A New Species of Hypostomus (Siluriformes: Loricariidae) from the Upper Rio
Paraguay Basin, Brazil
Fernanda O. Martins
, Manoela M. F. Marinho
, Francisco Langeani
and Jane P. Serra
A new small Loricariidae, Hypostomus careopinnatus, is described from the Rio Taquari drainage, upper Rio Paraguay
basin, Mato Grosso, Brazil. The new species can be easily distinguished from all congeners, except Hypostomus levis, by
the absence of adipose fin. Hypostomus careopinnatus is distinguished from H. levis mainly by the presence of slender bifid
teeth, with mesial cusp large and rounded, and lateral cusp small and pointed (vs. spoon-shaped teeth). The new species
described herein completely lacks the adipose fin and also lacks the median pre-adipose plates in almost all specimens
examined. The absence of adipose fin is probably an independent acquisition for Hypostomus careopinnatus and
Um pequeno novo Loricariidae, Hypostomus careopinnatus,e´ descrito das drenagens do Rio Taquari, bacia do alto Rio
Paraguai, Mato Grosso, Brasil. A nova espe´cie distingue-se facilmente de todas as demais congeˆneres, exceto de
Hypostomus levis, pela auseˆncia da nadadeira adiposa. Hypostomus careopinnatus distingue-se de H. levis pela presenc¸ a de
dentes delgados e bicuspidados, com a cu´spide mesial maior e arredondada, e a cu´ spide lateral menor e afilada (vs.
dentes em forma de colher). A nova espe´cie aqui descrita na˜o possui nadadeira adiposa, e tambe´ m perdeu as placas
medianas pre´-adiposa, na maioria dos exemplares examinados. A auseˆncia da nadadeira adiposa e´ provavelmente uma
aquisic¸a˜o independente para Hypostomus careopinnatus eHypostomus levis.
THE genus Hypostomus La Ce´pe`de is the most species-
rich genus within the family Loricariidae, comprising
140 valid species according to Armbruster (2004),
Armbruster et al. (2007), and recent descriptions (Zawadzki
et al., 2008a, 2008b, 2010; Carvalho et al., 2010), occurring
widely throughout the neotropics. The high intraspecific
variation in its morphology results in many nominal species
with unclear status in Hypostomus (Weber, 2003; Birindelli
et al., 2007; Jerep et al., 2007). Also, the limits of the genus
are yet to be unequivocally defined. Armbruster (2004)
recovered a monophyletic Hypostomus, though not support-
ed by exclusive features, and recognized Aphanotorulus,
forma, and Watawata as junior synonyms of Hypostomus.
However, based on molecular data, Montoya-Burgos et al.
(2002) considered Aphanotorulus,Isorineloricaria, and Squali-
forma as valid genera.
A new Hypostomus described herein was recently collected
in the headwaters of the Rio Taquari, Rio Paraguay basin.
The new finding corroborates the statement of Lima and
Britto (2001) that the ichthyofauna of this region, as the
headwaters of several major river basin in the Neotropics, is
still poorly known.
MATERIALS AND METHODS
Measurements were taken according to Boeseman (1968)
and Armbruster (2003), with the inclusion of prepectoral
length, taken from the snout tip to the pectoral-fin origin;
premaxillary–ramus width, taken transversely from outer
edge to inner edge; and preanal length, taken from the
snout tip to the anal-fin origin. All measurements were
taken point to point with digital calipers to the nearest
0.1 mm. Body measurements are given as percents of the
standard length (SL), except when noted; subunits of the
head are given as percents of head length (HL). Counts
followed Armbruster (2003) and Zawadzki et al. (2008b),
with the addition of ventral plates along the anal-fin base.
Vertebrae counts included five from the Weberian appara-
tus, and the compound caudal centrum was counted as a
single element. Body plate nomenclature was based on
Schaefer (1997) with modifications of Oyakawa et al.
