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2001 by the American Society of Ichthyologists and Herpetologists
Copeia, 2001(4), pp. 1058–1063
New Species of Moenkhausia (Characiformes: Characidae) from
the Rio Negro of Brazil
F
LA
´
VIO
C. T. L
IMA AND
M
O
ˆ
NICA
T
OLEDO
-P
IZA
A new Moenkhausia species is described from the middle rio Negro of Brazil. This
species shares with Moenkhausia oligolepis, Moenkhausia sanctaefilomenae, and Moenk-
hausia pyrophthalma a reticulate body coloration and a red pigmentation on the
dorsal margin of the eye. It can be distinguished from those species by the presence
of a distinctive stripe on the caudal peduncle, a uniform anterior humeral blotch,
and meristic and morphometric data. Priocharax ariel, Serrabrycon magoi, Microster-
narchus fimbripinnus, and Acestridium martini are recorded for the first time from the
middle rio Negro in Brazil.
Uma espe´cie nova de Moenkhausia e´ descrita do me´dio rio Negro no Brasil. Esta
espe´cie compartilha com Moenkhausia oligolepis, Moenkhausia sanctaefilomenae e
Moenkhausia pyrophthalma um padra˜o de colorido reticulado e a pigmentac¸a˜o ver-
melha na margem dorsal do olho. Distingui-se destas espe´cies pela presenc¸a de uma
faixa perspı´cua no pedu´nculo caudal, uma mancha umeral anterior uniforme e por
dados merı´sticos e morfome´tricos. Priocharax ariel, Serrabrycon magoi, Microsternar-
chus fimbripinnus e Acestridium martini sa˜o registrados pela primeira vez no me´dio
rio Negro no Brasil.
A
S presently defined the characid genus
Moenkhausia Eigenmann consists of ap-
proximately 50 species. The monophyly of
Moenkhausia as well as the species level taxono-
my are yet to be critically examined. Species cur-
rently assigned to Moenkhausia are widespread
throughout rivers of cis-andean South America.
We herein describe a new, distinctive characid
that we include in the genus Moenkhausia. The
new species was collected in the middle rio Ne-
gro in 1972 by the Expedic¸a˜o Permanente da
Amazoˆnia (EPA), a project conducted by
MZUSP that made extensive collecting expedi-
tions in the Amazon region from 1967–1975.
The 1972 EPA expedition focused on an inten-
sive sampling of the small igarape´s tributaries to
the rio Negro and the surroundings of Santa
Isabel do Rio Negro (formerly Tapurucuara).
That effort yielded excellent collections of small
fishes, part of which were reported on by pre-
vious authors (Weitzman and Balph, 1979;
Weitzman and Fink, 1983; de Pinna, 1989). Two
additional specimens were collected by Tuyuka
Indians in the rio Tiquie´, a tributary of the rio
Uaupe´s, during an expedition in which the first
author took part. In addition to the description
of the new species, we also report on new re-
cords for two characiforms, one gymnotiform,
and one siluriform for the middle rio Negro.
M
ATERIALS AND
M
ETHODS
Counts and measurements were taken ac-
cording to Fink and Weitzman (1974:1–2). In
the description, the frequency of each count is
provided in parentheses after the respective
count, with the count of the holotype indicated
by an asterisk. The total number of specimens
counted was 14, except when indicated. Verte-
bral counts, pterygiophore insertions relative to
neural and hemal spines, and posterior dentary
teeth were recorded from a single cleared-and-
stained (C&S) specimen (MZUSP 62615, 44.2
mm SL). Vertebrae incorporated into the We-
berian apparatus were counted as four elements
and the fused PU1
1
U1 was considered a single
element.
Institutional abbreviations follow Leviton et
al. (1985). Institutional catalog numbers are fol-
lowed by the number of specimens in a lot,
range of standard lengths, and locality infor-
mation.
Moenkhausia diktyota n. sp.
