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Accepted by M.R. de Carvalho: 7 Mar. 2016; published: 6 May 2016
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4109 (2): 227
–
238
http://www.mapress.com/j/zt/
Article
227
http://doi.org/10.11646/zootaxa.4109.2.7
http://zoobank.org/urn:lsid:zoobank.org:pub:3CDD7AA6-1511-4396-AAEE-EF80471EACB5
A new species of Hisonotus (Siluriformes: Otothyrinae) from the upper rio
Paraná and rio São Francisco basins, Brazil
FÁBIO F. ROXO
1,4
, GABRIEL S.C. SILVA
1
, BRANDON T. WALTZ
2
& JORGE E.G. MELO
3
¹Universidade Estadual Paulista, Departamento de Morfologia, Laboratório de Biologia e Genética de Peixes, Botucatu, São Paulo
State, Brazil
2
University of Louisiana at Lafayette, Department of Biology, Lafayette, LA, USA
3
Pontificia Universidad Javeriana, Departamento de Biología, Facultad de Ciencias, Unidad de Ecología y Sistemática (UNESIS),
Bogotá, Cundinamarca State, Colombia
4
Corresponding author. E-mail: roxoff@hotmail.com.br
Abstract
A new species of Hisonotus is described from the upper rio Paraná and rio São Francisco basins. The new species is dis-
tinguished from congeners by (1) completely exposed abdomen with no development of dermal plates (with the exception
of extremely small platelets present near the urogenital pore in some specimens) and the combination of the following
characters: (2) lack of a conspicuous tuft of enlarged odontodes on posterior tip of parieto-supraoccipital; (3) rectangular
dorsal-fin spinelet; (4) complete mid-lateral plate series; (5) higher number of vertebrae, 29–30; (6) dark brown coloration
on caudal fin with one pair of circular hyaline colored regions at center of both lobes; (7) absence of broad light stripes on
dorsolateral surface of head; (8) odontodes not forming longitudinally aligned rows on head and trunk; and (9) apex of
teeth yellowish in color.
Key words: Cascudinhos, Catfishes, Ichthyology, Neotropical fishes, taxonomy
Introduction
Within the subfamily Otothyrinae (family Loricariidae), the genus Hisonotus (type species H. notatus Eigenmann
& Eigenmann, 1889) includes 35 species widely distributed throughout the hydrographic systems of South
America. The distribution of Hisonotus ranges from Amazonian drainages in Brazil (Britski & Garavello, 2007;
Silva et al., 2014) to tributaries of the La Plata basin in Argentina (Azpelicueta et al., 2004) and Atlantic coastal
drainages, (Carvalho & Reis, 2011).
In the last 10 years, 21 species of Hisonotus have been described (Britski & Garavello, 2007; Carvalho et al.,
2008; Carvalho & Reis, 2009; Carvalho & Reis, 2011; Carvalho & Datovo, 2012; Martins & Langeani, 2012; Roxo
et al., 2013; Roxo et al., 2014a, Silva et al., 2014). The polyphyletic nature of the genus has been described in
previous molecular (e.g., Chiachio et al., 2008; Cramer et al., 2011; Roxo et al., 2014b) and morphological studies
(e.g., Martins et al., 2014), with the species H. acuen, H. bocaiuva and H. chromodontus distantly related to the
type species, H. notatus. Furthermore, Roxo et al. (2015) recently described the genus Curculionichthys, comprised
of five species previously included within Hisonotus (Curculionichthys insperatus, C. luteofrenatus, C. oliveirai,
C. paresi and C. piracanjuba).
A recent collection expedition to tributaries of the upper rio Grande and rio São Francisco basins has led to the
discovery of a novel species within the genus Hisonotus. The description of the newly discovered taxa is described
herein.
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Material and methods
Measurements and counts were taken preferentially from the left side of specimens. Measurements were made as
point to point distances to the nearest 0.01 mm with a digital caliper. Abbreviations and nomenclature of bones
followed Carvalho & Reis (2009) and Schaefer (1997). Vertebral counts include the five vertebrae that comprise
the Weberian apparatus and the compound caudal centrum (PU1 + U1) as one element. Dorsal fin-ray counts
include the spinelet as the first unbranched ray. Specimens were cleared and double stained (C&S) according to the
method of Taylor & Van Dyke (1985). Institutional acronyms follow Fricke & Eschmeyer (2015). Specimens are
deposited at the Academy of Natural Sciences (ANSP), Philadelphia, PA, USA; Laboratório de Biologia e Genética
de Peixes (LBP), Universidade Estadual Paulista, Botucatu, Brazil; Museo de Pontifica Universidad de Javeriana
(MPUJ), Bogota, Colombia; Museu de Zoologia, Universidade de São Paulo (MZUSP), São Paulo, Brazil; and
Coleção Ictiológica do Nupelia (NUP), Universidade Estadual de Maringá, Maringá, Brazil. Zoological
nomenclature follows the International Code of Zoological Nomenclature (International Commission on
Zoological Nomenclature 1999).
