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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4144 (3): 443
–
450
http://www.mapress.com/j/zt/
Article
443
http://doi.org/10.11646/zootaxa.4144.3.10
http://zoobank.org/urn:lsid:zoobank.org:pub:B103BEBB-93A8-4EA8-B32B-DD68EBEC6DA5
Pareiorhaphis vetula, a new armored catfish from the headwaters of the Rio Doce
basin, Brazil (Siluriformes: Loricariidae)
EDSON H. L. PEREIRA
1
, PABLO LEHMANN A.
2
& ROBERTO E. REIS
1
1
PUCRS—Laboratório de Sistemática de Vertebrados. Av. Ipiranga, 6681, Caixa Postal 1429, 90619-900 Porto Alegre, RS, Brazil.
E-mail: ehlpereira@gmail.com and reis@pucrs.br
2
Universidade do Vale do Rio dos Sinos, Laboratório de Ictiologia, Av. Unisinos 950, 93022-000 São Leopoldo, RS, Brazil.
E-mail: pablole@unisinos.br
Abstract
A new species of Pareiorhaphis is described from the upper Rio Doce basin. The description is based on a series of spec-
imens recently collected in small headwater tributaries to the Rio Guanhães, a tributary of the Rio Santo Antonio, left bank
of the Rio Doce in Minas Gerais State, eastern Brazil. Pareiorhaphis vetula, new species, is a small loricariid catfish with
dark brown spots irregularly scattered over a brown background on the dorsal surface of body and along flanks. The new
species differs from all other Pareiorhaphis species by having the maxillary barbel completely adnate to the lower lip and
by adult males possessing a particularly elongate, sharply pointed, conical urogenital papilla. In addition, Pareiorhaphis
vetula is further distinguished from most congeners by having a shorter pelvic-fin spine, or by possessing more numerous
premaxillary teeth, and by lacking a dorsal-fin spinelet. A comparison with congeners P. nasuta, P. scutula and P.
proskynita, which also occur in headwater streams of the Rio Doce basin is also presented.
Key words: biodiversity, cascudos, neotropical, systematics, taxonomy
Introduction
The genus Pareiorhaphis Miranda Ribeiro, 1918 is presently composed of 23 species occurring from coastal river
basins in Bahia to northeastern Rio Grande do Sul, as well as in the upper tributaries to the São Francisco and
Paraná rivers. Pareiorhaphis species, including the new species being described herein, are primarily diagnosed
from remaining neoplecostomines by having the canal plate firmly articulated to the preopercle, and by a series of
sexually dimorphic features of hypertrophied odontodes of adult males (Pereira et al., 2007, 2010).
Three species of Pareiorhaphis are so far recognized from the Rio Doce basin, Pareiorhaphis nasuta Pereira,
Vieira & Reis, 2007, P. scutula Pereira, Vieira & Reis, 2010 and P. proskynita Pereira & Britto, 2012. These species
are diagnosed from each other based on features of eye diameter, snout length, arrangement of abdominal plates,
presence of an adipose fin, and by variations in the color pattern (Pereira & Britto, 2012).
In the present paper we report the discovery and describe another species of Pareiorhaphis from a pristine
headwater tributary to the Rio Doce basin in Santo Antônio do Itambé, Minas Gerais State. This discovery raises to
four the number of Pareiorhaphis species in the Rio Doce basin, making that river drainage the one with highest
diversity in this genus.
Material and methods
Measurements and counts were taken according to Pereira et al. (2007). Morphometric features were obtained with
digital calipers to the nearest 0.1 mm and were made from point to point under a stereomicroscope. Standard length
is expressed in millimeters while all other measurements are given as percent of standard length or head length.
Counts of procurrent caudal-fin rays, preadipose azygous plates, and vertebrae were made in cleared and
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counterstained specimen (c&s), which were prepared according to Taylor & Van Dyke (1985). Vertebral counts
include five centra in the Weberian Apparatus and the fused preural plus ural centra counted as one element.
