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Zookeys 498: 127—143 (2015)

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A new species of Hisonotus (Siluriformes, Loricariidae)
from rio Sao Francisco basin, Brazil

Fabio E Roxo!, Gabriel S. C. Silva', Claudio Oliveira!

| Universidade Estadual Paulista, Departamento de Morfologia, Laboratério de Biologia e Genética de Peixes,
Rubiao Junior s/n, 18618970 Botucatu, Sado Paulo State, Brazil

Corresponding author: Fabio F Roxo (roxoff@hotmail.com.br)

Academic editor: VV. Bogutskaya | Received 14 January 2015 | Accepted 16 April 2015 | Published 21 April 2015
Attp://zoobank. ore/154FEB27-179B-4DE0-B2B4-5BF7A61D44D3

Citation: Roxo FF, Silva GSC, Oliveira C (2015) A new species of Hisonotus (Siluriformes, Loricariidae) from rio Sao
Francisco basin, Brazil. ZooKeys 498: 127-143. doi: 10.3897/zookeys.498.6896

Abstract

A new species of Hisonotus is described from the rio Sao Francisco basin. The new species can be dis-
tinguished from congeners by having (1) a unique coloration pattern of caudal fin with one black spot
extending from its origin to the ventral lobe and two dark spots at the end of the lobe’s rays; (2) odontodes
forming longitudinally aligned rows on head and trunk; (3) a functional V-shaped spinelet; (4) a single
rostral plate at the tip of the snout; (5) by lacking contrasting dark geometric spots on the anterodorsal
region of the body; (6) a lower caudal-peduncle depth; and (7) lower counts of the lateral median plates
and (8) higher premaxillary and dentary teeth. The new species is the second described species of the genus
Hisonotus in the rio Sao Francisco basin. It was found inhabiting the marginal vegetation of the rio Sao

Francisco and three of its tributary, rio das Velhas, rio Paraopeba and rio Formoso.

Keywords
Cascudinhos, Fresh-Water, Minas Gerais State, Neotropical Fish, Otothyrinae

Introduction

Loricariidae is one of the largest and most diverse families of Neotropical freshwa-
ter fish with about 900 valid species (Eschmeyer and Fong 2014). The loricariids are
subdivided into seven subfamilies: Delturinae, Hypoptopomatinae, Hypostominae,
Lithogeninae, Loricariinae, Neoplecostominae, and Otothyrinae (Armbruster 2004;
Reis et al. 2006; Chiachio et al. 2008; Roxo et al. 2014a). Currently, Otothyrinae

Copyright Fabio F Roxo et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY
4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

128 Fabio E. Roxo et al. / ZooKeys 498: 127-143 (2015)

(sensu Chiachio et al. 2008 and Roxo et al. 2014a) is composed of about 100 valid
species classified in twelve genera (Eschmeyer 2014).

Within the Otothyrinae, the genus Hisonotus Eigenmann & Eigenmann, 1889 (type
species 1. notatus Eigenmann & Eigenmann, 1889) was resurrected by Schaefer (1998)
with the following combination of characters: snout plates in the anterior portion of
the nostril reduced or absent, the rostrum having enlarged odontodes, and the lateral
rostral margin composed of thickened plates. However, Britski and Garavello (2007)
considered the rostrum with enlarged odontodes as a character very polymorphic and
present in several other genera and species of Otothyrinae, as in species of Parotocinclus
Eigenmann & Eigenmann, 1889. Furthermore, Britski and Garavello (2007) suggested
that the other two characters are not satisfactory to define the genus Hisonotus.

Currently, Hisonotus contains 34 valid species, 19 of which were described in the
past decade (Eschmeyer 2014), representing an increase of 127% of the diversity of
this genus. Herein, we add a new species to the genus Hisonotus found during recent
collection expeditions to the rio Sao Francisco basin. This is the second species of the
genus described from this hydrographic system.

Material and methods

Measurements and counts were taken from the left side of the fish, and were made
from point to point to the nearest 0.1 mm with a digital caliper. Abbreviations used
in the text and the measurements followed Carvalho and Reis (2009). Specimens were
cleared and double stained (c&s) according to the method of ‘Taylor and Van Dyke
(1985). Vertebral counts also 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. Institutional acronyms
follow Fricke and Eschmeyer (2014). Specimens are deposited at the LBP, Laboratorio
de Biologia e Genética de Peixes, Universidade Estadual Paulista, Botucatu; MZUSP,
Museu de Zoologia, Universidade de Sao Paulo, Sao Paulo. Zoological nomenclature
follows the International Code of Zoological Nomenclature (International Commis-
sion on Zoological Nomenclature 1999).