(2005). In the description, the mode of each count is given
in parentheses, after the respective count. Specimens were
cleared and stained (CS) according to Taylor and Van Dyke
(1985). Institutional abbreviations are as listed at http://
www.asih.org/node/204, with the addition of the DZSJRP
fish collection, Departamento de Zoologia e Botaˆnica,
Universidade Estadual Paulista, Sa˜o Jose´ do Rio Preto,
Hypostomus careopinnatus, new species
Figure 1, Table 1
Holotype.—DZSJRP 12231, 57.9 mm SL, Brazil, Mato Grosso
State, Alto Araguaia, tributary of Rio Ariranha, Rio Taquari
drainage, upper Rio Paraguay basin, 17u189370S,
53u329220W, F. Langeani, J. P. Serra, M. M. F. Marinho, A.
Manzale, 11 August 2009.
Paratypes.—Brazil, Mato Grosso State, Alto Araguaia:
DZSJRP 12447, 70, 21.2–57.0 mm SL; MCP 45987, 5, 28.7–
52.0 mm SL; MNRJ 38309, 5, 30.3–50.6 mm SL; collected
with the holotype. ANSP 190960, 5, 29.5–51.7 mm SL; AUM
51645, 5, 25.9–55.9 mm SL; CPUFMT 690, 5, 33.9–49.0 mm
UNESP—Universidade Estadual Paulista, Laborato´rio de Ictiologia, Departamento de Zoologia e Botaˆnica, Rua Cristo´va˜ o Colombo, 2265,
15054-000 Sa˜o Jose´ do Rio Preto, SP, Brazil; E-mail: (FOM) firstname.lastname@example.org; and (FL) email@example.com. Send
reprint requests to FOM.
Instituto Federal de Educac¸a˜o, Cieˆncia e Tecnologia Sul de Minas Gerais, estrada de Muzambinho, km 35, Caixa Postal 02, 37890-000
Muzambinho, MG, Brazil; E-mail: firstname.lastname@example.org.
Submitted: 25 January 2011. Accepted: 15 February 2012. Associate Editor: R. E. Reis.
F2012 by the American Society of Ichthyologists and Herpetologists DOI: 10.1643/CI-11-011
Museu de Zoologia da Universidade de Sa˜o Paulo, Caixa Postal 42494, 04299-970 Sa˜o Paulo, SP, Brazil; E-mail: email@example.com.
Copeia 2012, No. 3, 494–500
SL; DZSJRP 12611, 56, 16.9–59.1 mm SL; INPA 35151, 5,
32.2–56.3 mm SL; NUP 11257, 5, 30.2–53.8 mm SL; UFRGS
14043, 5, 34.4–50.8 mm SL; same locality as holotype, F.
Langeani, J. P. Serra, M. M. F. Marinho, F. O. Martins, 3
June 2010. MZUSP 41483, 10, 19.5–47.0 mm SL, Co´rrego do
Mato, Fa zenda Co´ rrego do Mato, Rio Taquari drainage,
upper Rio Paraguay basin, L. P. S. Portugal, F. Langeani, 9
Fig. 1. Hypostomus careopinnatus, holotype, DZSJRP 12231, 57.9 mm SL, Rio Taquari drainage, upper Rio Paraguay basin, Mato Grosso State, Brazil.
Martins et al.—New Hypostomus from Paraguay River basin 495
Diagnosis.—Hypostomus careopinnatus can be readily distin-
guished from all congeners, except Hypostomus levis, by the
complete absence of adipose fin (vs. presence). Hypostomus
careopinnatus is distinguished from H. levis mainly by the
slender bifid teeth, with mesial cusp large and rounded, and
lateral cusp small and pointed (vs. spoon-shaped teeth), by
having 23–42 premaxillary and 25–42 dentary teeth (vs. 11–
17 for both premaxillary and dentary); and body covered
with black spots (vs. spots absent). Additionally, the new
species differs from all species of Hypostomus which occur in
the La Plata basin (except H. ancistroides,H. angipinnatus,H.
aspilogaster,H. borellii,H. boulengeri,H. brevis,H. commersoni,
Table 1. Morphometric and Meristic Data of the Holotype and 29 Paratypes of Hypostomus careopinnatus. Means are presented for measurements,
and modes for counts. The range includes the holotype. SD =standard deviation.
