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A New Species of the Electric Fish Genus Hypopygus (Gymnotiformes:
Hypopomidae) from the Lower Amazon Basin, Brazil
Author(s): Luiz Antnio Wanderley Peixoto , Guilherme Moreira Dutra , Carlos David de Santana , and
Wolmar Benjamin Wosiacki
Source: Copeia, 2013(2):232-237. 2013.
Published By: The American Society of Ichthyologists and Herpetologists
DOI: http://dx.doi.org/10.1643/CI-12-087
URL: http://www.bioone.org/doi/full/10.1643/CI-12-087
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A New Species of the Electric Fish Genus Hypopygus
(Gymnotiformes: Hypopomidae) from the Lower Amazon Basin, Brazil
Luiz Antonio Wanderley Peixoto
1
, Guilherme Moreira Dutra
1
,
Carlos David de Santana
1,2
, and Wolmar Benjamin Wosiacki
1
Hypopygus benoneae, new species, is described from streams in the lower Rio Anapu, in the eastern portion of the
Amazon basin, Brazil. The new species is diagnosed from all congeners by a unique set of characters, including the
absence of the sixth infraorbital bone and 115134 total anal-fin rays. Hypopygus benoneae, new species, is a member of
a clade also comprising H. hoedemani, H. lepturus, and H. minissimus. The description of H. benoneae, new species, brings
the number of species in Hypopygus to nine. A dichotomous key to the species of Hypopygus is provided.
Hypopygus benoneae, nova especie, e descrita para igarapes do baixo Rio Anapu, na porcao oriental da bacia Amazonica,
Brasil. A nova especie e diagnosticada de todos os congeneres por uma combinacao u nica de caracteres, tais como a
ausencia do sexto osso infraorbital e o nu mero total de raios da nadadeira anal 115134. Hypopygus benoneae, nova
especie, e um membro do clado que tambem inclui H. hoedemani, H. lepturus, and H. minissimus. A descricao de H.
benoneae, nova especie, eleva o nu mero de especies em Hypopygus para nove. Uma chave dicotomica para as especies de
Hypopygus e fornecida.
E
LECTRIC fishes of the genus Hypopygus are small-
sized species of less than 150 mm total length and
include the smallest known gymnotiform species, H.
minissimus, which reaches only approximately 55 mm total
length. Species of Hypopygus occur in streams and small
rivers of the Amazon and Orinoco basins, drainages of the
Guiana Shield, and limited portions of the R o Paraguay
basin (de Santana and Crampton, 2011). All species of
Hypopygus can be promptly recognized within the Gymno-
tiformes by the possession of a postpectoral accessory
electric organ (de Santana and Crampton, 2011:fig. 15).
Monophyly of the genus is supported by 13 synapomor-
phies, some of which are related to a set of reductive
morphological characters associated with miniaturization
(de Santana and Crampton, 2011).Their electric organs
generate tetraphasic pulse-type discharges ranging from 34
to 85 Hz (de Santana and Crampton, 2011).
Hypopygus was proposed by Hoedeman (1962) for H.
lepturus from the Marowijne River drainage in Suriname
and was later reported from several localities in the
Amazon and Orinoco basins, the major drainages of the
Guianas, and some northern tributaries of the R o Paraguay
basin. The genus remained monotypic for 32 years, until
Mago-Leccia (1994) described H. neblinae from the R o
Bar a in Venezuela, with that species now known to be
widespread in the Orinoco and Rio Negro basins, and Rio
Preto da Eva in Brazil. Recently, de Santana and Crampton
(2011) revisited the genus and recognized six additional
species: H. cryptogenes, originally described as Stegostenopos
cryptogenes by Triques (1997), from the lower or middle Rio
Negro and the Rio Preto da Eva, both in the Amazon basin;
H. hoedemani also from the Rio Negro and Rio Preto da Eva;
H. isbruckeri from the upper R o Orinoco; H. minissimus
from the upper R o Orinoco and Rio Negro; H. nijsseni from
the upper Amazon near Tefe; and H. ortegai from localities
near Iquitos and Jenaro Herrera, in the upper Amazon
basin.
