, 14 Feb 2017.
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Paratypes. LBP 23323, 8 (20.76-46.53 mm SL), MZUSP 121682 (2 ex): 2 CS (20.76, 22.30 mm SL), 1 MS
(46.53), 1 genetype (20.76 mm, Genbank accession number: MF034463-COI, MF034462-16S); same data as
holotype.
Diagnosis. Trichomycterus pascuali is readily distinguished from its congeners by two remarkable features:
possession of five pectoral-fin rays (vs. six or more) and absence of pelvic fin, girdle, and muscles (vs. presence).
Within Trichomycterus, only T. candidus (Rio Grande basin, Brazil), T. catamarcensis (Laguna Blanca basin,
Argentina), and T. tropei ro (laguna dos Patos basin, Brazil) also lack pelvic structures. Trichomycterus pascuali
differs from these three species in having the aforementioned five pectoral-fin rays (vs. six, eight, and seven,
respectively) and seven branchiostegal rays (vs. eight, five or six, and nine, respectively). It addition, it differs from
T. candidus by the possession of anterior and posterior cranial fontanels completely separated from each other by a
broad epiphyseal bar (vs. fontanels partially or completely fused, with epiphyseal bar absent or minute and
incomplete) and 37 post-Weberian vertebrae (vs. 38–39 in T. candidus); from T. catamarcensis by the first pectoral
fin prolonged as a small filament (vs. pectoral filament absent in most specimens), 14 ribs (vs. 18–20 in T.
catamarcensis), 17–18 dorsal procurrent caudal-fin rays (vs. 12–13), and 15–16 ventral procurrent caudal-fin rays
(vs. 11–12); and from T. t rope i ro by the presence of pectoral filament (vs. absence), 17–18 dorsal procurrent
caudal-fin rays (vs. 13–14), 15–16 ventral procurrent caudal fin rays (vs. 9–10), and absence of pores i
1
and i
3
(vs.
presence).
Members of the so-called “Trichomycterus” hasemani group, a clade of highly derived Amazonian miniature
trichomycterids, also exhibit a low count of pectoral-fin rays and one of its species, “T.” anhanga, further lacks
pelvic fins. However, the “T.” hasemani group is demonstrably a new subfamily that is more closely related to the
Vandelliinae-group than to trichomycterines (de Pinna 1989; Dutra et al. 2012; Datovo 2014, Wosiacki 2002). The
nominal assignment of these species to the genus Trichomycterus is merely due to taxonomic inertia, as the formal
description of the new subfamily is still underway (by W. Wosiacki and M.C.C. de Pinna, pers. comm.). In any
event, T. pascuali differs from all species of the T. hasemani group by the relatively large body size, with up to 49.4
mm SL (vs. miniature body, with less than 18.6 mm SL); presence of an ossified neurocranial roof with two small
fontanels (vs. roof of neurocranium mostly unossified and forming a single wide fontanel); and 14 ribs (vs. 1–3).
Description. Morphometric data are given in Table 1. Body elongate, maximum total length 52.2 mm SL,
cross section of trunk roughly cylindrical at pectoral girdle and progressively more compressed towards caudal-fin.
In lateral view, dorsal profile of body straight from snout tip to interorbital region, slightly convex to dorsal-fin
origin, and slightly convex to end of caudal peduncle. Ventral surface of body convex from tip of snout to anal-fin
insertion. Caudal peduncle smoothly continues profile of trunk region, with dorsal and ventral profiles slightly
convex. Greatest body depth at dorsal-fin origin. Head wide, longer than wide, nearly trapezoidal in dorsal view.
Eyes elliptical, covered by thin and translucent skin separate from surface of eyeball. Anterior nostril larger than
posterior one, surrounded by fleshy flap of integument continuous with base of nasal barbel. Posterior nostril
partially surrounded anteriorly by thin flap of skin. Seven branchiostegal rays visible in c&s individuals. Mouth
subterminal, its corners laterally or slightly posteriorly oriented. Lower lip with conspicuous fleshy lobes along
lateral limits, lobes situated internal to base of rictal barbels. Lower lip with anterior and, to lesser degree,
anteroventral surfaces covered by small papillae. Anterior margin of upper lip rounded with numerous papillae.