Holotype nRange Mean SD
Standard length (mm) 57.9 30 41.3–57.9 — —
Percents of standard length
Predorsal length 43.2 30 40.2–46.0 43.4 1.3
Preanal length 62.2 30 60.9–65.6 63.2 1.3
Head length 32.5 30 31.7–37.5 34.0 1.3
Thoracic length 25.4 30 19.8–25.4 22.3 1.3
Abdominal length 19.0 30 18.0–22.8 20.4 1.1
Dorsal-fin unbranched ray length 23.7 30 21.2–26.8 23.6 1.3
Dorsal-fin base length 20.0 30 19.0–24.5 20.8 1.2
Pectoral-fin unbranched ray length 26.3 30 23.9–27.9 25.8 1.1
Pelvic-fin unbranched ray length 23.3 30 21.1–25.6 23.5 1.0
Anal fin length 14.7 29 12.7–16.4 14.0 0.8
Cleithral width 29.9 30 26.6–30.3 28.3 1.0
Caudal peduncle length 32.6 30 30.5–36.9 33.3 1.5
Caudal peduncle depth 8.1 30 7.4–8.6 7.9 0.3
Prepectoral length 26.4 29 25.1–29.5 27.0 1.1
Dorsal–pectoral length 28.7 29 26.6–29.6 28.3 0.9
Dorsal–pelvic length 19.5 29 15.5–22.8 19.2 1.6
Dorsal–anal length 32.3 29 29.0–34.7 31.4 1.2
Percents of head length
Head depth 52.1 30 48.6–58.2 52.3 2.6
Head width 83.0 30 70.3–84.8 78.8 4.1
Snout length 67.0 30 54.8–68.9 63.8 3.1
Orbital diameter 14.9 30 14.3–18.6 16.0 1.0
Interorbital width 46.8 29 36.8–46.8 42.7 2.3
Mandibulary ramus length 11.2 30 7.8–15.6 12.9 1.7
Premaxillary ramus width 13.8 30 9.2–18.1 14.4 1.7
Eye–nostril distance 11.7 29 9.9–14.6 11.6 1.1
Snout–nostril distance 47.3 29 38.1–48.7 45.3 2.3
Internostril width 18.6 29 14.8–20.0 17.3 1.2
C. peduncle depth/C. peduncle length 24.9 30 20.1–26.3 23.9 1.3
Holotype nRange Mode
Median plates series 27 32 26–27 27
Predorsal plates 3 32 3–3 3
Dorsal plates below dorsal-fin base 7 32 6–9 7
Ventral plates in anal-fin base 2 32 2–3 2
Ventral plates between end anal-fin base and caudal fin 13 32 12–14 13
Dorsal-fin branched rays 7 29 7–7 7
Anal-fin branched rays 4 29 4–5 4
Caudal-fin branched rays 14 29 14 14
Pectoral-fin branched rays 6 29 6–6 6
Pelvic-fin branched rays 5 29 5–5 5
Premaxillary teeth 30 29 23–42 35
Dentary teeth 25 29 25–42 32
496 Copeia 2012, No. 3
H. cordovae, H. denticulatus,H. dlouhyi,H. fluviatilis,H.
heraldoi,H. hermanni,H. isbrueckeri,H. laplatae,H. latifrons,
H. mutucae,H. nigromaculatus,H. peckoltoides,H. piratatu,H.
topavae, H. uruguayensis, and H. variostictus) by the presence
of darker spot on body (vs. spot on body lighter than
background or absent). It can be distinguished from H.
brevis,H. denticulatus,H. derbyi,H. isbrueckeri,H. mutucae,H.
nigromaculatus, and H. peckoltoides by the abdomen entirely
covered by small plates (vs. completely naked or with wide
naked areas); from H. aspilogaster,H. commersoni,H. cordovae,
H. derbyi, and H. laplatae by having 26–27 lateral median
plates (vs. 29–31, 28–30, 28–30, 28, and 31–32, respectively);
from H. angipinnatus,H. borellii,H. boulengeri,H. fluviatilis,H.