During an expedition to the lower Rio Anapu basin,
Floresta Nacional de Caxiuana, in the lower Amazon basin
in Brazil, a previously unknown species of Hypopygus was
collected and is described herein. This brings the total
species of Hypopygus to nine.
MATERIALS AND METHODS
All measurements were taken point-to-point to the nearest
0.1 mm with digital calipers under a stereomicroscope,
preferably on the left side. Measurements followed de
Santana and Crampton (2011). The nomenclature for the
lateral line system followed Arratia and Huaquin (1995). The
frequency of pectoral-fin rays and scales count are given in
parentheses after each count. An asterisk indicates counts
for the holotype. Specimens were cleared and counter-
stained (CS) according to Taylor and Van Dyke (1985).
Institutional abbreviations are as listed at http://www.asih.
org/node/204. Measurement abbreviations used in the text
are: LEA 5 length to the end of the anal fin, HL 5 head
length, and CFL 5 caudal filament length. For additional
comparative material examined see de Santana and Cramp-
ton (2011). Key to the species of Hypopygus was modified
from de Santana and Crampton (2011).
RESULTS
Hypopygus benoneae, new species
Figure 1, Table 1
Steatogenys elegans [in part].Montag et al., 2008:32 [in
listing of species].
Hypopygus lepturus [in part].de Santana and Crampton,
2011:1134 [in material examined].
1
Museu Paraense Em lio Goeldi, Caixa Postal 399, 66040-170, Belem, PA, Brazil; E-mail: (LAWP) luizwp@yahoo.com.br; (GMD)
guilhermedutr@yahoo.com.br; (CDS) csantana@museu-goeldi.br; and (WBW) wolmar@museu-goeldi.br. Send reprint requests to WBW.
2
Division of Fishes, Department of Vertebrate Zoology, MRC-159, National Museum of Natural History, P.O. Box 37012, Smithsonian
Institution, Washington, D.C. 20013-7012.
Submitted: 19 July 2012. Accepted: 25 November 2012. Associate Editor: R. E. Reis.
F 2013 by the American Society of Ichthyologists and Herpetologists DOI: 10.1643/CI-12-087
Copeia 2013, No. 2, 232237
Holotype.MPEG 23603, 1, 46.6 mm LEA, Brazil, Para,
Melgaco, tributary of Igarape Acu , lower Rio Anapu
drainage, 01u48915.40S, 51u28932.20W, N. Benone, 17 No-
vember 2010.
Paratypes.All from Brazil, Para, lower Rio Anapu drainage:
MPEG 23604, 3, 28.652.3 mm LEA, USNM 408391, 1,
31.0 mm LEA, collected with holotype; MPEG 3729, 1,
59.1 mm LEA, MZUSP 112205, 1, 49.6 mm LEA, Melgaco,
Igarape Curuazinho, M. I. Assuncao, 15 August 1993; INPA
38237, 1, 27.6 mm LEA, MPEG 18713, 1, 35.6 mm LEA,
Portel, Igarape Caquajo , 01u59947.40S, 51u379210W, L. F. A.
Montag and others, 2 September 2009; MPEG 23605, 1 CS,
41.1 mm LEA, Melgaco, Igarape Pereira, 01u47917.00S,
51u28954.20W, N. Benone, 27 November 2010; MPEG 3731,
2 CS, 29.331.1 mm LEA, Igarape Curuazinho, M. I.
Assuncao, 14 August 1993; MPEG 24295, 1, 51.2 mm LEA,
Igarape Caju , 01u46925.70S, 51u30919.50W, N. Benone, 5 July
2010; MPEG 24296, 4, 26.231.0 mm LEA, Igarape Santa
Maria, 01u46938.60S, 51u28957.80W, N. Benone, 25 Novem-
ber 2010.