Figure 1
Holotype.—MZUSP 62614 (52.4 mm SL), Brazil,
Amazonas, rio Negro, igarape´atSa˜o Joa˜o, near
Santa Isabel do Rio Negro, 0
8
24
9
S; 65
8
02
9
W;
EPA, 23 October 1972.
Paratypes.—MZUSP 62615 (9, 1 C&S, 28.4–52.3
mm SL); INPA 16200 (1, 51.2 mm SL); USNM
363175 (1, 51.5 mm SL); same data as holotype.
MZUSP 64107 (2, 35.5–45.1 mm SL), Brazil,
Amazonas, igarape´ Yoariwasotoamaku´ya, tribu-
tary of rio Tiquie´, comunidade de Cachoeira
1059LIMA AND TOLEDO-PIZA—NEW MOENKHAUSIA FROM RIO NEGRO
Fig. 1. Moenkhausia diktyota, holotype, MZUSP 62614, 52.4 mm SL, Brazil, Amazonas, rio Negro, igarape´at
Sa˜o Joa˜o, near Santa Isabel do Rio Negro.
Comprida, 0
8
15
9
44.0
0
N, 70
8
01
9
05.2
0
W; Tuyuka
Indians, 23 October 2000.
Diagnosis.—Among the species traditionally as-
signed to the Tetragonopterinae (Ge´ry, 1977),
M. diktyota is most similar to M. oligolepis (Gu¨n-
ther), Moenkhausia sanctaefilomenae (Steindach-
ner), and Moenkhausia pyrophthalma Costa, with
which it shares a reticulate body color pattern
formed by dark pigmentation along the poste-
rior portion of the scales. M. diktyota is distin-
guished from these three species by having a
horizontally elongate black stripe extending
from a vertical through the last anal fin ray to
tips of middle caudal fin rays (Fig. 1). In M.
oligolepis, M. sanctaefilomenae, and M. pyrophthal-
ma, the caudal peduncle has a large, vertically
elongate, irregular rectangular blotch that ex-
tends between its dorsal and ventral margins
and horizontally from the end of the caudal pe-
duncle to the caudal fin base. In M. diktyota, the
caudal peduncle stripe is not preceded by a
light area such as occurs in M. oligolepis, M. sanc-
taefilomenae, and M. pyrophthalma. The anterior
humeral blotch in all four species is vertically
elongate. However, in M. diktyota, the anterior
humeral blotch is uniformly pigmented along
its length, whereas in M. oligolepis, M. sanctaefi-
lomenae, and M. pyrophthalma, it is more intense-
ly pigmented dorsally, becoming fainter ventral-
ly.
Moenkhausia diktyota is further distinguished
from M. sanctaefilomenae and M. oligolepis by its
shallower body (31.0–36.3 vs 37.9–44.0, n
5
14
and 42.2–48.4, n
5
25, respectively) and by its
incomplete lateral line (vs usually complete, see
Discussion). Although some overlap occurs, M.
diktyota usually has a deeper body (31.0–36.3 vs
27.5–31.6), more scales along the lateral series
(31–34 vs 27–31) and more branched anal fin
rays (20–22 vs 17–19) than M. pyrophthalma.
Description.—Morphometric data for the holo-
type and paratypes are presented in Table 1.
Body compressed, moderately elongate, greatest
body depth slightly anterior to dorsal fin origin.
Dorsal profile of head convex from upper lip to
vertical through anterior nostril; straight to
slightly concave from latter point to tip of su-
praoccipital spine. Predorsal body profile con-
vex, posteroventrally inclined along dorsal fin
base, straight to slightly convex from that point
to vertical through last anal fin ray; slightly con-
cave between latter point and origin of dorsal-
most procurrent caudal fin ray. Ventral profile
of head and body gently convex to anal fin or-
igin. Body profile along anal fin base convex
and posterodorsally slanted. Ventral profile of
caudal peduncle straight to concave.