Barbels relatively short and with larger bases. Nasal barbel thick always surpassing posterior margin of eye but not
reaching opercular patch of odontodes. Maxillary barbel reaching posterior tip of interopercular patch of
odontodes. Rictal barbel reaching interopercular patch of odontodes. Interopercular patch of odontodes posteriorly
elongate with 11 or 12 conic odontodes imbedded in flesh. Opercular patch of odontodes very small, with 10–12
thin odontodes with thick tips.
Cephalic laterosensory system including incomplete supraorbital and infraorbital canals. Infraorbital canal
restricted to its posterior fragment corresponding to pores i
10
and i
11
. Supraorbital pores s
1
, s
3
and s
6
. Two bilaterally
paired pores s
6
. Lateral line of trunk reduced, comprising two pores dorsal to middle of pectoral fin. Vertebra 37, ribs 14.
Pectoral fin narrow, with rounded margin. Pectoral-fin rays i,4, first one longest, unbranched, prolonged as
small filament. Dorsal fin with distal margin rounded, ii,5-7 rays (modally ii,6,i). Anal fin slightly smaller than
dorsal fin, with distal margin rounded, ii,4-5,i rays, origin at vertical through posterior portion of dorsal fin. Pelvic
fin, girdle and muscles absent. Caudal fin with distal margin truncate with rounded corners and dorsal and ventral
edges slightly convex, i,5+6,i principal rays, branched rays splitting tree times. Procurrent caudal-fin rays 17 or 18
dorsally and 15 or 16 ventrally.
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NEW TRICHOMYCTERUS IN PARANAPANEMA BASIN
FIGURE 1. Trichomycterus pascuali, MZUSP 121681, holotype 48.8 mm SL; Brazil: São Paulo State, Municipality of
Itatinga; unnamed river tributary of Tamandua river, Paranapanema basin, in ventral, lateral, and dorsal views.
Color in alcohol. Two main body-coloration patterns. First pattern (Fig. 2A) observed in specimens 20.8–48.8
mm SL. Ground color of body yellowish. Head covered by irregular brown spots and two regular longitudinal
stripes along lateral region of body, one dorsolateral and another midlateral. Ventrolateral row of spots observed at
abdominal region and caudal peduncle. Dorsal, anal and caudal fins slightly pigmented.
Second color pattern (Fig. 2B) found in specimens 49.4–52.2 mm SL. Head with some regular brown spots in
the dorsal region, gradually lighter laterally; barbels slightly pigmented with the same coloration as head spots.
Trunk yellowish with dorsal and abdominal lateral portion of the body covered by irregular spots from the lateral
head region to caudal peduncle without defined pattern. Pectoral fin unpigmented, dorsal, anal, and caudal fins rays
pigmented with the same coloration as the body spots.
Etymology. The name “pascuali” was given in honor to José Pascual Ochoa, L. Ochoa’s father.
Distribution. Known only from the type locality, a small stream tributary of Tamanduá river, Paranapanema
river, Upper Paraná, close to the town of Itatinga, São Paulo state, southeastern Brazil (Fig. 4).
Habitat, ecological notes, and conservation status. The new species was discovered in a small stream with
clear water and moderate water flow at the altitude of approximately 600 meters above sea level. The specimens
were collected under rocks and aquatic vegetation in a muddy bottom area bordered by riparian vegetation. The
region is highly impacted by agricultural activity, more specifically eucalyptus monoculture. Rivers in the region
are poor in both alpha diversity and specimen abundance. Trichomycterus pascuali was the only species of fish
found in the type locality and the number of individuals sampled was low. This species has a restricted area of
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occupancy and its vulnerability to the threats from anthropic impacts could drive the taxon to critically endangered
in a short time. According to the IUCN (2001) criteria, T. pascuali should be classified in the category vulnerable
VU D2.