Nomenclature and counts for body plates follow Schaefer (1997). Specimens examined belong to the Auburn
University Museum, Auburn (AUM), and Museu de Ciências e Tecnologia, Pontifícia Universidade Católica do
Rio Grande do Sul, Porto Alegre (MCP). Comparative material of Pareiorhaphis species listed in Pereira et al.
(2012), except for Pareiorhaphis garbei, MCP 26524, 10, Rio Aleixo, tributary to Rio Macacu, Itaboraí, Rio de
Janeiro, Brazil, and P. nudulus, MCP 29152, 23, Arroio Bananeira, tributary to Rio Três Forquilhas, Itati, Rio
Grande do Sul, Brazil. In the lists of type material museum abbreviation and catalog number come first, followed
by the number and SL range of specimens in that lot, and locality data.
Pareiorhaphis vetula, new species
Fig. 1 Table 1
Holotype. MCP 49693, 48.2 mm SL, male. Brazil, Minas Gerais State, Santo Antônio do Itambé, Rio Mãe d’Água
at Cachoeira da Ponte de Pedra, tributary to Rio Guanhães, Rio Doce basin, 18°27’16.5”S 43°18’29.7”W, 26 July
2015, R. E. Reis, P. Lehmann A., & E. H. L. Pereira col.
Paratypes. All from Brazil, Minas Gerais State, Santo Antônio do Itambé: AUM 67215, 2 females, 44.9–45.4
mm SL; MCP 49169, 2 males and 5 females, 42.1–48.6 mm SL (1 female with caudal peduncle broken not
measured), same data as holotype. MCP 49149, 3 males and 2 females, 40.5–49.3 mm SL + 1 male c&s, 44.5 mm
SL, Córrego Beirão da Areia below Cachoeira da Fumaça, tributary to Rio Mãe d’Água, itself tributary to Rio
Guanhães, Rio Doce basin, 18°27’27.7”S 43°19’54.3”W, 26 July 2015, R. E. Reis, P. Lehmann A., & E. H. L.
Pereira col.
Diagnosis. Pareiorhaphis vetula is distinguished from all Pareiorhaphis species by having the maxillary
barbel completely adnate to the lower lip (a vestigial nub present in one specimen), Fig. 2 (vs. maxillary barbel free
and conspicuous in other species) and by adult males possessing a particularly elongate, sharply pointed, conical
urogenital papilla, which distinctly passes the level of the anal opening, Fig. 3 (vs. urogenital papilla short and
extending to approximately the same level of the anal opening). The new species can be distinguished from all
Pareiorhaphis species except, P. calmoni, P. garapia, and P. splendens by having a shorter pelvic-fin spine, 15.5–
17.6% SL (vs. 18.3–26.4% SL). Additionally, the new species is distinguished from P. c a l m o n i and P. garapia by
having more numerous premaxillary teeth, 84–103 (vs. 45–81 and 64–81, respectively); and from P. s p l e n d e n s by
lacking a dorsal-fin spinelet (vs. spinelet present). The new species is further distinguished from remaining
Pareiorhaphis species in the Rio Doce basin by having a much smaller adult body size, with largest mature
specimen examined 49.3 mm SL, contrary to 94.7 mm SL (P. nasuta), 90.5 mm SL (P. sc u tu la ) and 89.7 mm SL (P.
proskynita), and by lacking a dorsal-fin spinelet (vs. spinelet present).