heraldoi,H. hermanni,H. latifrons,H. piratatu,H. topavae, and
H. uruguayensis by the caudal peduncle depth, 20.1–26.3%in
its length (vs. 29–40%); from H. variostictus by the orbital
diameter 14.3–18.6%of head length (vs. 20%), and tip of
pelvic fin not reaching middle anal-fin base (vs. reaching);
from H. dlouhyi by having five longitudinal plate series at the
end of caudal peduncle (vs. four); and from H. ancistroides by
the weak keels on dorsal and mid-dorsal plate series, formed
by one or two well-developed odontodes on each plate (vs.
conspicuous keels, formed by a cluster of three or more well-
developed odontodes on each plate), and by the caudal
peduncle depth 7.4–8.6%in the SL (vs. 9.9–12.3%).
Description.—Morphometric and meristic data in Table 1.
Largest specimen examined 57.9 mm SL. Body deepest at
dorsal-fin origin. Dorsal profile of body straight from tip of
snout to nares; convex from nares to dorsal-fin origin;
straight along dorsal-fin base; slightly concave from base of
last dorsal-fin ray to caudal-fin base. Ventral profile of body
almost straight from tip of snout to pelvic-fin origin;
posterodorsally inclined from pelvic-fin origin to anal-fin
origin; straight from anal fin to caudal-fin origin. Caudal
peduncle rounded in cross-section, with shallow concave
area near caudal-fin origin. Anterior profile of snout
rounded in dorsal view. Head with lateral ridge from nares
to posterior margin of compound pterotic; distinctly raised
on posterodorsal corner of orbit, and poorly developed
along compound pterotic. Posterior process of supraoccip-
ital bordered by single predorsal plate, partially divided in
some specimens. Opercle hatchet-shaped, with small to
well-developed odontodes (up to 40). Preopercle without
odontodes, partially covered by three to four plates. Cheek
plates slightly evertible with large and strong odontodes.
Eye moderate in size, dorsolaterally placed; iris operculum
present, well developed; space between orbits concave;
dorsal edge of orbit slightly elevated. Nares separated by
flap of skin. Oral disk round, medium-sized. Outer edge of
upper lip without platelets and odontodes. Maxillary barbel
shorter than eye diameter, linked to lip proximally,
ornamented with small papillae. Lower lip with fringed
edges, not reaching transversal line through gill openings
and scapular bridge; its ventral surface covered with
numerous diminute papillae decreasing in size posteriorly,
except for smooth region posterior to symphysial region.
Dentary angle averaging 90uor more. Premaxillary teeth 23
to 42 (mode 35); dentary teeth 25 to 42 (mode 32). Teeth
bicuspid, curved inward distally; medial cusp considerably
larger than lateral cusp.
Tip of snout mostly naked, with two lateral pointed
patches of small plates with odontodes. Ventral surface of
head mostly naked, except for few small plates anterior to
gill opening. Five complete lateral series of plates. Dorsal
plate series with six to nine (mode seven) plates, limiting
naked area along dorsal-fin base. Median series with 26–27
(mode 27) perforated plates. Mid-ventral plates keeled up to
vertical through pelvic-fin origin. Ventral plate series
starting slightly posterior to vertical through pelvic-fin
origin, with two to three ventral plates along anal-fin base,
and 12 to 14 ventral plates from end of anal-fin base to
caudal fin. Dorsal, mid-dorsal, and mid-ventral plate series
with poorly developed longitudinal keels. Abdomen com-
pletely covered by small plates, except near pectoral-fin
insertion, around pelvic-fin base, and urogenital opening.
Pre-anal plate absent.