TABLE 1 . Morphometric data for holotype and paratype of Trichomycterus pascuali (n=8 including the holotype). SD=
Standard deviation.
FIGURE 2. Body coloration patterns observed in Trichomycterus pascuali, LBP 23323, (A) 45.4 mm SL, (B) 52.2 mm SL.
Holotype Range Mean SD
Standard length (mm) 48.8 31.37-49.36 40.4 6.1
Percents of Standard Length
Head length 15.5 14.30-16.77 15.8 0.9
Predorsal length 67.3 65.24-68.31 67.2 1.1
Preanal length 73.3 68.27-73.81 71.8 1.9
Scapular girdle width 10.8 10.68-12.08 11.4 0.5
Pectoral-fin length 8.2 8.21- 12.56 10.4 1.5
Caudal peduncle length 17.5 14.49-21.33 17.8 1.9
Caudal peduncle depth 11.5 9.66-11.53 10.7 0.7
Body depth 13.7 8.19-14.87 12.5 2.0
Length of dorsal-fin base 10.9 6.57-13.13 10.3 1.8
Length of anal-fin base 8.4 5.77-9.54 8.1 1.1
Percents of Head Length
Head width 96.0 80.66-96.03 91.2 6.4
Nasal barbel length 66.1 64.20-79.92 68.9 4.9
Maxillary barbel length 55.4 55.36-76.31 62.7 7.1
Rictal barbel length 66.0 53.30-71.43 63.2 6.9
Snout length 12.9 7.14-17.35 12.8 3.4
Interorbital 27.7 25.27-31.86 29.0 2.2
Mouth width 33.0 26.72-47.08 35.3 5.9
Supra-orbital pore distance 11.7 10.52-16.07 12.9 1.9
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Genetic identification. Aligned sequences were obtained for 573 bp of COI from the reference set of 11 tissue
samples of trichomycterines, the translation of sequences did not result in stop codons, indicating the amplification
of functional domains. The nucleotide composition of the COI matrix was 29.8% (T), 26.8% (C), 25.1% (A),
18.3% (G), with a proportion of 400 invariables sites, 173 variables sites and 113 parsimony informative sites. The
overall mean genetic distance based in the COI gene was 0.138±0.011. The genetic divergence between species
was superior to 3% and the results show concordance with the morphological identification. The pairwise species
comparison ranged from 0.5 to 24.1% (Table 2) and the divergence genetic within the Trichomycterus species was
0.097±0.009. Trichomycterus pascuali presented a genetic divergence with its congeneric species from 9.3 to
11.7%, with T. candidus and T. brasiliensis respectively. the genetic divergence between E. mutisii and T. pascuali
was (10.8%) similar with other species of the Trichomycterus, the maximum likelihood analysis using a
concatenated matrix with COI and 16S genes (1122 pb, 30.2%(T), 25.0 (C), 23.7%(A), 21.1%(G), with 112
conserved sites, 254 variables sites and 145 parsimony informative sites) shows that T. pascuali is more related
with T. inhering and T. variegatus than other genera included in this analysis with a bootstrap support of 77% (Fig.
5).
FIGURE 3. Dorsolateral view of head of Trichomycterus pascuali, LBP 23323 (51.2 mm SL); arrow indicates levator internus
IV originating from the dorsal face of posttemporo-supracleithrum.