Description. Counts and proportional measurements in Table 1. Small-sized loricariid with standard length of
measured specimens 40.5–49.3 mm SL. Body elongate, moderately depressed. Greatest body width at cheeks
(males) or cleithrum (females), progressively tapering to end of caudal peduncle. Dorsal profile of body convex
from snout tip to dorsal-fin origin, straight to slightly concave from that point to origin of adipose fin, and slightly
concave from adipose spine to caudal fin. Greatest body depth at dorsal-fin origin. Least body depth at shallowest
portion of caudal peduncle. Trunk and caudal peduncle mostly oval in cross-section, slightly flattened ventrally and
more compressed caudally. Lateral-line canal in median series complete, pored tube visible from compound
pterotic to caudal-fin base. Ventral profile nearly straight between snout tip and pelvic girdle, slightly convex
between pelvic and anal fins, and slightly concave along caudal peduncle. Dorsal and lateral surfaces of body
covered by dermal plates. Predorsal area covered by plates arranged in three or four series. Five lateral rows of
dermal plates covering body, not forming keels. Ventral surface of head and abdomen completely naked up to
urogenital opening.
Head broad and moderately depressed. Outline of head widely round (males) to squarish (females) in dorsal
view (Fig. 4). Interorbital space slightly concave. Three weakly elevated ridges between orbits and snout tip
formed by underlying bones, without emerging hyperthrophied odontodes. Central ridge on snout more prominent.
Snout convex in lateral profile; mostly covered by dermal plates, though lacking rostral plate. Adult male with
small soft fleshy lobe extending along lateral portion of head. Soft fleshy area of cheek and opercle ornamented
with hypertrophied odontodes; lateral process of cleithrum without hypertrophied odontodes. Eye small,
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dorsolaterally placed; orbital diameter 9.9–12.9% HL. Iris operculum absent, pupil circular. Nares ovoid, slightly
longer than wide, positioned much closer to anterior margin of orbit than to snout tip. Lips well developed, widely
oval transversely and occupying most of ventral surface of head; lips wider than snout in females, visible dorsally.
Lower lip wide and long, reaching to pectoral girdle in some specimens, upper lip narrow. Lower lip densely
covered by minute papillae; papillae decreasing in size towards edge. Margin of lower lip finely fringed. Maxillary
barbel completely adnate to the lower lip (a vestigial nub present in one specimen). Teeth series in both premaxillae
and dentaries with mesial ends slightly curved inwards. Teeth slender, asymmetrically bifid, medial cusp long and
pointed; lateral cusp small and pointed, with about one fifth length of medial cusp in unworn teeth.
Dorsal-fin origin along vertical passing through origin of pelvic-fin ray. Dorsal fin short, not contacting
preadipose azygous plates when adpressed. Nuchal plate exposed in front of dorsal fin. Dorsal-fin spinelet absent.
Dorsal-fin spine moderately flexible, followed by seven branched rays. Adipose fin with well-ossified leading
spine bearing odontodes. Adipose-fin membrane very short and not extended beyond adipose-fin spine. Adipose
fin preceded by one median, azygous, preadipose plate. Pectoral fin small, with spine slightly curved and flattened,
covered by minute odontodes in females, immature males and juveniles. Adult male with pectoral-fin spine
broadened and bearing straight to slightly curved, short hypertrophied odontodes on entire outer face. Pectoral fin
with six branched rays, first slightly longer than spine. Subsequent branched rays decrease gradually in size, last
ray two thirds length of first ray. Distal margin of pectoral fin slightly rounded, just surpassing origin of pelvic fin
when adpressed. Pelvic fin with one unbranched and five branched rays, not reaching to anal-fin origin when
adpressed. Pelvic-fin unbranched ray depressed, covered by minute odontodes ventrally and laterally and with
small dermal flap on dorsal surface in males. Odontodes on anal-fin spines somewhat flat and spatulate. Anal fin
short with one unbranched and five branched rays. Anal-fin origin along vertical passing between base and tip of
depressed last dorsal-fin ray. Caudal fin concave; lower lobe slightly longer than upper; 14 branched rays. Upper
caudal-fin lobe with five and lower lobe with four plate-like procurrent rays, posteriormost elongate. Odontodes on
principal and procurrent rays small and irregularly arranged. Total vertebral centra 30; hypural plate symmetrical.