Hisonotus alberti n. sp.
(Figs. 1–4, Table 1)
Holotype. MZUSP 118845 (male, 33.5 mm SL), Brazil, Minas Gerais State, municipality of São Roque de Minas,
ribeirão das Posses, upper rio Paraná basin, 20°17'15"S, 46°34'53"W, 19 May 2011, col. J.A. Senhorini, P.S.
Cecarelli, J.O. Junqueira, R. Devidé, R. Rocha, M.N. Mehanna.
Paratypes. All from Minas Gerais State, Brazil (67 specimens). ANSP 200148, (2 females, 32.1–34.0 mm SL,
2 males, 29.8–31.7 mm SL), municipality of São Roque de Minas, ribeirão das Posses, upper rio Paraná basin,
20°14'54"S, 46°38'24"W, 10 April 2010, col. J.A. Senhorini, P.S. Cecarelli, J.O. Junqueira, R. Devidé, R. Rocha,
M.N. Mehanna. LBP 10245 (6 males, 28.9–30.5 mm SL), municipality of São Roque de Minas, córrego Luciano,
rio São Francisco basin, 20°18'44"S, 46°31'51"W, 08 April 2010, col. J.O. Junqueira, R. Devidé, R. Rocha, M.N.
Mehanna. LBP 10266 (9 females, 25.6–34.9 mm SL, 11 males, 28.5–31.8 mm SL, 7 C&S, sex and measurements
not determined), municipality of São Roque de Minas, ribeirão das Posses, upper rio Paraná basin, 20°14'54"S,
46°38'24"W, 10 April 2010, col. J.A. Senhorini, P.S. Cecarelli, J.O. Junqueira, R. Devidé, R. Rocha, M.N.
Mehanna. LBP 11847 (2 females, 24.3–33.4 mm SL, 2 males, 30.4–30.3 mm SL) collected with holotype. LBP
17388 (6 females, 24.9–33.4 mm SL, 2 males, 29.3–30.1 mm SL), municipality of Oliveira, córrego Caxambú,
upper rio Paraná basin, 20°47'52"S, 44°52'35"W, 19 February 2013, col. R. Devidé, G.S.C. Silva, C. Araya, V.F.
Sene. MPUJ 8199, (2 females, 27.6–31.4 mm SL, 4 males, 28.5–30.5 mm SL), municipality of São Roque de
Minas, ribeirão das Posses, upper rio Paraná basin, 20°14'54"S, 46°38'24"W, 10 April 2010, col. J.A. Senhorini,
P.S. Cecarelli, J.O. Junqueira, R. Devidé, R. Rocha, M.N. Mehanna. NUP 9437 (2 females, 25.9–32.4 mm SL),
municipality of Oliveira, córrego Caxambú, upper rio Paraná basin, 20°48'06"S, 44°52'41"W, 23 July 2009, col. F.
Correa. NUP 9440 (3 females, 32.7–34.6 mm SL, 2 males, 30.2–31.9 mm SL), municipality of Oliveira, córrego
Caxambú, upper rio Paraná basin, 20°48'06"S, 44°52'41"W, 14 January 2009, col. F. Correa. NUP 9441 (4 males,
27.4–30.0 mm, 1 C&S, sex and measurements not determined), municipality of Oliveira, córrego Caxambú, upper
rio Paraná basin, 20°48'06"S, 44°52'41"W, 14 January 2009, col. F. Correa.