Diagnosis.Hypopygus benoneae is diagnosed from H. crypto-
genes by the absence of the sixth infraorbital bone (vs.
presence); the total number of anal-fin rays, 115134
(vs. 148174); the head length, 13.720.2% of LEA (vs.
10.312.1%); the interocular width, 26.135.4% of HL (vs.
15.021.1%); the caudal filament depth, 1.86.8% of CFL
Fig. 1. Hypopygus benoneae, MPEG 23603, 46.6 mm LEA, holotype, Brazil, Para, Melgaco, lower Rio Anapu drainage, 01u48915.40S, 51u28932.20W.
(A) Head; (B) left lateral view; (C) dorsal view.
Peixoto et al.New species of Hypopygus 233
(vs. 7.815.3%); the absence of the extrascapular canal (vs.
presence); the absence of the fourth mandibular canal (vs.
presence); the absence of the fifth mandibular canal (vs.
presence); the number of precaudal vertebrae, 1516 (vs.
14); and the second basibranchial ossified (vs. cartilaginous).
Hypopygus benoneae is distinguished from H. hoedemani by
the absence of the anteriormost branchiostegal ray (vs.
presence; for the homology of the branchiostegal rays in
Gymnotiformes see de Santana and Vari, 2010:246247); the
presence of the third mandibular canal (vs. absence); the
second basibranchial ossified (vs. cartilaginous) and the
presence of the supraorbital canal (vs. absence). Hypopygus
benoneae is diagnosed from H. isbruckeri by the absence of
the sixth infraorbital canal (vs. presence); the total number
of anal-fin rays, 115134 (vs. 136152); the number of scales
above the lateral line, four to six (vs. seven or eight); the
absence of the extrascapular canal (vs. presence); the
absence of the fourth mandibular canal (vs. presence); the
absence of the fifth mandibular canal (vs. presence); and the
number of precaudal vertebrae, 1516 (vs. 14). Hypopygus
benoneae is distinguished from H. lepturus by the number of
pectoral-fin rays, 1113 (vs. 9 or 10); the interocular width,
26.134.5% of HL (vs. 18.724.4%) and the absence of the
extrascapular canal (vs. presence). Hypopygus benoneae is
differentiated from H. minissimus by the presence of dark
oblique bands on the lateral surface of the body (vs.
absence); the presence of the scales along the middorsal
surface of the body (vs. absence); the number of scales below
the lateral line, 57 (vs. 4); the caudal filament length, 21.2
37.7% of LEA (vs. 45.977.3%); the interocular width, 26.1
34.5% of HL (vs. 15.021.1%); the anteriormost branchios-
tegal ray absent (vs. present); the presence of the second
mandibular canal (vs. absence); the presence of the third
mandibular canal (vs. absence); and the second basibran-
chial ossified (vs. cartilaginous). Hypopygus benoneae is
distinguished from H. neblinae by the absence of the sixth
infraorbital canal (vs. presence); the total number of anal-fin
rays, 115134 (vs. 137145); the interocular width, 26.1
34.5% HL (vs. 14.223.3%); the absence of the extrascapular
canal (vs. presence); the absence of the fourth mandibular
canal (vs. presence); the absence of the fifth mandibular
canal (vs. presence); and the number of transitional
precaudal vertebrae, 2 (vs. 3). Hypopygus benoneae is
diagnosed from H. nijsseni by the absence of the sixth
infraorbital canal (vs. presence); the total number of anal-fin
rays, 115134 (vs. 145154); the head depth at the eye, 44.0
62.8% HL (vs. 64.669.9%); the absence of the extrascapular
canal (vs. presence); the absence of the fourth mandibular
canal (vs. presence); the absence of the fifth mandibular
canal (vs. presence); and the number of transitional
precaudal vertebrae, 2 (vs. 4). Hypopygus benoneae is
diagnosed from H. ortegai by having the dorsal rami of
intermittent branch of anterior lateral line nerve visible as
two parallel lines (vs. not visible); the absence of the sixth
infraorbital canal (vs. presence); the total number of anal-fin
rays, 115134 (vs. 137156); the interocular width, 26.1
34.5% of HL (vs. 18.724.4%); the pectoral-fin length, 45.2
59.3% of HL (vs. 64.669.9%); the absence of the extra-
scapular canal (vs. presence); the absence of the fourth
mandibular canal (vs. presence); the absence of the fifth
mandibular canal (vs. presence); and the number of
transitional precaudal vertebrae, 2 (vs. 3).