Color in alcohol. Overall background color of dorsal and lateral surface of head and trunk light to dark brown;
mostly unpigmented and yellowish pale ventrally. Predorsal region and area around dorsal fin slightly lighter than
head. Fleshy lateral margins of head in mature males pale yellow. Head, dorsum and flanks with irregular and
inconspicuous scattered dark brown dots; dots not forming transverse saddles and somewhat lined along lateral
line. Lower lip plain yellowish pale; upper lip with dark dots on dorsal surface. Ventral surface of head and
abdomen completely pale yellow, with some concentration of dark brown chromatophores on lateral portion of
head and caudal peduncle. Dorsal, pectoral and pelvic fins with two or three inconspicuous, transverse rows of dark
dots on rays. Anal fin mostly unpigmented. Caudal fin homogeneously dark or with scattered, inconspicuous,
lighter areas.
Sexual dimorphism. Mature males with small fleshy lobes on lateral margins of head, with hypertrophied
odontodes on cheek and opercle. Pectoral-fin spine distinctly thickened and bearing hyperthrophied odontodes. In
addition, males (Fig. 1) with much broader head while females with narrower head which allows lateral margins of
lower lip to be visible dorsally (Fig. 4). Males also with large and pointed urogenital papilla (Fig. 3), absent in
females, plates in dorsal and lateral portions of body covered by enlarged odontodes, giving hispid appearance, and
small dermal flap on dorsal surface of pelvic-fin spine.
Distribution. Pareiorhaphis vetula is known from two localities, one at Rio Mãe d’Água and one of its
tributary creeks, the Córrego Beirão da Areia, in the upper Rio Guanhães drainage, Rio Doce basin, eastern Brazil
(Fig. 5).
Etymology. Pareiorhaphis vetula, from the Latin vetulus, old or a little old man, in allusion to the small size of
adult specimens. An adjective.
Ecological notes. Pareiorhaphis vetula is known from two localities in the upper portion of Rio Guanhães, Rio
Doce basin, at an approximate altitude of 755 and 850 m above sea level, upstream the town of Santo Antonio de
Itambé, Minas Gerais State, Brazil (Fig. 6). The creeks where specimens have been collected possess tea-colored
dark water with pools of slow current alternated by rapids, with depths between 0.3 and 2 m, and approximately 5
to 6 m wide. The substrate is formed by big rocks, pebbles and sand, and the marginal vegetation is composed of
grasses and mostly riparian forest. On the Córrego Beirão da Areia, the new species was determined to be syntopic
with Astyanax rivularis, Trichomycterus sp., and another, probably undescribed, species of Pareiorhaphis,
currently under study by the authors. It was the only species collected at the Rio Mãe D’Água.
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TABLE 1. Morphometrics and meristics of the holotype and paratypes of Pareiorhaphis vetula. Measurement values are
given as percentage of standard length or head length. SD = standard deviation, H = holotype, n=number of specimens.