Dorsal fin with seven branched rays, posterior border
straight; tip of rays extending to vertical through median
portion of anal fin; well-developed, V-shaped spinelet
present, locking mechanism functional. Tip of first and last
proximal radials of dorsal fin contacting neural spine of
seventh and 16
vertebrae, respectively. Adipose fin absent;
typical azygous pre-adipose plates often absent (159 speci-
mens), some individuals with one to two (18 specimens;
Fig. 2). Pectoral fin with strong spine, covered by well-
developed odontodes distally on outer portion, and with six
branched rays; tip of adpressed pectoral fin reaching
posteriormost portion of pelvic-fin base. Cleithrum exposed
process located dorsally to pectoral-fin rays, tapering
posteriorly. Pelvic fin with thin and flexible spine, covered
by well-developed odontodes in ventral portion, and with
five branched rays; tip of adpressed pelvic fin surpassing
Fig. 2. Dorsal view of caudal peduncle of Hypostomus careopinnatus:
pre-adipose plates (A) absent, (B) present. DZSJRP 12447, paratypes,
42.4 and 56.3 mm SL, respectively.
Martins et al.—New Hypostomus from Paraguay River basin 497
anal-fin origin. Anal fin with thin and flexible unbranched
ray, and four or five branched rays; plate of first proximal
radial of anal fin exposed, with 10 to 20 odontodes; tip of
first and last proximal radials of anal fin contacting hemal
spine of 15
vertebrae, respectively. Caudal
fin concave, with two outer unbranched rays and 14 inner
branched rays; lower lobe longer than upper one; five dorsal
and four or five ventral procurrent rays. Vertebrae 31–32.
Color in alcohol.—Overall ground color light brown. Dorsal
and lateral surfaces of head and body covered by small dark
spots, about four times in orbital diameter; more numerous
on head. Ventral surface of body lighter than dorsal and
lateral, cream colored. Abdominal coloration varying, not
according to size; commonly, abdomen with dark spots
slightly larger, one to two times in orbital diameter,
occasionally extending to posteroventral portion of head;
some specimens with conspicuous dark vermiculations on
abdomen, other specimens with abdomen entirely pale.
Interradial membrane of fins mostly hyaline. Fin rays with
dark spots larger than those on ventral portion of body, also
over membrane, forming transverse bars more conspicuous
on the caudal fin.
Geographic distribution and habitat.—Only known from the
Rio Taquari drainage, upper Rio Paraguay basin, Alto
Araguaia, Mato Grosso State, Brazil (Fig. 3). The type locality
of Hypostomus careopinnatus is a small stream about two
meters wide and 0.4–0.5 m deep, silted, bottom predomi-
nantly with pebble, sand, and mud, running through a
grazing area, with very poor riparian vegetation.
Etymology.—The specific name careopinnatus comes from the
Latin careo, be deprived of, and the Latin pinnatus, with fin,
in reference to the absence of adipose fin.
Armbruster (2004) performed a morphological phylogenetic
analysis of Loricariidae emphasizing the Hypostominae and
redefined the genus Hypostomus based on three non-
exclusive synapomorphies: (1) hatchet-shaped opercle; (2)
anterior process of the compound pterotic passing halfway
through the orbit; and (3) pointed cleithral process.
Hypostomus careopinnatus share the three synapomorphies.
Several loricariids have lost the adipose fin (Pereira and
Reis, 1992; Fisch-Muller et al., 2005a, 2005b): all Loricar-
iinae, most of the Hypoptopomatinae, some Hypostominae,
and a species of Neoplecostominae (Neoplecostomus para-
nensis). Generally, the adipose fin is preceded by one or a
series of median pre-adipose plates. The presence of pre-
adipose plates is common in the Hypostominae which lack
adipose fin (e.g., most Pareiorhina,Rhinelepis spp.). Some of
them also have a raised crest on such plates (e.g., Ancistrus
Fig. 3. Type locality of Hypostomus careopinnatus, tributary of Rio Ariranha, Rio Taquari drainage, upper Rio Paraguay basin, Brazil.
498 Copeia 2012, No. 3
tombador,A. reisi,A. jataiensis,Corymbophanes spp., Pareio-
rhaphis vestigipinnis,Leptoancistrus spp., Pareiorhina carran-
cas); others lack the adipose fin and also the pre-adipose
plates (Pseudorinelepis and Hypostomus levis [Armbruster,
2003]). The new species herein described completely lacks
the adipose fin in all specimens collected. Also, Hypostomus
careopinnatus lacks the median pre-adipose plates in almost
all specimens examined (159), except for some (18) with one
or two, never with raised crest.