Results

Hisonotus vespuccii sp. n.
http://zoobank.org/CD9657F5-3F02-4B32-BF89-24AF86020201
Figure 1, 3; Table 1

Holotype. MZUSP 115274 (female 32.6 mm SL) Brazil, Minas Gerais State, muni-
cipality of Pirapora, rio Sao Francisco, 17°21'00"S, 44°57'08"W, 11 November 2014,
LE Ochoa, FF Roxo, LH Roxo, GSC Silva.

A new species of Hisonotus (Siluriformes, Loricariidae) from rio Sado Francisco basin, Brazil 129

Figure |. Holotype of H. vespuccii, MZUSP 115274, female, 32.6 mm SL, from Minas Gerais State,
municipality of Pirapora, rio Sao Francisco, 17°21'00"S, 44°57'08" W.

Paratypes. All from Brazil, Minas Gerais State, rio Sao Francisco basin (249 spe-
cimens in total). LBP 8960 (4 females 32.3—35.8 mm SL, 3 males 27.0—28.2 mm
SL), municipality of Presidente Juscelino, rio das Velhas, 18°40'21"S, 44°11'33"W, 01
October 2009, C Oliveira, FF Roxo, GJC Silva, BF Melo. LBP 10421 (9 females 27.7—
30.3 mm SL, 9 males 23.6—26.5 mm SL, 5 c&s sex not determined 20.2—29.6 mm SL),
municipality of Pirapora, rio Sao Francisco, 17°21'00"S, 44°57'08"W, 15 July 2010, JA
Senhorini, M Mehanna. LBP 19491 (9 females 19.9—32.4 mm SL, 12 males 21.1—27.9
mm SL, 1 female c&s 32.6 mm SL, 1 male c&s 27.7 mm SL), collected with holotype.
LBP 19495 (1 female 32.6 mm SL, 1 male 28.7 mm SL), municipality of Presidente
Juscelino, rio das Velhas, 18°40'21"S, 44°11'33"W, 11 November 2014, LE Ochoa,
FF Roxo, LH Roxo, GSC Silva. MZUSP 39208 (5 sex not determined 23.8—30.0 mm
SL), rio Sao Francisco, 29 November 1987, Y Sato (UHE Formoso project). MZUSP
39280, (3 sex not determined 28.4—30.2 mm SL), rio Sao Francisco, 20 January 1988,
Y Sato (UHE Formoso project). MZUSP 39351 (33 sex not determined 16.5—34.4

130 Fabio E. Roxo et al. / ZooKeys 498: 127-143 (2015)

mm SL), rio Sao Francisco, 23 January 1988, Y Sato (UHE Formoso project). MZUSP
39446 (9 sex not determined 22.9-30.5 mm SL), rio Formoso, 08 February 1988, Y
Sato (UHE Formoso project). MZUSP 39482 (1 sex not determined 24.5 mm SL), rio
Formoso, 08 to 10 February 1988, Y Sato (UHE Formoso project). MZUSP 39511
(1 sex not determined 30.4 mm SL), cérrego Marambaia, rio Sao Francisco, 09 Fe-
bruary 1988, Y Sato (UHE Formoso project). MZUSP 51507 (2 sex not determined
25.6-31.3 mm SL), municipality of Fortuna de Minas, rio Paraopeba, 13 July 1995,
CBM Alves. MZUSP 57587 (110 sex not determined 20.8—33.6 mm SL), street be-
tween municipalities of Itacarambi and Manga, cérrego das Missdes, 15 July 1993,
RE Reis. MZUSP 57588 (7 sex not determined 20.2—24.9 mm SL), street between
municipalities of Manga and Montalvania, rio Japuré, rio Sao Francisco, 15 July 1993,
RE Reis. MZUSP 57590 (20 sex not determined 22.8-31.4 mm SL), municipality
of Januaria, rio Sao Francisco, 15°29'25"S, 44°21'50"W, 14 July 1993, JC Garavello.
MZUSP 57591 (3 sex not determined 23.1—26.5 mm SL), street between municipali-
ties of Manga and Montalvania, rio Calindd, rio Sao Francisco, 15 July 1993, RE Reis.

Diagnosis. Hisonotus vespuccii differs from the congeners by having a unique col-
oration pattern of caudal fin with one black spot extending from its origin to the
ventral lobe and two dark spots at the end of the lobe’s rays and the following combi-
nation of character states (none is unique): odontodes forming longitudinally aligned
rows (one odontode after the other, but not necessarily forming parallel series) on head
and trunk; a functional V-shaped spinelet; the presence of a single rostral plate at tip
of the snout; the lack of contrasting dark geometric spots on the anterodorsal region of
the body; a low caudal peduncle (depth 6—8% SL); few lateral median plates (21-23);
and numerous premaxillary and dentary teeth (13-21 and 11-21, respectively).