Discussion
Osteological, myological and meristic characters have been used to classify species of the Trichomycteridae (e.g.,
Baskin 1973; de Pinna 1989; Datovo & Bockmann 2010). The subfamily Trichomycterinae belongs to a well-
supported clade that includes all trichomycterid subfamilies except Copionodontinae and Trichogeninae (de Pinna
1992, 1998; Wosiacki 2002; Datovo & Bockmann 2010). In traditional classifications, the Trichomycterinae
basically includes taxa lacking the synapomorphies for the TSVSG clade, a lineage formed by the most specialized
trichomycterid subfamilies (Sarcoglanidinae, Glanapteryginae, Tridentinae, Stegophilinae, and Vandelliinae; de
Pinna 1998). According to the most recent phylogenetic hypothesis based on myological data and proposed by
Datovo & Bockmann (2010), the only character that may potentially support the monophyly of the
Trichomycterinae is the posterior part of the levator internus IV originating from the dorsal face of the
posttemporo-supracleithrum. Trichomycterus pascuali simultaneously shows this synapomorphic condition (Figure
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3) and lacks the synapomorphies for the TSVSG clade, so that its allocation in the Trichomycterinae is
unequivocal.
FIGURE 4. A. Map showing the type localities of Trichomycterus pascuali, 23°13’27.06”S 48°31’45.34”W, and congeners
lacking pelvic fins and girdle. B. Type locality of T. pascuali, unnamed tributary of Tamanduá river, Paranapanema basin.
Whereas the genus Trichomycterus is a non-monophyletic group that includes those trichomycterines lacking
the diagnostic features of the remaining genera, Eremophilus has been traditionally distinguished by its lack of
pelvic fins (Eigenmann 1918; Myers 1944). However, several studies indicate that pelvic fins were lost
independently many times during the trichomycterid radiation and that species with such a feature are often more
closely related to taxa with pelvic fins (de Pinna 1988, 1989; Barbosa & Costa 2003b; de Pinna & Wosiacki 2003;
Datovo 2014). As a result, the genus Eremophilus has been eroded by the removal of some species originally
assigned to this genus, namely E. candidus and E. camposi (Miranda-Ribeiro 1949; 1957) that were transferred to
Trichomycterus (T. candidus, de Pinna & Wosiacki 2003) and Listrura (L. camposae, Glanapteryginae; de Pinna
1988), respectively. In the last decade, two new trichomycterines lacking pelvic fins and girdles were described and
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allocated to the genus Trichomycterus following similar reasoning: T. catamarcensis (Fernandéz & Vari 2000) from
Catamarca, Argentina, and T. tropei ro (Ferrer & Malabarba 2011) from rio das Antas, laguna dos Patos system. As
a result, Eremophilus is currently a monotypic genus that includes only its type species, E. mutisii, from the Bogotá
River, Colombia.
The allocation of Trichomycterus pascuali to the genus Trichomycterus is supported by DNA-barcode and
phylogenetic analyses that indicate the new species as being more closely related to other species of this genus than
to Eremophilus mutisii (Fig. 4, Table 2). These analyses also include T. candidus, which is the congener lacking
pelvic fins occurring in closest geographical proximity (Rio Grande basin, Upper Paraná) to T. pascuali (Rio
Paranapanema basin, Upper Paraná). Interestingly, the two species are not resolved as sister taxa, indicating that
their pelvic structures were lost independently, thus reinforcing the view that such a feature has an erratic
distribution across the Trichomycteridae.
TABLE 2. Pairwise comparison of nucleotide divergence (K2P distances) at COI from 10 species representatives
of four Trichomycterinae genera
Comparative material
Bullockia maldonadoi (Eigenmann 1920): LBP 3112 (8), municipality of Monte Aguila, rio Bio Bio, La laja basin,
Chile. Eremophilus mutisii Humboldt 1805: ANSP 11306, municipality of Ubaté, Laguna de Fuquene, Colombia;
MZUSP 35409 (1 c&s), no locality data. Ituglanis parahybae (Eigenmann 1918): LBP 10730 (3) municipality of
Conceição do Macabu, rio Macabu, Paraíba do Sul basin, Brazil. Scleronema minutum (Boulenger 1891): LBP
3310 (11) municipality of Pelotas, arroio dos Corrientes, Atlántico, Brazil. Trichomycterus areolatus Valenciennes,
1846: LBP 3118 (29) municipality of Cararrehue, Lago Villarica, rio Maichin, Chile. Trichomycterus brasiliensis
Lütken, 1874: LBP 11778 (2) municipality of São Roque de Minas, Córrego da Cachoeira, São Francisco basin,
Brazil. Trichomycterus candidus (Miranda Ribeiro, 1949): LBP 11630 (18) municipality of Uberaba, riacho
Grotão, Paraná basin, Brazil; MZUSP 55641 (4, 3 c&s), municipality of Delfinópolis, tributary of rio Grande,
Paraná basin, Brazil; MZUSP 55642 (3), municipality of Delfinópolis, tributary of rio Grande, Paraná basin,
Brazil; MZUSP 109234 (5), municipality of Carmo do Rio Claro, tributary of rio Grande, Paraná basin, Brazil.