Within the genus, the loss of the adipose fin is a derived
feature shared by H. levis,aHypostomus cochliodon group
member. According to Armbruster (2003) and Armbruster
and de Souza (2005), the Hypostomus cochliodon group is
defined by the distinctly spoon-shaped teeth, the preoper-
culo-hyomandibular ridge deflected posterior to the main
body of the hyomandibula, and the dentary angle averaging
less than 80u.Hypostomus careopinnatus has elongate, not
spoon-shaped teeth, and dentary angle averaging 90uor
more, but, as in the members of the Hypostomus cochliodon
group, the preoperculo-hyomandibular ridge is deflected
posterior to the main body of the hyomandibula. We are
unable to establish any relationships of Hypostomus careo-
pinnatus and congeners. Nevertheless, the absence of the
adipose fin is probably an independent acquisition for the
new species and Hypostomus levis.
All from Brazil.
Rio Amazonas drainage: Pseudorinelepis genibarbis: MZUSP
26833, 1, 129.3 mm SL.
Rio Paraguay drainage: Hypostomus boulengeri:MZUSP
40092, 4, 98.0–142.7 mm SL; MZUSP 95075, 8, 15.3–
88.8 mm SL. Hypostomus cochliodon:DZSJRP3077,2,
100.9–110.4 mm SL; MZUSP 44313, 4, 31.2–112.9 mm SL;
MZUSP 60009, 17, 26.8–78.8 mm SL; MZUSP 87803, 1,
53.8 mm SL. Hypostomus dlouhyi: MZUSP 46036, 1, paratype,
153.0 mm SL. Hypostomus latifrons: MZUSP 38193, 2, 178.2–
232.3 mm SL. Hypostomus peckoltoides: MZUSP 105226,
holotype, 108.0 mm SL. Hypostomus piratatu: MZUSP 3234,
4, 107.13–129.7 mm SL; MZUSP 46038, holotype, 167.6 mm
SL. Hypostomus regani: MZUSP 78880, 2, 124.2–150.8 mm SL.
Plecostomus variostictus: MNRJ 1072, holotype, 31.9 mm SL.
Rio Parana´ drainage: Hypostomus ancistroides: DZSJRP 5806,
7, 30.6–92.5 mm SL; DZSJRP 6334, 3, 73.7–97.8 mm SL;
DZSJRP 7112, 18, 15.9–96.2 mm SL; DZSJRP 10788, 32, 15.9–
69.9 mm SL; DZSJRP 13311, 23, 34.1–71.2 mm SL.
159.5 mm SL. Hypostomus heraldoi: MZUSP 98771, holotype,
204.7 mm SL. Hypostomus margaritifer: MZUSP 43776, 1,
205.6 mm SL. Hypostomus regani: MZUSP 85891, 13, 70.1–
141.2 mm SL. Hypostomus topavae: MZUSP 97834, 8, 57.2–
114.3 mm SL. Plecostomus scaphyceps:MZUSP1014,1
paratype, 38.3 mm SL. Rhinelepis aspera: DZSJRP 4478, 2,
103.1–132.3 mm SL; DZSJRP 4495, 1, 114.1 mm SL.
Rio Uruguay drainage: Hypostomus isbrueckeri:MZUSP
40257, 3 of 5 paratypes, 81.6–150.8 mm SL. Hypostomus
roseopunctatus: MZUSP 40344, holotype, 211.5 mm SL.
C. Zawadzki provided some rare literature, A. Manzale
helped in the field expedition, and F. Pupo examined the
holotype of H. variostictus. Ame´rica Latina Logı´stica (ALL),
Ferronorte S/A, and Tetraplan Consultoria e Planejamento
gave financial support for field work. The authors were
supported by the Fundac¸a˜o de Amparo a` Pesquisa do Estado
de Sa˜o Paulo (2009/11873-0, FOM and 2009/15075-0,
MMFM) and Conselho Nacional de Desenvolvimento
Cientı´fico e Tecnolo´ gico (CNPq 306.988/2008-9, FL).
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