Description. Counts and measurements are presented in Table 1. Maximum body
size 35.7 mm SL. Dorsal profile of head, in lateral view, slightly convex from snout tip
to margin of posterior naris; strongly convex to posterior margin of parieto-supraoc-
cipital; and almost straight to dorsal-fin origin. Dorsal profile of trunk, in lateral view,
straight and descending from dorsal-fin origin to insertion of caudal-fin. Ventral pro-
file, in lateral view, straight from snout tip to anal-fin origin; concave and ascending to
caudal-fin insertion. Greatest body depth at dorsal-fin origin (14-18% SL). Greatest
body width at cleithral region (21-25% SL), progressively narrowing towards to both
snout and caudal fin. Cross-section of body between pectoral and pelvic fins dorsally
rounded and ventrally flat; cross-section of caudal peduncle ellipsoid, rounded laterally
and almost flat dorsally and ventrally.

Head rounded in dorsal view; snout round and slightly pointed. Dorsal and ven-
tral series of odontodes along anterior margin of snout completely covering its tip;
odontodes larger than remaining ones on head. Odontodes on head and trunk hyper-
trophied and arranged in longitudinal rows. Head without conspicuous crests. Some
specimens with a poor developed tuft of odontodes in posterior portion of parieto-su-
praoccipital. Eyes small (13-17% HL), dorsolaterally positioned. Iris operculum pre-
sent and developed. Premaxillary teeth 13-21; dentary teeth 11-21. Teeth bifid, major

A new species of Hisonotus (Siluriformes, Loricariidae) from rio Séo Francisco basin, Brazil 131

Figure 2. Coloration pattern of caudal fin of Hisonotus species. a H. vespuccii, LBP 19491, 26.2 mm
SL b H. acuen, LBP 16279, 26.1 mm SL ¢c A. armatus, LBP 14461, 38.2 mm SL d H. bocaiuva, LBP
17402, 24.8 mm SL e H. chromodontus, LBP 7964, 27.6 mm SL f H. depressicauda, LBP 1293, 26.1 mm
SL gH. francirochai, LBP 8356, 31.6 mm SL h ZH. heterogaster, LBP 14580, 38.7 mm SL i H. insperatus,
LBP 17432, 26.9 mm SLj H. iota, LBP 13072, 41.5 mm SLk ZH. leucofrenatus, LBP 2039, 40.7 mm SL
1 7. leucophrys, LBP 13071, 41.1 mm SL m 4. luteofrenatus, LBP 19534, 30.5 mm SLn H. megaloplax,
LBP 13108, 35.8 mm SL 0 H. montanus, LBP 13055, 31.8 mm SL p H. nigricauda, LBP 14652, 38.5
mm SL q H. notatus, LBP 18472, 32.0 mm SL r . oliveirai, LBP 13333, 24.0 mm SLs H. paresi, LBP
13351, 24.6 mm SLt H. piracanjuba, LBP 17256, 22.1 mm SL u Z. vireo, LBP 14452, 35.5 mm SL.

132 Fabio E. Roxo et al. / ZooKeys 498: 127-143 (2015)

Tablel. Morphometrics and meristic data for Hisonotus vespuccii, holotype and paratypes measured are
from LBP fish collection.

Character Holotype Males, n = 21 Females, n = 24 al fi eee

SD | Mean | SD
SL 32.6 28.3 | 3.0

Percents of SL <a Sse.

Head length 33.1 35.8 | 1.1

Predorsal length 44.5 44.2-47.3 | 45.8 | 0.8 | 44.1-48.6| 46.1 | 1.2] 46.0 | 1.0

Dorsal-fin spine length 22.6 24.6 | 1.3 | 19.5-25.4 23.8 1.4

Anal-fin unbranched ray length 15.5 16.0-19.2 14.9-18.1} 16.1 |0.7] 16.7 | 1.0
Pectoral-fin spine length 23.9 | 23.9-29.1 2 lLS2E/y| 252A ek P25" |S
Pelvic-fin unbranched ray length} 16.9 17.3-21.5 15.1-18.5 | 16.8 | 0.8} 18.0 | 1.5

Cleithral width pies 23.2 | 0.5 | 21.2-24.9 | 23.0 | 0.8| 23.1 | 0.6
Thoracic length 18.0 | 15.2-183] 16.5 | 0.7] 14.9-18.4| 16.7 |0.9| 16.6 | 0.8
Abdominal length 22.6 | 20.7-23.1| 21.6 | 0.5 | 20.1-24.6| 22.5 [1.1] 22.1 | 1.0

Body depth at dorsal-fin origin | 14.7 15.7 | 0.9 | 14.3-17.5| 15.9 [0.8] 15.8 | 0.9

Caudal-peduncle length 39.0 | 36.7-40.3 | 38.6 34.0-39.4 | 37.3 | 1.2| 37.9 | 1.2
Caudal-peduncle depth 6.2 3| 6.2-7.9 | 7.0 |04} 7.2 | 0.4
Nares opening 12.5 TAL Oe? |e ASO | #23
Percents of HL oe oad
Snout length 48.8 45.5-50.3 | 48.9 | 1.1] 48.8 | 1.2
Orbital diameter 16.1 13.4-16.9 | 14.9 |0.9} 15.2 | 0.9
Interorbital width 36.1 B95. 1E=387 371 34.1-39.0 | 36.2 | 1.3] 36.6 | 1.2