H n Low High Mean SD
Standard length (mm) 48.2 15 40.5 49.3 45.5
Percent of standard length
Head length 35.6 15 34.7 37.8 36.3 0.87
Predorsal length 45.1 15 44.3 47.9 45.7 0.89
Postdorsal length 36.5 15 35.4 40.1 37.5 1.37
Preanal length 67.5 15 64.4 68.0 66.7 1.21
Preadipose length 77.7 15 73.8 78.0 76.8 1.13
Dorsal-fin spine length 17.9 15 16.2 21.3 19.5 1.57
Anal-fin unbranched ray length 10.9 15 9.5 12.7 11.1 0.86
Pectoral-fin spine length 15.8 15 15.8 20.8 17.9 1.57
Ventral-fin unbranched ray length 15.8 15 15.5 17.6 16.6 0.61
Upper principal caudal-fin ray 23.4 15 20.9 25.5 23.7 1.26
Lower principal caudal-fin ray 26.3 15 25.2 30.6 27.4 1.55
Adipose-fin spine length 7.5 14 6.8 10.0 8.6 0.99
Adipose to caudal fin distance 22.2 15 20.6 26.3 22.5 1.46
Trunk length 17.6 15 17.1 20.4 19.2 0.98
Abdominal length 19.4 15 19.2 21.7 19.9 0.73
Cleithral width 29.2 15 29.2 33.0 30.8 1.02
Body depth at dorsal-fin origin 18.1 15 17.5 21.3 18.7 1.08
Body width at dorsal-fin origin 20.5 14 18.9 22.8 20.6 1.12
Body width at anal-fin origin 11.4 14 9.8 12.8 11.6 0.78
Caudal peduncle length 34.0 15 32.4 35.9 33.6 1.03
Caudal peduncle depth 7.8 15 7.5 9.5 8.5 0.63
Caudal peduncle width 4.6 15 3.8 5.0 4.5 0.32
Percent of head length
Snout length 68.2 15 67.3 73.1 70.0 1.51
Orbital diameter 11.4 15 9.9 12.9 11.7 0.87
Interorbital width 25.0 15 25.0 31.2 28.0 1.87
Head depth 48.3 15 46.0 53.3 49.5 2.18
Mandibular ramus left 26.2 15 24.2 26.8 25.7 0.79
Mandibular ramus rigth 26.2 15 22.1 26.6 25.6 1.17
Counts Mode
Premaxillary teeth left 86 14 84 99 86
Premaxillary teeth right 84 14 84 103 92
Dentary teeth left 84 15 84 97 89
Dentary teeth right 94 15 83 97 93
Plates in median lateral series left 24 14 24 26 25
Plates in median lateral series right 25 15 24 26 25
Plates at dorsal-fin base 7 15 5 7 6
Plates between dorsal and adipose 5 15 5 7 6
Plates between adipose and caudal 5 15 4 6 4
Plates at anal-fin base 4 15 2 4 3
Plates between anal and caudal 10 15 9 12 10
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FIGURE 1. Pareiorhaphis vetula, MCP 49693, holotype, 48.2 mm SL, male. Brazil, Minas Gerais State, Santo Antônio do
Itambé, rio Mãe d’Água at Cachoeira da Ponte de Pedra, tributary to Rio Guanhães, Rio Doce basin, 18°27’16.5”S
43°18’29.7”W.
All type specimens were collected upstream the town of Santo Antônio de Itambé, where the river is still well
preserved. However, the tourism at the Cachoeira da Fumaça waterfall is intensive and can pose some threat to the
environment. The recent ecological disaster which occurred in Mariana, middle Rio Doce in 2015 highlighted the
severe anthropic impact that has been occurring in that basin, and additional inventory efforts to adequately map
the species distribution and possibly discover additional unraveled diversity would be highly welcome.
Conservation notes. Pareiorhaphis vetula appears to be restricted to small, headwater tributaries to the Rio
Guanhães, in the region of Santo Antônio de Itambé, Minas Gerais State, Brazil. Considering that specific threats
to the species were not detected in the area, and the existence of a Conservation Unit (Pico do Itambé State Park)
encompassing a portion of those headwaters, P. vetula can be categorized as Least Concern (LC) according to the
International Union for Conservation of Nature (IUCN) categories and criteria (IUCN Standards and Petitions
Subcommittee, 2016).
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FIGURE 2. Maxillary barbel of Pareiorhaphis. (A) Barbel mostly free, P. garbei, MCP 26524, 89.1 mm SL. (B) Barbel mostly
adnate to lower lip, P. nudulus, MCP 29152, 37.0 mm SL. (C) Barbel completely adnate to lower lip, P. vetula, MCP 49169, 47.9
mm SL, paratype. Scale bar = 2 mm.