Diagnosis. The new species can be distinguished from all congeners by possessing an exposed abdomen (i.e.,
area bounded by cleithral region and urogenital opening) with no dermal plate development (Fig. 2a) and with the
exception of extremely small platelets present near urogenital opening in some specimens (Fig. 2b) (vs. abdomen
partially or completely covered by plates, Figs. 2c,d, respectively). Moreover, Hisonotus alberti differs from H.
bocaiuva, H. carreiro, H. francirochai, H. iota, H. leucophrys and H. prata by the absence of a conspicuous tuft of
enlarged odontodes on posterior tip of parieto-supraoccipital (vs. presence); from H. chromodontus, H. acuen, H.
vespuccii and H. bockmanni by possessing a rectangular dorsal-fin spinelet (vs. a functional V-shaped spinelet);
from H. leucophrys, H. megaloplax, H. montanus, H. nigricauda, H. laevior, H. notopagos, H. carreiro, H. prata,
H. vireo, H. brunneus, H. heterogaster, and H. bocaiuva by possessing a complete mid-lateral plate series (i.e.,
plate series not truncated), Fig. 3b (vs. mid-lateral plate series incomplete and truncated); from H. megaloplax, H.
montanus, H. nigricauda, H. carreiro, H. prata, H. brunneus, H. iota, H. acuen, and H. bocaiuva by larger number
of vertebrae (29–30 vs. 25–28); from H. nigricauda and H. prata, H. leucofrenatus, and H. notatus by possessing a
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DESCRIPTION OF A NEW SPECIES OF HISONOTUS
dark brown pigmented caudal fin with paired rounded regions of hyaline color near the center of both caudal lobes
(Fig. 1) (vs. almost completely dark caudal fin); from H. leucophrys and H. montanus by absence of broad light
stripes on dorsolateral surface of head (vs. presence of broad stripes on dorsolateral surface of head); from H.
vespuccii by possessing non longitudinally aligned rows of odontodes on head and trunk (vs. odontodes forming
longitudinally aligned rows); and from H. chromodontus by possessing yellow teeth apices (vs. reddish-brown
teeth).
Description. Morphometric and meristic data are presented in Table 1. Small body size, holotype 33.5 mm SL;
paratypes 26.3−34.1 mm SL. Body slender, dorsoventrally depressed, widest near cleithrum. Dorsally, tip of snout
rounded, short (45.0−50.2% of HL), and depressed directly in front of each nostril on dorsal surface. Greatest body
depth at dorsal fin origin (14.9−19.7% of SL) and greatest body width at cleithral region (22.3−26.1% of SL),
gradually decreasing towards snout and caudal fin. In lateral view, dorsal profile of head angled upwards linearly
from tip of snout to interocular area, shallowly concaved from interocular region to dorsal fin insertion. Dorsal
profile of trunk nearly straight, descending from base of dorsal-fin origin to caudal peduncle. Ventral profile
slightly concave from snout tip to anal-fin origin, linear from anal-fin origin to caudal peduncle.
FIGURE 1. Hisonotus alberti, holotype, MZUSP 118845, male, 33.5 mm SL, from ribeirão das Posses, upper rio Paraná basin
in municipality of São Roque de Minas, Brazil.
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FIGURE 2. Cleared and stained images showing the variation in abdominal plates. (a) Hisonotus alberti, LBP 10266, 32.1 mm
SL, abdomen completely exposed, with absence of dermal plates; (b) H. alberti, LBP 10266, 31.5 mm SL, abdomen with
extremely small plate (black arrow) near the urogenital opening; (c) H. chromodontus, LBP 7948, 23.1 mm SL, abdomen
partially covered by plates; (d) H. chromodontus, LBP 7948, 27.2 mm SL, abdomen completely covered by plates.
Head without conspicuous crests, except a small tuft of enlarged odontodes on posterior tip of the parieto-
supraoccipital (more evident in juvenile specimens). Anterior margin of snout completely covered by dorsal and
ventral series of odontodes; odontodes on tip of snout larger than on rest of head. Snout tip with delicate band
lacking odontodes (not present in some specimens). Odontodes on head and trunk well defined, not forming
longitudinal rows. Eyes small (14.1−20.9% of HL), dorsolaterally positioned. Lips round and papillose; papillae
uniformly distributed on base of dentary and premaxilla, tapering distally. Lower lip larger than upper lip; lower
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DESCRIPTION OF A NEW SPECIES OF HISONOTUS
and upper lip margins fringed. Maxillary barbel present and poorly developed. Teeth slender and bicuspid; medial
cusp larger than lateral cusp. Left premaxillary teeth 14−22 (mode 20). Left dentary teeth 13−21 (mode 18).