Head depth 40.5 | 39.0-45.0 42.3 | 17
Suborbital depth 172 16.6 | 0.9 | 14.9-20.3 16.8 | 1.3
Mandibular ramus D7. 81-105 | 9.3 | 0.6) 8.3-15.6 | 9.6 | 1.4]. 9.5 LA

Meristics Se SS ESS ae)

Left lateral scutes 22 21-23 22 22 -
Left premaxillary teeth 19 15-21 19 17 -
Left dentary teeth 19 11-19 16 16 -

(medial) cusp large and rounded, minor (lateral) cusp minute and pointed. Accessory
patch of teeth absent on dentary and premaxilla. Oral disk oval, covered with papillae
uniformly distributed on base of dentary and premaxilla and slightly decreasing in size
distally. Lower lip larger than upper lip; its border fringed. Maxillary barbel present
and joined to lower lip. Presence of conspicuous V-shaped buccal papilla located im-
mediately anterior to buccal valve. Tip of snout with large rostral plate.

Dorsal fin II,7; its origin slightly posterior to pelvic-fin origin. Tip of adpressed
dorsal fin surpassing vertically through end of anal-fin origin. Dorsal-fin spinelet short
and V-shaped; dorsal-fin lock functional. Pectoral fin 1,6; its tip reaching middle of
pelvic-fin length when depressed. Pectoral-axillary slit present between pectoral-fin
insertion and lateral process of cleithrum. Pectoral spine supporting sharp odontodes
on dorsal and ventral surfaces (well developed posteriorly). Pelvic fin I,5; its tip reach-

A new species of Hisonotus (Siluriformes, Loricariidae) from rio Sao Francisco basin, Brazil 133

Figure 3. Live specimen of H. vespuccii, LBP 19491, paratype, 32.4 mm SL, from Minas Gerais State,
municipality of Pirapora, rio Sao Francisco.

Ss gp ; \ pee wY NS y
Nar” ~ » - a fie ji So a6

Figure 4. (a) (b) Type locality of H. vespuccii at rio Sao Francisco, municipality of Pirapora, Minas Ger-
ais State, 17°21'00"S, 44°57'08" W. (c) (d) Paratypes locality of H. vespuccii at rio das Velhas, municipality
of Presidente Juscelino, Minas Gerais State, 18°40'21"S, 44°11'33"W. Photo: LH Roxo.

ing anal-fin origin when depressed in males and far from reaching anal-fin origin in
females. Pelvic-fin unbranched ray with dermal flap along its dorsal surface in males.
Pectoral spine supporting sharp odontodes on ventral surface turned mesially.

134 Fabio E. Roxo et al. / ZooKeys 498: 127-143 (2015)

Figure 5. Map of the distribution of H. vespuccii. Red Star = type locality, at rio Sao Francisco, munici-

pality of Pirapora, Minas Gerais State. Red Circles = paratypes localities of H. vespuccii at rio das Velhas,

rio Paraopeba and rio Formoso, Minas Gerais State.

Anal fin i,5; its tip reaching seventh or eighth plate from its origin. Caudal-fin i,7—
7,1; distal margin forked. Adipose fin absent. Total vertebrae 27 (in 7 c&s specimens).
Body almost entirely covered by bone plates, except on ventral portion of head, around
pectoral- and pelvic-fin origins, on dorsal-fin base and area around anus. Abdomen
partially covered by bony plates randomly distributed and surrounded by naked areas
(in some specimens abdomen is completely covered by bony plates). Laterally, body
completely covered by plates; mid-dorsal and mid-ventral plate series well developed
reaching vertical through half of caudal peduncle; median plate series continuous in
median portion of body. Coracoid and cleithrum completely exposed, covered with
odontodes. Arrector fossae partially enclosed by ventral lamina of coracoids.

Color in alcohol. Ground color of dorsal surface of head and body dark gray to
lighter brown (juveniles lighter than adults). Ventral surface light brown to yellow
in juveniles. All body and fins covered by scattered chromatophores, more visible on
ventral portions and around fins insertions (Fig. 1). Caudal-fin hyaline, except for one
black spot at its origin extending to ventral lobe and two dark spots at end of rays (Fig.
2a). In some specimens, caudal-fin with chromatophores irregular distributed and
sometimes badly forming two dark strips (more visible in juveniles). Neither variation
nor variability of caudal-fin coloration patter found in samples we examined (holotype
and 249 specimens widely distributed in rio Sao Francisco basin) with specific empha-
sis to variability between populations and variation depending on feeding.