Jaws equal, mouth terminal, maxilla reaching
vertical slightly anterior to anterior limit of or-
bit. Snout length shorter than eye diameter. Eye
large, with very small adipose eyelid along an-
terior margin of orbit.
Premaxillary teeth in two rows (Fig. 2). Outer
row with 3 (4), 4* (8) or 5 (1) tricuspid teeth.
Inner row with 5* (13) or 6 (1) teeth; all teeth
pentacuspid, except for tricuspid posteriormost
tooth. Maxillary teeth 2 (8) or 3*(4), tricuspid.
Dentary with four large pentacuspid teeth fol-
lowed by eight smaller teeth, the anterior tooth
tricuspid and all others conic.
Scales cycloid, moderately large. Lateral line
incomplete, with 8 (1), 9 (1), 10* (3), 11 (8) or
1060 COPEIA, 2001, NO. 4
T
ABLE
1. M
ORPHOMETRIC
D
ATA OF
Moenkhausia diktyota. n
5
14.
Character Holotype Range Mean
Standard length (mm) 52.4 28.4–52.4
Percentages of standard length
Body depth at dorsal fin origin
Snout to dorsal fin origin
Snout to pectoral fin origin
Snout to pelvic fin origin
Snout to anal fin origin
34.0
49.8
26.9
46.9
66.6
31.0–36.6
49.8–53.9
26.8–29.2
46.6–50.0
65.4–68.4
34.0
52.4
27.7
47.4
66.6
Caudal peduncle length
Caudal peduncle depth
12.6
11.5
11.9–18.3
10.4–11.9
12.2
12.1
Pectoral fin length
Pelvic fin length
20.2
16.2
20.2–23.6
15.1–18.4
22.0
16.8
Dorsal fin length
Anal fin base
Head length
25.4
28.4
26.0
25.4–29.6
26.0–28.5
25.2–29.2
27.2
27.6
26.6
Percentages of head length
Snout length
Orbital diameter
Interorbital width
Upper jaw length
24.3
39.0
33.8
49.3
20.0–24.3
39.0–43.4
31.3–37.7
41.7–49.3
24.5
38.7
34.6
45.7
Fig. 2. Jaws and teeth of Moenkhausia diktyota,
paratype, MZUSP 62615, 44.2 mm SL. Premaxilla,
maxilla, and lower jaw. Lateral view, anterior to right.
Scale bar: 1 mm.
12 (1) perforated scales. Lateral series scales in-
cluding perforated scales 31 (1), 33* (10) or 34
(3). Scales of two longitudinal rows above lat-
eral line series larger; scales of other rows of
scales decreasing in size dorsally and ventrally.
Scales in transverse series between dorsal fin or-
igin and lateral line 5, not including half scale
of predorsal series situated just anterior to first
dorsal fin ray in all specimens. Four scales in
transverse series between lateral line and pelvic
fin origin in all specimens. Predorsal scales
9*(4), 10 (9) or 11 (1). Twelve scale rows
around caudal peduncle in all specimens. Cau-
dal fin scaled, with small scales extending along
approximately two-thirds length of outermost
caudal fin rays. Scales on middle caudal fin rays
restricted to fin base.
Dorsal fin rays ii,9. Dorsal fin origin at mid-
body. Last dorsal fin ray at vertical through anal-
fin origin. First dorsal fin pterygiophore insert-
ing behind neural spine of ninth vertebra. Ad-
ipose fin present. Anal fin rays iv, 20 (7), 21*(3)
or 22 (1). First anal fin pterygiophore inserting
anterior to hemal spine of 16th vertebra. Anal
fin margin concave, with five anteriormost rays
more elongate, rays gradually decreasing in
length posteriorly. Pectoral fin rays i,10 (3), 11
(8) or 12* (3). Tip of pectoral fin reaching ver-
tical through pelvic fin origin. Pelvic fin rays i,7
(i, 6 in one specimen). Caudal fin forked, with
upper and lower lobes similar in size. Epibran-
chial seven; ceratobranchial 8, and hypobran-
chial 2 gill rakers. Thirty-three vertebrae.