Dorsal fin ii,7; dorsal-fin spinelet short and laterally extended (Fig. 3a); dorsal-fin locking mechanism not
functional. Dorsal-fin origin posterior to pelvic-fin origin. Tip of branched dorsal-fin rays reaching vertical line
through center of anal-fin base. Pectoral fin i,6; tip of longest pectoral-fin ray almost reaching vertical line through
center of horizontal pelvic-fin length when depressed. Pectoral axial slit present between pectoral-fin base and
lateral process of cleithrum. Anteroventral margin of pectoral spine possessing odontodes. Pelvic fin i,5; tip not
exceeding anal-fin origin when depressed (in both sexes). Males with soft tissue flap along dorsal margin of
unbranched pelvic-fin ray. Anal fin i,5; tip of unbranched anal-fin ray reaching 7th and 8th plate from anal-fin
origin. Caudal fin i,14,i; distal margin forked. Cross-section of caudal peduncle near ellipsoid. Adipose fin absent.
Total vertebrae 29 (3 C&S)−30 (1 C&S).
FIGURE 3. Hisonotus alberti, paratype, LBP 10266, 32.1 mm SL, sex not determined. (a) Dorsal view illustrating the
rectangular shape of the spinelet; (b) Lateral view illustrating lateral trunk plates; (c) dorsal and (d) lateral views illustrating the
configuration of cranial bones and dermal plates of the head.
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DESCRIPTION OF A NEW SPECIES OF HISONOTUS
Body mostly covered by dermal plates. Ventral margin of head, small area around pectoral and pelvic fin
origins, narrow band along dorsal-fin base, and abdominal region all lacking plates (Fig. 2a); minute platelets on
pre-anal area rarely present (Fig. 2b). Cleithrum and coracoid partially to completely exposed in some specimens.
Arrector fossae partially enclosed by ventral lamina of coracoids. Lateral side of body completely covered by
plates; mid-dorsal and mid-ventral series of plates well developed, reaching center of caudal peduncle; median
plates continuous through median portion of body, with 1 or 2 plates unperforated prior to termination of plate
series (Fig. 3b).
Details of osteological features of the head are shown in Fig. 3c in dorsal view. Snout tip formed by one square
rostral plate (r). Parallelogram-shaped nasal plate (n) forming medial border of nostril and extending anteriorly and
medially with several prenasal plates (pn). Prenasal plates formed by three to four polyhedric and variable plates,
extending posteriorly to nostrils and nasal plates, anteriorly with rostral plate (n), and laterally with first postrostral
plate. Prenasals present between nares comprised of small platelets and one large triangular shaped plate.
Posterodorsal head plates consisting of compound pterotic (cpt), parieto-supraoccipital (soc), frontal (f), prefrontal
(pf) and sphenotic (sp) plates. Compound pterotic (cpt) plates bearing relatively large and irregularly distributed
fenestra. Details of osteological features of the head are shown in Fig. 3d in lateral view. First posterior rostrum
plate (pr1) rectangular in shape; Second posterior rostrum plate (pr2) and third posterior rostrum plate (pr3) larger
than pr1, triangular and roughly pentagonal in shape, respectively. Complete series of canal-bearing infraorbital
plates (io1-io5); io1 positioned between pr1 and pr2; io2 positioned between each nostril and pr2; pr3, pr4 and pr5
forming ventral and posterior limit of orbit. Canal-bearing preopercle (pop) elongate, positioned anterior to pr3,
dorsal to io4 and io5, posterior to cp2, and ventral to first subocular check plate (cp1). First and second subocular
cheek plates (cp1−cp2) and opercle (op) form posteriolateral margin of head.
Coloration in alcohol. Dorsal surface dark brown on head to light brown on margins of trunk (juveniles
lighter than adults). Ventral surface light brown to yellow in juveniles. Body and fins covered by scattered
chromatophores, which are more visible on dorsal and lateral portions of head (Fig. 1). Caudal fin black from fin
base to tip of rays, except extended hyaline area on dorsal lobe and hyaline spot on ventral lobe.
Sexual dimorphism. Males possess urogenital papillae and soft-tissue flaps along the shaft of the unbranched
pelvic fin-ray, conditions that are not observed in females. Narial openings appear to be wider in males than
females (mean: 12.4 % of HL in males vs. 10.7 % of HL in females).
Distribution and Habitat. The new species Hisonotus alberti is known from two tributaries of the upper rio
Paraná basin: ribeirão das Posses and córrego Caxambú, and one tributary of the upper rio São Francisco basin at
córrego Luciano (Fig. 4). The new species is found to be associated with marginal vegetation.