Trichomycterus iheringi (Eigenmann, 1917): LBP 4512 (7), municipality of Santo André, rio Paranapiacaba,
Paraná basin, São Paulo, Brazil. Trichomycterus tropeiro Ferrer & Malabarba 2011: MZUSP 108296 (1 paratype),
MZUSP 108297 (1 paratype), MZUSP 108298 (1 paratype), municipality of São José dos Ausentes, rio das Antas,
I. parahybae T. areolatus B. maldonadoi S. minutum E. mutisii T. brasiliensis T. candidus T. pascuali T. variegatus
Ituglanis parahybae
Trichomycterus
areolatus 0.145
Bullockia maldonadoi 0.144 0.047
Scleronema minutum 0.142 0.149 0.150
Eremophilus mutisii 0.130 0.125 0.130 0.135
Trichomycterus
brasiliensis 0.132 0.134 0.135 0.125 0.100
Trichomycterus
candidus 0.125 0.129 0.133 0.132 0.101 0.032
Trichomycterus
pascuali 0.162 0.150 0.150 0.108 0.117 0.095 0.093
Trichomycterus
variegatus 0.183 0.138 0.132 0.112 0.127 0.100 0.106 0.057
Trichomycterus
inheringi 0.182 0.138 0.132 0.114 0.131 0.104 0.111 0.062 0.005
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Brazil. Trichomycterus variegatus Costa 1992: LBP 10289 (3), municipality of São Roque de Minas, Córrego do
Lavapés, tributary rio São Francisco, Brazil.
FIGURE 5. Maximum likelihood tree built with concatenated matrix using cytochrome oxidase sub-united I (COI)
and 16S genes, showing the relationships of Trichomycterus pascuali within Trichomycterinae. Numbers on
branches of tree denote bootstrap (B) values.
Acknowledgements
This research was supported by the Brazilian agencies FAPESP (Fundação de Amparo à Pesquisa do Estado de São
Paulo, proc. 2014/06853-8 and proc. BEPE 2015/13382-4 to LEO) and CNPq (Conselho Nacional de
Desenvolvimento Científico e Tecnológico, proj. 402866/2014-2 to GJCS)
References
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SUPPLEMENTARY 1. Voucher information and GenBank accession numbers for the samples analyzed in the present study
Genus Species Specimen Catalog # GenBank
COI 16S
Copionodon Copionodon pecten LBP 17357 KY857929 KY807205
Bullockia Bullockia maldonadoi LBP 3112 KY857926 KY807202
Scleronema Scleronema minutum LBP 3310 KY857957 KY807234
Ituglanis Ituglanis parahybae LBP 10730 KY807219
Eremophilus Eremophilus mutisii ANSP 11306 KY857931 KY807207
Trichomycterus Trichomycterus pascuali LBP 23323 MF034463 MF034462
Trichomycterus areolatus LBP 3118 KY857964 KY807241
Trichomycterus brasiliensis LBP 11778 KY857993 KY807271
Trichomycterus candidus LBP 11630 KY857965 KY807242
Trichomycterus iheringi LBP 4512 KY858008 KY807286
Trichomycterus variegatus LBP 10289 KY857991 KY807269