FIGURE 3. Urogenital papilla of Pareiorhaphis. (A) P. vetula, MCP 49149, paratype, 43.9 mm SL, male. (B) P. nudulus. MCP
29152, 37.0 mm SL, male. ao = anal opening; up = urogenital papilla. Scale bar = 2 mm.
Discussion
In general body shape Pareiorhaphis vetula resembles the other species occurring in the Rio Doce basin, P. nasuta,
P. scutula, and P. proskynita, by having a long snout with small eyes, and a broadly rounded head with thin, short
hyperthrophied odontodes on the cheeks of mature males. The new species, however, has a much smaller adult
size, with the largest mature specimen examined reaching to less the 50 mm SL, contrary to the three other species
that reach 94.7 mm SL (P. n a s ut a ), 90.5 mm SL (P. scutula) and 89.7 mm SL (P. proskynita) as maximum
measured size. Further, on its smaller size and contrary to the three larger species, P. vetula lacks a dorsal-fin
spinelet, a feature which may be related to the size reduction.
The maxillary barbel of Pareiorhaphis vetula is the smallest and least conspicuous of all Pareiorhaphis
species, not being visible externally (Fig. 2C). In fact, the cartilage at the distal extremity of the maxilla is present,
but it is completely included inside the lower lip. In most loricariids, however, the maxillary barbel possess a
conspicuous free portion as in P. garbei (Fig. 2A), while in other species it is mostly adnate to the lateral margin of
the lower lip as in P. nudulus (Fig. 2b).
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FIGURE 4. Pareiorhaphis vetula, MCP 49169, paratype, 47.0 mm SL, female. Brazil, Minas Gerais State, Santo Antônio do
Itambé, rio Mãe d’Água at Cachoeira da Ponte de Pedra, tributary to Rio Guanhães, Rio Doce basin, 18°27’16.5”S
43°18’29.7”W.
FIGURE 5. Drainage map of eastern Brazil, showing the known distribution of Pareiorhaphis species. Pareiorhaphis vetula
(circle), P. proskynita (dot), P. scutula (squares), and P. n a s u t a (triangles). Each symbol may represent more than one lot or
locality.
Among Loricarioids, several groups exhibit secondary sexual dimorphism in the possession of a distinct
urogenital papilla behind the anal opening. Males of Callichthyidae, Astroblepus, Lithogenes, and the Delturinae
share the possession of a large urogenital papilla, which clearly protrudes as an elongate projection behind the anal
opening. Among remaining loricariids, the urogenital papilla of males is mostly absent in both the Loricariinae and
the Hypostominae, and is always present in the Neoplecostominae and the Hypoptopomatinae as a small, conical
papilla typically concealed behind the anal tube. Adult males of Pareiorhaphis vetula, however, possess a
particularly large, sharply pointed, conical urogenital papilla. Such a papilla is not only proportionally larger than
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in other Pareiorhaphis species, but it is also somewhat projected ventrally, distinctly passing the level of the anal
opening (Fig. 3), contrary to other species where it points more posteriorly and reaches to approximately the same
level of the anal opening.
FIGURE 6. The Rio Mãe d’Água at Cachoeira da Ponte de Pedra, tributary to Rio Guanhães, Rio Doce basin, Santo Antônio
do Itambé, Minas Gerais State, Brazil.
Acknowledgements
We are grateful to Alfredo Carvalho for calling our attention to the Pareiorhaphis species in the Rio Mãe D’Água.
We thank Carlos and Margarete Lucena for help provided at the MCP fish collection, Bárbara Calegari for the
photographs in Figs. 1 and 4, Juliano Romanzini for helping with the microphotography (Figs. 2 and 3), and Carlos
Oliveira and Priscilla C. Silva for the identification of Astyanax rivularis. RER and PLA are partially funded by the
Conselho Nacional de Desenvolvimento Científico e Tecnológico - CNPq (process #306455/2014-5 and #483060/
2013-5, respectively).
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