Description.Head, body shape, and pigmentation illustrat-
ed in Figure 1. Morphometric data for examined specimens
in Table 1. Body size small, maximum examined total
Table 1. Morphometric Data for Holotype and Paratypes of Hypopygus benoneae, New Species. Range includes the values for the holotype (n,
number of specimens; SD, standard deviation).
Holotype n Min Max Mean SD
Total length 72.8 16 37.3 92.1
Length to end of anal fin (mm) 46.6 16 26.2 59.1
Caudal filament length (mm) 26.4 16 10.8 36.2
Head length (mm) 7.0 17 4.7 8.1
Body cavity (mm) 3 9.5 14.4
Percentage of length to end of anal fin
Anal fin length 79.9 16 76.9 89.0 79.4 3.0
Anus to anal-fin base 10.2 15 9.4 16.0 11.9 2.1
Body depth at anal-fin origin 14.4 16 13.3 18.5 15.8 1.5
Body width at anal-fin origin 5.7 16 5.2 8.9 7.3 1.0
Caudal filament length 56.6 15 45.9 77.3 55.4 8.9
Head length 15.0 16 13.7 20.2 16.5 1.8
Snout to occiput 11.0 16 9.8 15.4 12.3 1.7
Percentage of head length
Head depth at eye 50.6 17 44.0 62.8 50.8 4.2
Head width at eye 42.3 17 38.6 51.7 43.3 3.4
Interocular width 27.7 17 26.1 34.5 28.8 2.0
Pectoral fin length 51.4 16 45.2 59.3 53.1 4.5
Orbital diameter 12.8 17 9.8 15.7 13.2 1.6
Postorbital length 66.3 17 47.2 66.3 59.7 5.4
Snout length 21.9 17 20.3 28.6 24.9 2.0
Percentage of caudal filament length
Caudal filament depth 2.9 16 1.8 6.8 3.0 1.2
234 Copeia 2013, No. 2
length 92.1 mm (n 5 16). Body elongate and distinctly
laterally compressed, more so posterior to abdominal region.
Greatest body depth along, or slightly posterior to, abdom-
inal cavity. Dorsal profile of body straight or slightly convex.
Ventral profile slightly convex. Caudal filament elongate.
Anus and urogenital papilla adjacent, positioned through
vertical between posterior margin of eye and anterior
margin of opercle.
Head compressed with greatest width at opercular region
and greatest depth through nape. Snout rounded in lateral
view. Mouth subterminal, upper jaw extends more anteriorly
than lower jaw. Mandibular canal bone, 3(1). Rictus extend-
ing posteriorly to point slightly anterior to vertical through
anterior margin of eye. Anterior naris inside upper lip.
Posterior naris absent. Eye small, circular, completely covered
by thin membrane, located on anterior half of head length,
and laterally oriented. Gill opening constricted to short
vertical aperture along posterior margin of opercle and
slightly anterior to vertical through pectoral-fin origin. Gill
opening extends above and below pectoral-fin base. Branchi-
al membranes joined at isthmus. Branchiostegal rays, 4(2).
Postpectoral electric organ with two columns and six rows
of electrocytes, its dorsal groove extending anteriorly to
approximately one and a half to two orbital diameters
behind posterior border of eye.
Scales small, cycloid, extending from immediately poste-
rior of head to caudal filament. Scales present on mid-dorsal
region of body. Scales above lateral line at midbody 4(2),
5*(14), or 6(1). Scales below lateral line 5(2), 6*(10), or 7(5).