A new species of Hisonotus (Siluriformes, Loricariidae) from rio Sao Francisco basin, Brazil 135

Color in life. Similar to pattern described for alcohol individuals, but with ground
color light green (Fig. 3).

Sexual dimorphism. Adult males distinguished from females by five characters:
(1) presence of a papilla at urogenital opening in males (vs. papilla absent in females);
(2) pelvic-fin extending beyond anal-fin origin in males, mean 19% SL (ws. pelvic fin
far from reaching anal-fin origin in females, mean 17% SL); (3) unbranched pelvic-fin
ray supporting a dermal flap (flap slightly wider in basal portion and progressively nar-
rowing distally) on proximal dorsal surface in males (vs. dermal flap absent in females);
(4) nares opening wider in males (13-18% HL) than females (10-13% HL); (5) body
size smaller in males (mean 26 mm SL) and larger in females (mean 30 mm SL). See
Table 1 for values of morphometric characters between males and females.

Habitat and distribution. Hisonotus vespuccii was found associated with marginal
vegetation (Fig. 4) in the rio Sao Francisco and in three of its tributaries, rio das Vel-
has, rio Paraopeba and rio Formoso (Fig. 5). The new species seems to be abundant
through all rio Sao Francisco basin.

Etymology. The specific name “vespuccii” comes from Italian and is in reference
to Américo Vesptcio (Amerigo Vespucci in Italian), navigator and explorer to whom
is attributed the discovery of the rio Sao Francisco in 1501.

Comparative remarks and discussion

The new species H. vespuccii has one character proposed by Schaefer (1998) to diag-
nose the genus Hisonotus: the rostrum with enlarged odontodes. Moreover, the new
species also shares three characters with many species of Hisonotus: a single rostral plate
on the tip of the snout, an arrector fossae partially enclosed by a ventral lamina of the
coracoid, a character also used by Schaefer (1998) as synapomorphy of all Otothyrini
except the New Taxon 3, and a functional V-shaped spinelet. ‘This last character was
firstly proposed by Carvalho and Datovo (2012) with pers. comm. of Roberto E. Reis,
and posteriorly was reported by Silva et al. (2014) as a possible synapomorphy that
may help delimit a new genus within Hisonotus. However, to better understand the
relationship of the new species with other species assigned to Hisonotus a phylogenetic
analysis is still necessary.

We used seven characters to distinguish the new species H. vespuccii from conge-
ners. The first character was the caudal fin with one black spot extending from its ori-
gin to the ventral lobe and two dark spots at the end of the lobe’s rays, a pattern unique
among Hisonotus species (see Fig. 2b—u for caudal-fin coloration pattern of some spe-
cies of Hisonotus). Britski and Garavello (2007) discussing about the coloration pattern
of the teeth in H. chromodontus Britski & Garavello, 2007 suggested that this aspect
of the organism could be a result of physiological features changing according to the
individual’s foraging success and physiological efficiency, as well as according to the
characteristics of the water where the species lives. However, the pattern of caudal-fin
coloration seems to be conserved among species of Hisonotus (Fig. 2), with a pattern

136 Fabio E Roxo et al. / ZooKeys 498: 127-143 (2015)

varying more drastically in H. acuen Silva, Roxo & Oliveira, 2014, a species widely
distributed through headwaters of the rio Xingu basin (see Fig. 5 in Silva et al. 2014).
In H. vespuccii, the pattern of the caudal-fin with one black spot extending from its
origin to the ventral lobe and two dark spots at the end of the lobe’s rays is present in
the holotype and the 249 paratypes analyzed.

Moreover, the new species H. vespuccii was distinguished from congeners by
present a combination of the following characters: (2) odontodes forming longitudi-
nally aligned rows (one odontode after the other, but not necessarily forming parallel
series) on head and trunk, a character shared with H. insperatus, H. luteofrenatus,
H. oliveirai, and H. paresi; (3) a functional V-shaped spinelet as reported previously
(see Fig. 2 in Silva et al. 2014 to description of this character state) shared with H.
acuen, H. bockmanni, H. chromodontus, H. insperatus, H. luteofrenatus, H. oliveirai,
H. paresi, and H. piracanjuba; (4) the presence of a single rostral plate at tip of the
snout, a character also present in H. insperatus, H. luteofrenatus, H. oliveirai, H.
paresi, and H. piracanjuba; (5) the lacking of contrasting dark geometric spots on
the anterodorsal region of the body, a character used to distinguish the new spe-
cies of H. bockmanni and H. paresi; (6) a lower caudal-peduncle depth (6.2—8.0%
of SL) used to distinguish the new species of H. acuen (8.6-11.1% of SL); and (7)
lower counts of the lateral median plates (21—23) and higher counts of premaxillary
(13-21) and dentary teeth (11—21) used to distinguish H. vespuccii of H. insperatus
(25-26; 10-12; 8-11, respectively).