Color in alcohol.—Examined specimens lack gua-
nine on body and head, except opercle which
is silvery. Ground coloration tan, darker dorsal-
ly, becoming gradually paler ventrally. Dorsal
portion of head including snout along with tip
of lower jaw with dark pigmentation. Ventral
portion of head and infraorbital region pale.
1061LIMA AND TOLEDO-PIZA—NEW MOENKHAUSIA FROM RIO NEGRO
Upper half of opercle dark forming inconspic-
uous, vertically elongate blotch. Scales on body
except ventral region anterior to pelvic fins and
dorsal and ventral portions of caudal peduncle
bordered by dark pigment and forming con-
spicuous reticulate pattern. Humeral region
with two vertically elongate dark blotches, an-
terior blotch more evident and bordered ante-
riorly and posteriorly by regions of paler color-
ation. Posterior humeral blotch much less con-
spicuous. Caudal peduncle with elongate black
stripe extending from vertical through last anal
fin ray to tips of middle caudal fin rays. Stripe
with rounded anterior margin, wider on caudal
peduncle than on caudal fin rays. All other fins
hyaline. Anterior portion of upper and lower
caudal fin lobes yellowish on region covered by
scales. Smaller specimen (28.4 mm SL) with
overall lighter ground body coloration and with
humeral blotches and blotch on the upper half
of opercle more conspicuous.
Color in life.—Notes on the coloration of two
freshly preserved paratypes (MZUSP 64107)
were taken in the field. Top of head and dor-
sum brownish; sides of head and body copper-
greenish; dorsal portion of the eye and a small
blotch on the upper portion of caudal peduncle
red; caudal peduncle stripe black; pectoral, pel-
vic, caudal, adipose, and anal fins yellow-ochra-
ceous, the latter with some red pigmentation;
and dorsal fin reddish.
Sexual dimorphism.—Anal fin of mature males
with hooks on posterior margin of posterior
branch of the four anterior branched rays. Two
hooks per ray segment (one on each side). Anal
fin shape similar in specimens with and without
hooks.
Geographic distribution.—The species is recorded
from Sa˜o Joa˜o, near Santa Isabel do Rio Negro
(formerly Tapurucuara), in the middle rio Ne-
gro, and from a igarape´ tributary of the upper
rio Tiquie´, a tributary of the rio Uaupe´s, near
the Brazilian/Colombian border. Both localities
lie in the rio Negro basin in Amazonas, Brazil.
Popular names.—The new species is called ‘‘tu-
niupe’’ by the Tukano Indians of the upper rio
Tiquie´, and ‘‘tumuape´,’’ by the Tuyuka Indians
from the same region. Usage of both names is
apparently restricted to the new species, be-
cause other names are employed by the Indians
to identify several other small characids that oc-
cur in the upper rio Tiquie´.
Etymology.—The name diktyota, from the Greek
diktyotos, reticulated, in allusion to the color pat-
tern of the new species. An adjective.
D
ISCUSSION
The species of Moenkhausia as defined by Ei-
genmann (1917:65) are characterized by the
possession of a completely pored lateral line
and a scaled caudal fin. Although M. diktyota has
scales on the caudal fin, its lateral line is incom-
pletely pored. The combination of those two
features would usually result in the assignment
of the new species with the tetragonopterine ge-
nus Hemigrammus Gill. Our decision to place
this species in Moenkhausia rather than Hemi-
grammus is based mainly on its resemblance in
overall body form with some species currently
included in Moenkhausia. Species assigned to
Moenkhausia show a wide range of variation in
features such as body shape and coloration,
which raises the question about the monophy-
letic condition of the genus as currently de-
fined. Therefore, our comparisons were restrict-
ed only to those Moenkhausia species sharing a
reticulate color pattern and red eye pigmenta-
tion.