Anteriormost perforated lateral line scale along vertical
through pectoral-fin origin. Lateral line interrupted.
Pectoral-fin rays 11*(12), 12(4), or 13(1). Distal margin of
fin rounded. Tip of pectoral-fin reaching vertical through
bases of second to seventh anal-fin ray. Pectoral-fin radials,
4. Anal-fin rays ixii,109131(115134 total anal-fin rays;
vi,111*). Anal-fin origin located at vertical through fourth
pleural rib. Precaudal vertebrae 15(2) or 16(1) with 13(2) or
14(1) anterior and 2(3) transitional vertebrae. Pleural ribs,
8(1) or 9(2). Anterior displaced hemal spines, 3(1). Posterior
displaced hemal spines, 1(1).
Coloration in alcohol.Background coloration on body dark
brown, darker over mid-dorsal region. Fourteen to 21 pale
bands apparent from dorsal view between point through
opercular opening and distal portion of caudal filament.
Abdomen dark brown, with tiny diffuse clear areas and pale
yellowregion around anus. Dorsal region of head dark brown
with tiny pale area between orbits. Head dark brown laterally
with pale area posteroventral to orbit. Cheek with irregular
dark brown region, occasionally forming sub-orbital stripe,
extending from ventral margin of orbit to ventral margin of
head. Lips with same pigmentation pattern as head, becom-
ing gradually darker posteriorly. Preopercle pale overall but
with posterior margin dark brown. Opercle uniformly dark.
Opercular membrane hyaline.
Body with 5 to 12 irregular clear bands in lateral view.
Clear bands in small specimens (26.231.1 mm LEA)
extending from insertion of anal-fin rays to lateral line.
Larger specimens (41.159.1 mm LEA) with clear bands
restricted to region of basal anal-fin pterygiophores, with
diffuse clear spots between proximal portion of anal-fin
pterygiophores and lateral line. Dorsal and ventral margin of
pectoral-fin insertion densely darkened and central portion
unpigmented. Pectoral-fin rays in small specimens (up to
31.1 mm LEA) with dark pigmentation along length of
dorsal and ventral surfaces of rays. Larger specimens (more
than 40 mm LEA) with dark pectoral-fin pigmentation
restricted to medial portions of fin. Anal-fin rays with dark
pigmentation on anterior and posterior portions, occasion-
ally only on posterior portion of fin. Interradial membranes
of anal and pectoral fins hyaline.
Dorsal rami of intermittent branch of anterior lateral line
nerve visible as two black parallel lines on each side of dorsal
portion of body approximately along vertical through
pectoral-fin insertion. Lateral line nerve visible as dark line
extending from above pectoral-fin insertion posteriorly into
approximately basal one-third of caudal filament.
Distribution.Hypopygus benoneae is only known from rain-
forest streams in the lower Rio Anapu in the Floresta
Nacional de Caxiuana, lower Amazon basin, Para, Brazil
(Fig. 2).
Habitat.Specimens of Hypopygus benoneae were collected in
shallow areas along the margins (0.60.9 m deep and 2.5
Fig. 2. Map of the lower Amazon basin showing known distribution of
Hypopygus benoneae. Star indicates type locality.
Peixoto et al.New species of Hypopygus 235
18.8 m of width) of the lower Rio Anapu. The substrate was
composed of submerged root-tangle and rafts with leaf litter
(N. Benone, pers. comm.). Hypopygus benoneae co-occurs
with H. lepturus in the Rio Anapu (see Material Examined).
Remarks.Hypopygus benoneae was collected syntopically
with specimens of H. lepturus. Hypopygus benoneae can be
easily distinguished from the syntopic population of H.
lepturus by the interocular width (26.134.5% vs. 22.4
25.7% HL, respectively) and the number of pectoral-fin rays
(1113 vs. 910, respectively).
Etymology.The species name, benoneae, is in honor of
Naraiana Benone who collected the majority of the
specimens that served as the basis for this description.