FIGURE 4. Distribution map of Hisonotus alberti in the upper rio Paraná basin (red) and upper rio São Francisco basin
(yellow). The green dot represents the type locality at ribeirão das Posses, municipality of São Roque de Minas, 20°17'15"S,
46°34'53"W. The blue dots represent paratype localities.
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Etymology. The specific epithet “alberti” is a patronym honoring professor James S. Albert, from the
University of Louisiana at Lafayette (UL) in recognition of his dedication and contributions to the studies of
Neotropical freshwater fishes.
Discussion
The new species is included in the genus Hisonotus based on the synapomorphy of the presence of enlarged
odontodes on rostrum and thickened plates forming the lateral rostral margin. However, Britski and Garavello
(2007) argue that the characters used by Schaefer (1998) to distinguish Hisonotus from other genera of Otothyrinae
need to be redefined. The utilization of poor diagnostic characters has resulted in several species being included in
the genus Hisonotus that are not closely related with the type species, H. notatus (see works of Cramer et al., 2011;
Martins et al., 2014; Roxo et al., 2014b). Moreover, despite the previously improved studies designed to better
elucidate the evolutionary history of Hisonotus, a more detailed phylogenetic analysis has yet to be executed. It is
imperative that a more robust analysis including more representatives of Otothyrinae is performed in order to better
understand the phylogenetic relationships among members of Hisonotus. It would be particularly helpful to analyze
the relationship between Otothyropsis and Hisonotus, genera that exhibit similar/shared diagnostic characters
(Lippert et al., 2014).
Hisonotus alberti possesses two characters used to diagnose the genus Otothyropsis as proposed by Calegari et
al. (2011): (1) an elongated posterior extension of the compound pterotic, which forms the dorsal margin of an
augmented lateral opening of the swimbladder capsule (characters 8 and 12 of Schaefer, 1998); and (2) the
truncation of the mid-dorsal series of lateral plates between the dorsal and the caudal fins. However, these two
character are present in several other species of Hisonotus. The first character is present in H. depressicauda and H.
francirochai, and the second character is present in almost all species of Hisonotus. Furthermore, Lippert et al.
(2014) emphasized that the mid-dorsal series of lateral plates is truncated more anteriorly in Hisonotus than in
species of Otothyropsis; however, the authors do not present a specific limit for the plate series. According to the
authors, the character state is similar to that observed in Otothyropsis marapoama, H. depressicauda, H.
francirochai, H. heterogaster, H. iota, H. laevior, H. leucophrys, H. megaloplax, H. montanus, H. notatus, and H.
vireo.
In a previous phylogenetic analysis based on a mitochondrial and nuclear DNA dataset, most species of
Hisonotus distributed throughout Southern of Brazil (i.e., H. armatus, H. taimensis, H. cf. charrua, H. leucophrys,
H. laevior, H. notopagos, H. leucofrenatus, H. nigricauda, H. ringueleti, H. prata, H. carreiro, H. megaloplax, H.
montanus, H. iota and H. aky), Otothyropsis marapoama (type species of Otothyropsis), and some species of the
genera Eurycheilichthys (i.e., Eurycheilichthys sp. 1 from arroio Jaboticaba, in rio Grande do Sul State) and
Epactionotus (i.e., E. bilineatus, E. itaimbezinho and E. gracilis) were grouped in a large clade closely related to
the type species of Hisonotus (H. notatus) (Roxo et al. 2014b). Considering the previous phylogenetic and
taxonomic results, we designate the new species, H. alberti, as a member of the genus Hisonotus. It is possible that
future studies of Otothyropsis will sync and synonymize the genus with Hisonotus.
Despite the questionable validity of Otothyropsis, the new species Hisonotus. alberti can be distinguished from
species of Otothyropsis by possessing and exposed abdomen with no development of dermal plates (Fig. 2a), with
the rare exception of extremely small platelets present near urogenital opening (Fig. 2b) (vs. the abdomen partly or
completely covered by plates, Figs. 2c,d, respectively). Moreover, H. alberti differs from species of Otothyropsis,
except O. alicula and O. polyodon, by possessing a rectangular-shaped dorsal fin-spinelet (vs. dorsal-fin triangular
or quadrangular); from O. alicula by possessing a dark brown caudal fin with one pair of rounded hyaline regions
at the center of both lobes (vs. a pigmented blotch occupying the proximal half of the caudal fin); and from O.
polyodon by the absence of a raised crest of enlarged odontodes on the supraoccipital (vs. presence of raised crest
of enlarged odontodes on the supraoccipital).