Within Moenkhausia, M. oligolepis, M. sanctae-
filomenae, and M. pyrophthalma share with the
new species two distinctive pigmentation pat-
tern features: the presence of dark pigmenta-
tion along the posterior margin of the body
scales resulting in a characteristic reticulate pat-
tern, and the occurrence of conspicuous red
pigmentation on the dorsal portion of the eye.
These characters, however, are not restricted to
these four species among Tetragonopterinae.
Red pigmentation on the eye also occurs in, for
example, Astyanax leopoldi Ge´ry et al. (Planquet-
te et al., 1996:251) and Hemigrammus ocellifer
(Steindachner) (Planquette et al., 1996:279).
The reticulate color pattern of the body is also
present in Hyphessobrycon reticulatus Ellis [and in
its presumably senior synonym, Hyphessobrycon
boulengeri (Eigenmann); see Malabarba, 1989:
135]. In addition to the absence of red eye pig-
mentation, the latter species lacks scales on the
caudal fin and has a longitudinal body stripe
that distinguishes it from the Moenkhausia spe-
cies mentioned above.
The reticulate color pattern on the body and
the red pigmentation on the dorsal portion of
the eye were previously proposed by Costa
(1994:28) as indicative of a possible relationship
between M. oligolepis, M. sanctaefilomenae, and M.
pyrophthalma. A third character proposed by
Costa (1994) for this group of species is the
lightly pigmented area followed by a dark
1062 COPEIA, 2001, NO. 4
blotch on the caudal peduncle. This feature is
also present in M. cotinho Eigenmann, which,
however, lacks the distinctive reticulate body
pattern of the other four species. However, the
value of these characters as potential synapo-
morphies for these species can only be evaluat-
ed after a detailed phylogenetic analysis includ-
ing additional characters and taxa.
The fact that M. diktyota has an incomplete
lateral line does not preclude its probable re-
lationship with M. pyrophthalma, M. oligolepis, or
M. sanctaefilomenae. Moenkhausia pyrophthalma
also has an incomplete lateral line (Costa, 1994)
as do populations of M. sanctaefilomenae (Eigen-
mann, 1917:83). As with the characters dis-
cussed above, the incomplete lateral line also
needs to be evaluated in a phylogenetic frame-
work.
Most Astyanax species [e.g., Astyanax fasciatus
(Cuvier), Astyanax bimaculatus (Linnaeus), and
Astyanax scabripinnis ( Jenyns)] have a caudal pe-
duncle stripe that is similar to that of M. diktyota.
However, in those Astyanax species the caudal
peduncle stripe is continuous with a silvery lon-
gitudinal band along the midbody. This longi-
tudinal band is dark in specimens that have not
retained guanine on the body, probably as a
consequence of a relatively longer fixation pe-
riod in formalin. Moenkhausia diktyota does not
possess any longitudinal band along the mid-
body.
The presence of a caudal peduncle stripe and
lack of longitudinal band along the midbody is
also found in Astyanax leopoldi (Ge´ry et al. 1988:
9; Planquette et al., 1996:251). However, A. leo-
poldi can be easily distinguished from M. diktyota
by having a complete lateral line, a caudal fin
not scaled, and the absence of a reticulate color
pattern on the body. In addition, A. leopoldi has
a distinctive chevron-like pigmentation pattern
along the longitudinal line of the body, a fea-
ture absent in M. diktyota.
During our sorting of the 1972 EPA rio Negro
collection, we discovered Priocharax ariel Weitz-
man and Vari (1987; Characidae), Microsternar-
chus fimbripinnus (Mago-Leccia, 1994; Hypopo-
midae), and Acestridium martini Retzer el al.
(1999; Loricariidae), species which were previ-
ously recorded only from localities in the upper
Rio Negro of Venezuela, in the Rı´o Casiquiare,
or upper Rı´o Orinoco. These records represent
an extension of the distribution range of these
species to the middle rio Negro of Brazil.