DISCUSSION
Formal taxonomic revisions have often proved to provide
the foundation for detecting additional intrageneric diver-
sity and developing a more accurate estimate of species-level
diversity and the distributional ranges of the described
species (e.g., Reis, 2004; de Santana and Vari, 2010a, 2010b).
That generality has proved to apply to Hypopygus, with the
analysis of samples of the genus recently collected in the
lower Amazon basin within the context of a recently
published revision (de Santana and Crampton, 2011)
revealing a previously undescribed species, H. benoneae.
Hypopygus benoneae shares the 13 synapomorphies pro-
posed for the genus: the anterior nares located inside the
upper lip (character 1, state 1), the loss of the posterior naris
(character 2, state 1), the loss of the fourth infraorbital bone
(character 13, state 1), the loss of the fifth infraorbital canal
bone (character 14, state 1), the loss of the parietal branch of
the supraorbital canal (character 17, state 1), the loss of the
vomer (character 22, state 1), the supracleithrum indepen-
dent from the posttemporal (character 29, state 1), the
absence of the mesocoracoid bridge (character 30, state 1),
the loss of the anteriormost branchiostegal ray (character 32,
state 1), the anterior portion of the anal-fin pterygiophores
completely covered by scales (character 43, state 2), the
presence of opaque tissue covering the base of the anal fin
(character 45, state 1), the intermittent lateral line (character
46, state 1), and the presence of the postpectoral accessory
electric organ (character 47, state 1). It also shares the eight
synapomorphies proposed by de Santana and Crampton
(2011) for the clade formed by H. hoedemani, H. lepturus, and
H. minissimus: the loss of the fourth mandibular canal bone
(character 8, state 1), the loss of the fifth mandibular canal
bone (character 9, state 1), the absence of the sixth
infraorbital (character 15, state 1), the narrow anterior
dorsomedial fontanel (character 21, state 1), the absence of
the medial ridge on the posterior portion of the dorsal
surface of the basihyal (character 33, state 1), the second
basibranchial ossified (character 34, state 1), the loss of the
ventral preopercular sensory canal (character 38, state 1),
the loss of the dorsal preopercular sensory canal (character
39, state 1), and the low number of anal-fin rays (115134;
character 41, state 1). Additionally, H. benoneae is included
in the clade of H. hoedemani and H. minissimus by the
presence of two of the eight proposed synapomorphies: the
loss of the infraorbital canal aperture (character 16, state 1)
and the partial or total loss of the extrascapular canal
(character 25, state 1). Consequently, we hypothesize that
H. benoneae is the sister group of the clade composed by H.
hoedemani and H. minissimus.
Nine of the ten synapomorphies shared between H.
benoneae and the clade that contains H. hoedemani, H.
lepturus, and H. minissimus represent reductive characters (8,
9, 15, 16, 21, 25, 38, 39, and 41). Thus, based on the
presence of these reductive characters and despite of the lack
of information on the minimum size of maturity, H.
benoneae is classified as a miniature gymnotiform as are
the other members of that clade.