Comparative material
All specimens are from Brazil unless stated otherwise.
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DESCRIPTION OF A NEW SPECIES OF HISONOTUS
Curculionichthys insperatus: LBP 4945, 5, 27.3−28.5 mm SL, 2 c&s, 28.2−29.9 mm SL, Botucatu, São Paulo
State; LBP 6770, 5, 25.1−28.2 mm SL, 3 c&s, 20.0−27.0 mm SL, ribeirão Cubatão, Marapoama, São Paulo
State; LBP 13336, 1 c&s, 26.0 mm SL, rio Capivara, Botucatu, São Paulo State; LBP 13337, 2 c&s,
27.4−28.6 mm SL, rio Araquá, Botucatu, São Paulo State; MZUSP 22826, 1, 25.4 mm SL, paratype, córrego
Água Tirada, Três Lagoas, Minas Gerais State; MZUSP 24832, 1, 23.8 mm SL, paratype, rio Corumbataí,
Corumbataí, São Paulo State; MZUSP 78957, 1, 29.6 mm SL, holotype, rio Capivara, Botucatu, São Paulo
State; MZUSP 78960, 31, 12.6−26.0 mm SL, paratypes, 5 c&s, 22.7−24.7 mm SL, rio Pardo, Botucatu, São
Paulo State; MZUSP 78965, 10, 15.6−28.6 mm SL, paratypes, rio Araquá, Botucatu, São Paulo State;
MZUSP 78968, 5, 24.1−27.3 mm SL, paratypes, córrego da Figueira, Lins, São Paulo State.
Curculionichthys oliveirai: MZUSP 115061, 1, 26.4 mm SL, holotype, ribeirão Cambira, tributary of rio Ivaí,
Cambira, Paraná State; LBP 13332, 1 23.2 mm SL, 1 c&s, 23.7 mm SL, paratype, rio Mourão, Campo
Mourão, Paraná State; LBP 17578, 5, 25.4−30.4 mm SL, paratypes, rio Mourão, between Engenheiro
Beltrão and Quinta do Sol, Paraná State; NUP 3578, 15, 24.7−28.1 mm SL, 2 c&s, 25.5−27.6 mm SL,
paratypes, ribeirão Salto Grande, Maria Helena, Paraná State.
Curculionichthys paresi: MZUSP 115062, 1, 26.2 mm SL, holotype, riacho Águas Claras, Santo Afonso, Mato
Grosso State; LBP 13351, 9, 14.7−24.3 mm SL, paratype, riacho Águas Claras, Santo Afonso, Mato Grosso
State; LBP 13352, 1, 23.7 mm SL, paratype, riacho Águas Claras, Santo Afonso, Mato Grosso State; NUP
10928, 2, 23.2−24.2 mm SL, paratype, 2 c&s, 23.6–24.2 mm SL, riacho Águas Claras, Santo Afonso, Mato
Grosso State; NUP 10976, 3, 16.7−20.5 mm SL, paratype, riacho São Jorge, Santo Afonso, Mato Grosso
State.
Curculionichthys piracanjuba: LBP 17256, 9, 17.2−26.3 mm SL, 1 c&s, 27.1 mm SL, córrego sem nome,
Morrinhos, Goiás State; NUP 5059, 1, 24.7 mm SL, córrego Posse, Anápolis, Goiás State; MZUSP 110491,
3, 17.5−24.4 mm SL, paratypes, rio Quente, Marcelãnia, Goiás State; NUP 10979, 3, 21.4−21.8 mm SL,
ribeirão Bocaina, Piracanjuba, Goiás State.
Hisonotus acuen: MZUSP 115350, 1, 25.9 mm SL, holotype, tributary of rio Toguro, Querência, Mato Grosso
State; LBP 15755, 16, 19.5–26.0 mm SL, paratypes, tributary of rio Suiá-Missu, ribeirão Cascalheira, Mato
Grosso State; LBP 16274, 27, 20.2–29.1 mm SL, 2 c&s, 23.6−24.2 mm SL, paratypes, tributary of rio
Culuene, Gaúcha do Norte, Mato Grosso State; LBP 16275, 29, 16.7–25.2 mm SL, 2 c&s, 19.3−20.8 mm
SL, paratypes, tributary of rio Feio, Querência, Mato Grosso State; LBP 16278, 12, 18.8–25.1 mm SL, 2
c&s, 26.8−27.1 mm SL, paratypes, córrego Xavante, Primavera do Leste, Mato Grosso State.