Members of Loricariidae are known to have intense sexual dimorphisms (Py-Dan-
iel and Fernandes 2005) as we can observe in species of the genera Ancistrus Kner, 1854
(Sabaj et al. 1999), Neoplecostomus Eigenmann & Eigenmann, 1888 (Zawadzki et al.
2008; Roxo et al. 2012; Andrade and Langeani 2014), Pareiorhaphis Miranda Ribeiro,
1918a (Pereira et al. 2007), Hisonotus (Martins and Langeani 2012), Hypostomus La-
cepéde, 1803 and Chaetostoma Tschudi, 1846 (Nomura and Mueller 1980; Lopez and
Roman-Valencia 1996), Farlowella Eigenmann & Eigenmann, 1889 (Retzer and Page
1996) and many other. In H. vespuccii, we observed five sexual dimorphic characters:
the presence of a papilla at the urogenital opening, a pelvic-fin that extends beyond the
anal-fin origin, the unbranched pelvic-fin ray supporting a dermal flap on their proxi-
mal dorsal surface, the nares opening wider and a body size that seems to be smaller
in males than in females. The first three characters are very common among species
of Hisonotus, however differences in size of nares were only previously reported in the
original description of H. piracanjuba (Martins and Langeani 2012) and differences
in body size in 1. ringueleti (Aquino et al. 2001). Carvalho and Reis (2009) reported
that the presence of the dermal flap on proximal dorsal surface of pelvic-fin on males
is a plesiomorphic character shared among most members of Otothyrinae and that the
derived condition evolved several times within this subfamily at the genera Schizolecis
Britski & Garavello, 1984, Epactionotus Reis & Schaefer, 1998, and within Hypopto-
pomatinae in Acestridium Haseman, 1911, Oxyropsis Eigenmann & Eigenmann, 1889

and Hypoptopoma Ginther, 1868.

A new species of Hisonotus (Siluriformes, Loricariidae) from rio Séo Francisco basin, Brazil 137

Comparative material

All from Brazil, except when stated otherwise.

Hisonotus acuen Silva, Roxo & Oliveira, 2014: MZUSP 115350, 1, 25.9 mm SL, ho-
lotype, tributary of rio Toguro, Queréncia, Mato Grosso State; LBP 15755, 16,
19.5-26.0 mm SL, paratypes, tributary of rio Suia-Missu, ribeirao Cascalheira, Mato
Grosso State; LBP 16274, 27, 20.2-29.1 mm SL, 2 c&s, 23.6-24.2 mm SL, para-
types, tributary of rio Culuene, Gaticha 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,
Queréncia, Mato Grosso State; LBP 16278, 12, 18.8-25.1 mm SL, 2 c&s, 26.8-27.1
mm SL, paratypes, cérrego Xavante, Primavera do Leste, Mato Grosso State.

Hisonotus aky (Azpelicueta, Casciotta, Almirén & Koerber, 2004): MHNG 2643.039,
2, 33.1-34.2 mm SL, paratypes, arroio Fortaleza, Argentina.

Hisonotus armatus Carvalho, Lehmann, Pereira & Reis, 2008: MZUSP 93884, 5, 37.6—
44.4 mm SL, paratypes, arroio Arambaré, Pedro Osdrio, Rio Grande do Sul State.

Hisonotus bocaiuva Roxo, Silva, Oliveira & Zawadzki, 2013: MZUSP 112204, 1, 24.2
mm SL, holotype, cérrego Cachoeira, Bocaitiva, Minas Gerais State; LBP 9817, 9, 3
c&Xs, 18.3-23.2 mm SL, paratypes, cérrego Cachoeira, Bocaitiva, Minas Gerais State.

Hisonotus brunneus Carvalho & Reis, 2011: MZUSP 104947, 4, 37.2-41.3 mm SL,
paratypes, rio Passo Novo, Cruz Alta, Rio Grande do Sul State.

Hisonotus carreiro Carvalho & Reis, 2011: MCP 40943, 3, 33.6-35.8 mm SL, arroio
Guabiju, Guabiju, Rio Grande do Sul State.

Hisonotus charrua Almiron, Azpelicueta, Casciotta & Litz, 2006: LBP 4861, 1, 35.9
mm SL, arroio Guaviyt, Artigas, Uruguay; MHNG 2650.051, 1, 34.2 mm SL,
paratype, arroio Aspinillar, Uruguay.

Hisonotus chromodontus Britski & Garavello, 2007: 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, Tangara
da Serra, Mato Grosso State; MZUSP 45355, 1, 25.9 mm SL, holotype, tribu-
tary 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 (Miranda Ribeiro, 1918b): MZUSP 5383, 1, 24.4 mm SL,
paralectotype, Sorocaba, Sao Paulo State; LBP 17474, 5 c&s, 18.1-24.0 mm SL,
rio Araqua, Botucatu, Sao Paulo State.