The lepidophagous characid fish Serrabrycon
magoi Vari (1986) was originally described from
the Rı´o Casiquiare and upper rio Negro. Goul-
ding et al. (1988:125) reported this species from
an unspecified locality in the rio Negro of Bra-
zil. We found specimens of S. magoi in the 1972
EPA collection.
Locality information of the new records fol-
low: Serrabrycon magoi: MZUSP 58697, 13, Brazil,
Amazonas, lagoon on rio Aiuana˜, tributary of
rio Negro; MZUSP 56534, 35, Brazil, Amazonas,
igarape´atSa˜o Joa˜o, near Santa Isabel do Rio
Negro, 0
8
24
9
S, 65
8
02
9
W. Priocharax ariel: MZUSP
55097, 6, Brazil, Amazonas, lagoon at margin of
rio Negro, Paricatuba, 0
8
31
9
S, 65
8
01
9
W; MZUSP
55099, 8, Brazil, Amazonas, igarape´atSa˜o Joa˜o,
near Santa Isabel do Rio Negro, 0
8
24
9
S,
65
8
02
9
W; MZUSP 62230, 4, Brazil, Amazonas, la-
goon in island on rio Negro, Paricatuba, 0
8
31
9
S,
65
8
01
9
W. Microsternarchus fimbripinnus: MZUSP
56536, 3, Brazil, Amazonas, igarape´atSa˜o Joa˜o,
near Santa Isabel do Rio Negro, 0
8
24
9
S,
65
8
02
9
W. Acestridium martini: MZUSP 61945, 4,
Brazil, Amazonas, rio Aiuana˜, near Santa Isabel
do Rio Negro.
Comparative materials are as follows: Moenk-
hausia pyrophthalma: MZUSP 45290, 7 paratypes,
19.8–29.8 mm SL; MZUSP 52246, 7, 20.2–30.6
mm SL. Moenkhausia oligolepis: MZUSP 38264,
21, 31.3–61.2 mm SL; MZUSP 52117, 15, 41.4–
55.5 mm SL; MZUSP 18193, 55, 20.2–73.2 mm
SL. Moenkhausia sanctaefilomenae: MZUSP 59805,
11, 36.6–56.4 mm SL; MZUSP 61160, 14, 17.0–
43.7 mm SL; MZUSP 58269, 25, 22.4–59.6 mm
SL. Moenkhausia cotinho: MZUSP 59219, 58,
28.6–46.6 mm SL. Hyphessobrycon reticulatus:
MZUSP 35241, 30, 37.1–54.4 mm SL.
A
CKNOWLEDGMENTS
We are grateful to R. C. Benine, R. M. C. Cas-
tro, C. R. Moreira, and S. H. Weitzman for crit-
ically reading the manuscript. H. Britski kindly
made available the slide of a freshly collected
specimen of M. diktyota. Specimens from the up-
per rio Tiquie´ were collected during an expe-
dition of the project ‘‘Peixes e pesca no alto
Tiquie´,’’ a joint collaboration among MZUSP,
Instituto Socioambiental, Federac¸a˜o das Orga-
nizac¸o˜es Indı´genas do Rio Negro (FOIRN), and
Associac¸a˜o das Tribos Indı´genas do alto rio Ti-
quie´ (ATRIART). We wish to thank A. Cabalzar,
M. Lopes, C.A. Ricardo, and the staff of FOIRN
and ATRIART for all the assistance and help
during this expedition. Thanks particularly to
the Tuyuka Indians from Cachoeira Comprida,
for their hospitality and generosity. FCTL re-
ceived financial support from FAPESP (grant
99/02402–0).
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———,
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(FCTL) M
USEU DE
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OOLOGIA DA
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NIVERSIDADE
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O
P
AULO
,C
AIXA
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OSTAL
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E
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OOLOGIA
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. E-mail: (FCTL)
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Send reprint requests to FCTL. Submitted: 23
Oct. 2000. Accepted: 30 March 2001. Section
editor: S. A. Schaefer.