Hisonotus aky: MHNG 2643.039, 2, 33.1−34.2 mm SL, paratypes, arroio Fortaleza, Argentina.
Hisonotus armatus: MZUSP 93884, 5, 37.6–44.4 mm SL, paratypes, arroio Arambaré, Pedro Osório, Rio Grande
do Sul State.
Hisonotus bocaiuva: MZUSP 112204, 1, 24.2 mm SL, holotype, córrego Cachoeira, Bocaiúva, Minas Gerais State;
LBP 9817, 9, 3 c&s, 18.3−23.2 mm SL, paratypes, córrego Cachoeira, Bocaiúva, Minas Gerais State.
Hisonotus brunneus: MZUSP 104947, 4, 37.2−41.3 mm SL, paratypes, rio Passo Novo, Cruz Alta, Rio Grande do
Sul State.
Hisonotus carreiro: MCP 40943, 3, 33.6−35.8 mm SL, arroio Guabiju, Guabiju, Rio Grande do Sul State.
Hisonotus charrua: LBP 4861, 1, 35.9 mm SL, arroio Guaviyú, Artigas, Uruguay; MHNG 2650.051, 1, 34.2 mm
SL, paratype, arroio Aspinillar, Uruguay.
Hisonotus chromodontus: LBP 7964, 25, 24.0−28.3 mm SL, 4 c&s, 24.9−28.9 mm SL, rio dos Patos, Nova Mutum,
Mato Grosso State; LBP 7974, 26, 17.7–24.8 mm SL, rio dos Patos, Nova Mutum, Mato Grosso State; LBP
12278, 2, 26.7−28.7 mm SL, 1 c&s, 26.7 mm SL, rio Sumidouro, Tangará da Serra, Mato Grosso State;
MZUSP 45355, 1, 25.9 mm SL, holotype, tributary of rio Preto, Diamantino, Mato Grosso State; MZUSP
70758, 7, 19.4−23.9 mm SL, paratype, riacho Loanda, Sinop, Mato Grosso State; NUP 10924, 24,
19.5−31.5 mm SL, rio Preto, Diamantino, Minas Gerais State.
Hisonotus depressicauda: MZUSP 5383, 1, 24.4 mm SL, paralectotype, Sorocaba, São Paulo State; LBP 17474, 5
c&s, 18.1−24.0 mm SL, rio Araquá, Botucatu, São Paulo State.
Hisonotus francirochai: LBP 13923, 22, 25.7−35.7 mm SL, córrego sem nome, Capetinga, Minas Gerais State;
MZUSP 3258, 1, 29.4 mm SL, lectotype, rio Grande, São Paulo State.
Hisonotus heterogaster: LBP 3335, 39, 20.8−30.1 mm SL, arroio sem nome, rio Grande, Rio Grande do Sul State;
MZUSP 104948, 3, 40.3−43.0 mm SL, paratypes, arroio Felício, Júlio de Castilho, Rio Grande do Sul State.
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Hisonotus iota: LBP 13072, 5, 32.3−33.0 mm SL, rio Chapecó, Coronel Freitas, Santa Catarina State.
Hisonotus laevior: LBP 3377, 1, 25.2 mm SL, arroio dos Corrientes, Pelotas, Rio Grande do Sul State; LBP 6037,
8, 33.4−47.0 mm SL, rio Maquiné, Osório, Rio Grande do Sul State; LBP 13187, 7, 19.4−45.8 mm SL,
córrego sem nome, Camaquá, Rio Grande do Sul State.
Hisonotus leucofrenatus: LBP 2085, 7, 38.3−50.6 mm SL, rio Sagrado, Morretes, Paraná State; LBP 6837, 36,
35.1−43.5 mm SL, rio Fau, Miracatu, São Paulo State.
Hisonotus leucophrys: LBP 13065, 6, 17.2−33.6 mm SL, rio Ariranhas, Xavantina, Santa Catarina State; LBP
13073, 1, 36.8 mm SL, rio Guarita, Palmitinho, Rio Grande do Sul State.
Hisonotus megaloplax: LBP 13108, 6, 36.4−37.8 mm SL, córrego sem nome, Saldanha Marinho, Rio Grande do
Sul State.