Hisonotus francirochai (Ihering, 1928): LBP 13923, 22, 25.7-35.7 mm SL, cérrego
sem nome, Capetinga, Minas Gerais State; MZUSP 3258, 1, 29.4 mm SL, lecto-
type, rio Grande, Sao Paulo State.

Hisonotus heterogaster Carvalho & Reis, 2011: 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 Felicio, Julio de Castilho, Rio Grande do Sul State.

138 Fabio E Roxo et al. / ZooKeys 498: 127-143 (2015)

Hisonotus insperatus Britski & Garavello, 2003: LBP 4945, 5, 27.3-28.5 mm SL, 2
c&s, 28.2-29.9 mm SL, Botucatu, S40 Paulo State; LBP 6770, 5, 25.1-28.2 mm
SL, 3 c&s, 20.0-27.0 mm SL, ribeirao Cubatao, Marapoama, Sao Paulo State;
LBP 13336, 1 c&s, 26.0 mm SL, rio Capivara, Botucatu, Sao Paulo State; LBP
13337, 2 c&s, 27.4-28.6 mm SL, rio Araqua, Botucatu, Sao Paulo State; MZUSP
22826, 1, 25.4 mm SL, paratype, cérrego Agua Tirada, Trés Lagoas, Minas Gerais
State; MZUSP 24832, 1, 23.8 mm SL, paratype, rio Corumbatai, Corumbatai,
Sao Paulo State; MZUSP 78957, 1, 29.6 mm SL, holotype, rio Capivara, Botu-
catu, Sao Paulo State; MZUSP 78960, 31, 12.6-26.0 mm SL, paratypes, 5 c&s,
22.7-24.7 mm SL, rio Pardo, Botucatu, S40 Paulo State; MZUSP 78965, 10,
15.6-28.6 mm SL, paratypes, rio Araqua, Botucatu, Sao Paulo State; MZUSP
78968, 5, 24.1-27.3 mm SL, paratypes, cérrego da Figueira, Lins, Sao Paulo State.

Hisonotus iota Carvalho & Reis, 2009: LBP 13072, 5, 32.3-33.0 mm SL, rio Chapecé,
Coronel Freitas, Santa Catarina State.

Hisonotus laevior Cope, 1894: LBP 3377, 1, 25.2 mm SL, arroio dos Corrientes, Pelo-
tas, Rio Grande do Sul State; LBP 6037, 8, 33.4-47.0 mm SL, rio Maquiné,
Osério, Rio Grande do Sul State; LBP 13187, 7, 19.4-45.8 mm SL, cérrego sem
nome, Camaqua, Rio Grande do Sul State.

Hisonotus leucofrenatus (Miranda Ribeiro, 1908): LBP 2085, 7, 38.3-50.6 mm SL,
rio Sagrado, Morretes, Parana State; LBP 6837, 36, 35.1-43.5 mm SL, rio Fau,
Miracatu, Sao Paulo State.

Hisonotus leucophrys Carvalho & Reis, 2009: LBP 13065, 6, 17.2-33.6 mm SL, rio
Ariranhas, Xavantina, Santa Catarina State; LBP 13073, 1, 36.8 mm SL, rio Guar-
ita, Palmitinho, Rio Grande do Sul State.

Hisonotus luteofrenatus Britski & Garavello, 2007: MZUSP 62593, 1, 28.6 mm SL, hol-
otype, cérrego Loanda, Claudia, Mato Grosso State; MZUSP 62594, 8, 22.4-30.5
mm SL, paratypes, riacho Selma, Sinop, Mato Grosso State; MZUSP 87144, 8,
16.8-27.9 mm SL, paratypes, cérrego Loanda, Claudia, Mato Grosso State.

Hisonotus megaloplax Carvalho & Reis, 2009: LBP 13108, 6, 36.4-37.8 mm SL, cér-
rego sem nome, Saldanha Marinho, Rio Grande do Sul State.

Hisonotus montanus Carvalho & Reis, 2009: 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 (Boulenger, 1891): LBP579, 16, 34.1-40.1 mm SL, rio Guaiba,
Eldorado do Sul, Rio Grande do Sul State.

Hisonotus notatus Eigenmann & Eigenmann, 1889: 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, Conceicao de Macabu, Rio de Janeiro State.

Hisonotus notopagos Carvalho & Reis, 2011: MZUSP 104943, 4, 35.3-37.3 mm SL,
arroio Boici, Pinheiro Machado, Rio Grande do Sul State.

Hisonotus oliveirai Roxo, Zawadzki & Troy, 2014b: MZUSP 115061, 1, 26.4 mm
SL, holotype, ribeirao Cambira, tributary of rio Ivai, Cambira, Parana State;
LBP 13332, 1 23.2 mm SL, 1 c&s, 23.7 mm SL, paratype, rio Mourao, Campo

A new species of Hisonotus (Siluriformes, Loricariidae) from rio Sao Francisco basin, Brazil 139

Mourado, Parana State; LBP 17578, 5, 25.4-30.4 mm SL, paratypes, rio Mourao,
between Engenheiro Beltrao and Quinta do Sol, Parana State; NUP 3578, 15,
24.7-28.1 mm SL, 2 c&s, 25.5-27.6 mm SL, paratypes, ribeirao Salto Grande,
Maria Helena, Parana State.