KEY TO THE SPECIES OF HYPOPYGUS
1a. Absence of sixth infraorbital bone; total number of
anal-fin rays, 102135 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 2
1b. Presence of sixth infraorbital bone, as a narrow
tube, positioned vertically, parallel to posterior
border of eye; total number of anal-fin rays, 136
174 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 5
2a. Presence of dark oblique stripes on lateral and
dorsal portions of body; presence of scales on mid-
dorsal region of body _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 3
2b. Absence of dark oblique stripes on lateral and
dorsal portions of body; absence of scales on mid-
dorsal region of body _ _ _ _ _ _ _ _ _ _ _ _ _ _ Hypopygus minissimus
Upper Orinoco, Venezuela; Upper Rio Negro,
Amazon basin, Brazil
3a. Presence of infraorbital canal aperture; presence of
extrascapular canal _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 4
3b. Absence of infraorbital canal aperture; absence of
extrascapular canal _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ Hypopygus hoedemani
widespread in Rio Negro and Rio Preto da Eva,
Amazon basin, Brazil
4a. Pectoral-finrays, 1113; interocular width, 26.134.5%
HL _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ Hypopygus benoneae, new species
Rio Anapu in the Floresta Nacional de Caxiuana ,
lower Amazon basin, Para, Brazil
4b. Pectoral-fin rays, 910; interocular width, 18.6
24.4% HL _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ Hypopygus lepturus
wide distribution embracing: most of the Amazon
basin in Bolivia, Brazil, Colombia, Ecuador, and
Peru; the Orinoco basin in Colombia and Venezuela;
major drainages of the Guianas in French Guiana,
Guyana, and Suriname; and some northern tributaries
of the Paraguay basin in Brazil
5a. Presence of scattered chromatophores on anal-fin
rays and membranes _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 6
5b. Absence of scattered chromatophores on anal-fin
rays and membranes _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 7
6a. Snout bulbous; 137145 anal-fin rays; 1011 pectoral-
fin rays _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ Hypopygus neblinae
widespread in R o Orinoco, Venezuela; widespread in
Rio Negro and Rio Preto da Eva, Amazon basin, Brazil
6b. Snout slightly curved; 155174 anal-fin rays; 14
pectoral-fin rays _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ Hypopygus cryptogenes
lower and middle Rio Negro, and Rio Preto da Eva,
Amazon basin, Brazil
7a. Bulbous head in a dorsal view; dorsal rami of
intermittent branch of anterior lateral line nerve
invisible _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ Hypopygus ortegai
known only from localities near Iquitos and Jenaro
Herrera, Loreto, upper Amazon basin, Peru
7b. Triangular head in a dorsal view; dorsal rami of
intermittent branch of anterior lateral line nerve
visible as two black parallel lines _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 8
236 Copeia 2013, No. 2
8a. Seven to eight scales above lateral line_ _ _ _ _ _ __ _ _ _ _ _ __ _ _ _ _ _ _ __ _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ Hypopygus nijsseni
known only from localities near Tefe, Amazonas,
upper Amazon basin, Brazil
8b. Five to six scales above lateral line _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ Hypopygus isbruckeri
known only from near San Fernando de Atabapo,
upper Orinoco basin, Venezuela
MATERIAL EXAMINED
Hypopygus lepturus: Brazil, Para, Floresta Nacional de Cax-
iuana, Rio Anapu, Igarape Curua: MPEG 6210, 1, 46.0 mm
LEA; MPEG 9945, 1, 55.8 mm LEA; MPEG 9965, 1, 55.3 mm
LEA; MPEG 9967, 1, 40.8 mm LEA; MPEG 10008, 2, 38.4
56.2 mm LEA; MPEG 10171, 9, 45.558.6 mm LEA; MPEG
12043, 1, 23.5 mm LEA; MPEG 12044, 1, 29.6 mm LEA;
MPEG 12045, 2, 20.220.4 mm LEA.
ACKNOWLEDGMENTS
We thank L. Smith and M. Rogers (FMNH), L. Rapp Py-
Daniel, R. de Oliveira, and M. Rocha (INPA), F. Provenzano
(MBUCV), D. Taphorn (MCNG), P. Buckup (MNRJ), M. de
Pinna (MZUSP), and R. Vari (USNM) for the loan of
specimens and assistance during visits to their institutions.
A. Dourado prepared Figure 1, and B. Prudente Figure 2.
LAWP and GMD thank Coordenacao de Aperfeicoamento de
Pessoal de N vel Superior (CAPES) for support. WBW thanks
Conselho Nacional de Pesquisa (CNPq) by the Bolsa de
Produtividade em Pesquisa. Collection of the type material
was aided by the Programa de Pesquisa em Biodiversidade
(PPBIO) of the Ministerio da Ciencia e Tecnologia (MCT),
Brazil. This paper benefited from suggestions by R. Vari.
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