Hisonotus montanus: LBP 13051, 3, 26.4−27.2 mm SL, rio Goiabeiras, Vargem, Santa Catarina State; LBP 13055,
5, 24.8−31.9 mm SL, rio Canoas, Vargem, Santa Catarina State.
Hisonotus nigricauda: LBP579, 16, 34.1−40.1 mm SL, rio Guaíba, Eldorado do Sul, Rio Grande do Sul State.
Hisonotus notatus: LBP 3472, 20, 21.0−34.3 mm SL, 3 c&s, 25.8−26.5 mm SL, rio Aduelas, Macaé, Rio de
Janeiro State; LBP 10742, 25, 24.4−43.3 mm SL, rio Macabu, Conceição de Macabu, Rio de Janeiro State.
Hisonotus notopagos: MZUSP 104943, 4, 35.3−37.3 mm SL, arroio Boici, Pinheiro Machado, Rio Grande do Sul
State.
Hisonotus prata: MCP 40492, 18, 19.5−33.2 mm SL, rio da Prata, Nova Prata, Rio Grande do Sul State; LBP 9918,
14, 21.7−32.6 mm SL, Laguna dos Patos system, Nova Prata, Rio Grande do Sul State.
Hisonotus ringueleti: FMNH 108806, 2, 25.7−32.2 mm SL, rio Quaraí basin, Uruguay; LBP 13148, 1, 24.5 mm
SL, arroio Putiá, Uruguaiana, Rio Grande do Sul State.
Hisonotus vireo: MZUSP 104946, 4, 30.4−39.5 mm SL, rio dos Sinos, Caraá, Rio Grande do Sul State.
Microlepidogaster arachas: LBP 10882, 3, 22.8−35.3 mm SL, rio Paraná basin, Arachas, Minas Gerais State.
Microlepidogaster dimorpha: LBP 10683, 2, 28.8−35.6 mm SL, rio Uberaba, Uberaba, Minas Gerais State.
Otothyropsis alicula: LBP 14642, 1, 31.4 mm SL, rio Sapucaí Mirim, Campos do Jordão, São Paulo State; LBP
17583, 3, 29.7−33.7 mm SL, unknown river, Sapucaí Mirim, Minas Gerais State; NUP 10170, 10, 24.0−34.1
mm SL, rio Baú, Sapucaí Mirim, Minas Gerais State.
Otothyropsis biamnicus: MZUSP 86192, 3, 33.1−33.3 mm SL, unknown river, Telêmaco Borba, Paraná State;
MZUSP 86167, 22, 17.2−42.1 mm SL, Telêmaco Borba, Paraná State.
Otothyropsis marapoama: LBP 4698, 6, 23.9−36.3 mm SL, ribeirão Cubatão, Marapoama, São Paulo State;
MZUSP 87892, 1, 29.4 mm SL, ribeirão Cubatão, Marapoama, São Paulo State; MZUSP 87893, 4,
19.3−22.2 mm SL, córrego Cubatão, Catanduva, São Paulo State.
Otothyropsis polyodon: NUP 9946, 10, 24.2−35.6 mm SL, unknown river, Jateí, Mato Grosso do Sul State; NUP
9393, 4, 34.3−35.1 mm SL, córrego da Lagoa, Dourados, Mato Grosso do Sul State.
Parotocinclus maculicauda: LBP 2869, 15, 20.2−44.7 mm SL, rio Fau, Miracatu, São Paulo State.
Parotocinclus prata: LIRP 1136, 38, 19.8−41.9 mm SL, paratypes, ribeirão Quiricó, Presidente Olegário, Minas
Gerais State.
Parotocinclus robustus: LBP 8258, 29, 18.7−39.1 mm SL, córrego Cachoeira, Bocaiúva, Minas Gerais State.
Acknowledgements
The authors thank C. Araya, P.S. Cecarelli, F. Correa, R. Devidé, J.O. Junqueira, M.N. Mehanna, R. Rocha, V.F.
Sene, J.A. Senhorini for their help during collection expeditions. The authors also thank V.M. Azevedo-Santos for
providing cleared and stained specimens. This research was supported by Brazilian agencies FAPESP (Fundação
de Amparo à Pesquisa do Estado de São Paulo, proc. 2014/05051-5 to FFR, and proc. 2012/01622-2 to GSCS),
MCT/CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico) (Edital Universal, proc. N.
441347/2014-2 coord. FFR) and COLCIENCIAS (Departamento Administrativo de Ciencia, Tecnología e
Innovación, Colombia) doctoral fellowship grant 567/2012.
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