Hisonotus paresi Roxo, Zawadzki & Troy, 2014b: MZUSP 115062, 1, 26.2 mm SL,
holotype, riacho Aguas Claras, Santo Afonso, Mato Grosso State; LBP 13351, 9,
14.7-24.3 mm SL, paratype, riacho Aguas Claras, Santo Afonso, Mato Grosso
State; LBP 13352, 1, 23.7 mm SL, paratype, riacho Aguas Claras, Santo Afon-
so, Mato Grosso State; NUP 10928, 2, 23.2-24.2 mm SL, paratype, 2 c&s,
23.6-24.2 mm SL, riacho Aguas Claras, Santo Afonso, Mato Grosso State; NUP
10976, 3, 16.7-20.5 mm SL, paratype, riacho Sao Jorge, Santo Afonso, Mato
Grosso State.

Hisonotus piracanjuba Martins & Langeani, 2012: LBP 17256, 9, 17.2-26.3 mm SL, 1
c&s, 27.1 mm SL, corrego sem nome, Morrinhos, Goids State; NUP 5059, 1, 24.7
mm SL, cérrego Posse, Anapolis, Goias State; MZUSP 110491, 3, 17.5-24.4 mm
SL, paratypes, rio Quente, Marcelania, Goids State; NUP 10979, 3, 21.4-21.8
mm SL, ribeirao Bocaina, Piracanjuba, Goias State.

Hisonotus prata Carvalho & Reis, 2011: 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 Aquino, Schaefer & Miquelarena, 2001: FMNH 108806, 2,
25.7-32.2 mm SL, rio Quarai basin, Uruguay; LBP 13148, 1, 24.5 mm SL, arroio
Putid, Uruguaiana, Rio Grande do Sul State.

Hisonotus vireo Carvalho & Reis, 2011: MZUSP 104946, 4, 30.4-39.5 mm SL, rio
dos Sinos, Caraa, Rio Grande do Sul State.

Hisonotus sp. 1 n.: LBP 8276, 1 c&s, 25.6 mm SL, rio Verde Grande, Jaiba, Minas
Gerais State.

Hisonotus sp. 2 n.: MZUSP 95687, 8, 19.8-21.5 mm SL, ribeirao da Anta, Gaticha do
Norte, Mata Grosso State.

Hisonotus sp. 3 n.: MZUSP 84157, 6, 20.1-23.6 mm SL, rio Manuel Alves, Porto
Alegre do Tocantins, Tocantins State.

Microlepidogaster arachas Martins, Calegari & Langeani, 2013: LBP 10882, 3,
22.8-35.3 mm SL, rio Parana basin, Arachas, Minas Gerais State.

Microlepidogaster dimorpha Martins & Langeani, 2011: LBP 10683, 2, 28.8-35.6 mm
SL, rio Uberaba, Uberaba, Minas Gerais State.

Otothyropsis marapoama Ribeiro, Carvalho & Melo, 2005: LBP 4698, 6, 23.9-36.3
mm SL, ribeirao Cubatao, Marapoama, Sao Paulo State.

Parotocinclus maculicauda (Steindachner, 1877): LBP 2869, 15, 20.2-44.7 mm SL, rio
Fau, Miracatu, Sao Paulo State.

Parotocinclus prata Ribeiro, Melo & Pereira, 2002: LIRP 1136, 38, 19.8-41.9 mm SL,
paratypes, ribeirao Quiricd, Presidente Olegario, Minas Gerais State.

Parotocinclus robustus Lehmann & Reis, 2012: LBP 8258, 29, 18.7-39.1 mm SL, cérrego
Cachoeira, Bocaitiva, Minas Gerais State.

140 Fabio E. Roxo et al. / ZooKeys 498: 127-143 (2015)

Acknowledgements

The authors wish to thank Bruno F Melo, Carlos BM Alves, Guilherme JC Silva, José A
Senhorini, Julio C Garavello, Luiz H Roxo, Luz E Ochoa, Mahmoud Mehanna, Rob-
erto E Reis and Yoshimi Sato for their help during the collection expeditions; Marga-
rete Penteado for English revision. Research was supported by Brazilian agencies FAPESP
(Fundacao de Amparo a Pesquisa do Estado de Sao Paulo, proc. 2014/05051-5 to FFR,
and proc. 2012/01622-2 to GSCS), MCT/CNPq (Conselho Nacional de Desenvolvi-
mento Cientifico e Tecnolégico) (Edital Universal, proc. N. 441347/2014-2 coord. FFR).

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