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Three new species of Moenkhausia (Teleostei, Characiformes, Characidae)

with the deﬁnition of the Moenkhausia jamesi species complex
Article in Zootaxa · July 2015

DOI: 10.11646/zootaxa.3986.4.1
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Zootaxa 3986 (4): 401–420 ISSN 1175-5326 (print edition)
www.mapress.com /zootaxa /
Copyright © 2015 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3986.4.1
http://zoobank.org/urn:lsid:zoobank.org:pub:5D6CD3DC-46B9-47F0-B518-AE29234F96F4
Description of three new species of Moenkhausia (Teleostei, Characiformes,

Characidae) with the definition of the Moenkhausia jamesi species complex
MARINA G. PETROLLI & RICARDO C. BENINE

Laboratório de Ictiologia, Departamento de Zoologia, IBB-UNESP Campus de Botucatu, Botucatu, SP, 18618-970, Brazil.
E-mail: magimenezz@gmail.com; rbenine@ibb.unesp.br
Abstract
From the examination of extensive comparative material currently identified as M. jamesi we verified that there are, at
least, three new species under this name. These, along with M. jamesi and M. justae, form what we herein called the M.
jamesi species complex, by sharing the following group of characters: a short maxilla, with its distal margin not exceeding
anterior third of the second infraorbital; first through third teeth of the inner row of premaxilla and first and second dentary
teeth with cusps arranged in a pronounced arch, humeral spot positioned between the fourth and seventh scales of the lat-
eral line and extending up to four scale rows above the lateral line and one scale row below the lateral line, and a vertically
oval to round spot at the base of the caudal fin rays. Moenkhausia ischyognatha sp. n., from Rio Xingu basin, differs from
the other species of the complex by its lower head depth. Moenkhausia alesis sp. n., from the river system Tocantins-Ara-
guaia, differs from M. jamesi, M. ischyognatha, and M. sthenosthoma by the number of scale rows above the lateral line.
Moenkhausia sthenosthoma sp. n., from the Rio Madeira basin, differs from M. jamesi by the number of scale rows be-
tween the lateral line and the midventral scale series. Moenkhausia justae can be diagnosed from the other species of the
complex by having a tri to pentacuspidate tooth on the maxilla.
Key words: Moenkhausia justae, taxonomic revision, Systematics, Neotropical region
Resumo
A partir do exame de extenso material correntemente identificado como Moenkhausia jamesi, verificamos que há pelo
menos três espécies novas sob este nome. Estas, juntas com M. jamesi e M. justae, formam o que aqui denominamos de
complexo de espécies M. jamesi por compartilharem do seguinte conjunto de caracteres: maxilar curto, sua margem distal
não ultrapassa o terço anterior do segundo infraorbital; primeiro ao terceiro dente da fileira interna do pré-maxilar e
primeiro e segundo dente do dentário com suas cúspides arranjadas em um arco pronunciado, mancha umeral localizada
entre a quarta e sétima escamas da série da linha lateral, e estendendo-se até quatro fileiras de escamas acima da linha
lateral e uma fileira de escamas abaixo da linha lateral, e uma mancha verticalmente ovalada a arredonda na base dos raios
medianos da nadadeira caudal. Moenkhausia ischyognatha sp. n., da bacia do Rio Xingu, difere das demais espécies do
complexo por sua menor altura da cabeça. Moenkhausia alesis sp. n., do sistema de rios Tocantins-Araguaia, difere de M.
jamesi, M. ischyognatha e M. sthenosthoma pelo maior número de escamas acima da linha lateral. Moenkhausia sthenos-
thoma sp. n., da bacia do Rio Madeira, difere de M. jamesi pelo maior número de escamas entre a linha lateral e a série
de escamas mediana ventral. Moenkhausia justae pode ser diagnosticada das outras espécies do complexo por ter um dente
tri a pentacuspidado no maxilar.
Palavras-chave: Moenkhausia justae, revisão taxonômica, Sistemática, região Neotropical
Introduction
The genus Moenkhausia currently consists of 76 valid species distributed in the main South-American
hydrographic basins, and with its greatest diversity in the Amazon basin (Lima et al. 2003; Eschmeyer 2015). The
genus was proposed by Eigenmann (1903) based on Tetragonopterus xinguensis and originally described as a
Accepted by M.R. de Carvalho: 18 Jun. 2015; published: 20 Jul. 2015 401

“species similar to Markiana, but with a naked anal fin (without the sheath of small scales covering the base of the
anal fin, as observed in Markiana) and caudal fin with scales”. Eigenmann (1917) presented a more detailed
definition through an identification key that characterized the genus by the presence of small scales covering the
base of the caudal fin; premaxilla with two rows of teeth with the inner row with five multicuspidade teeth; and
lateral line generally complete and without a sharp bend such as present in the genus Tetragonopterus.
Most species of Moenkhausia appear to be widespread in the river basins in which they occur (Lima et al.
2003). Benine et al. (2009), however, showed that M. oligolepis, taken to be widespread in the Amazon and
Paraguay river basins, may represent a complex of species with, at least, four different lineages. In fact, DNA
barcoding has evidenced cryptic species in different groups of neotropical fishes (e.g. Bellafronte et al. 2013;
Pereira et al. 2013; Paz et al. 2014). The employment of a combination of different techniques has shown that
different allopatric/sympatric populations of even morphologically very similar species, not diagnosable by
traditional means, as is the case of Astyanax scabripinnis, represent, actually, cryptic species (Castro et al. 2014a,
b). Purely morphologically based studies have evidenced sibling species (described as subspecies) in groups of
small characids since the second half of the nineteen century, largely in the genus Astyanax (see Eigenmann 1921).
For the genus Moenkhausia, Eigenmann (1908; 1910; 1917) described several subspecies of Moenkhausia lepidura
due to the characteristic black colored upper lobe of caudal fin, which was latter recognized by Géry (1977; 1992)
as the Moenkhausia lepidura group.
From the examination of extensive comparative material currently identified as M. jamesi, we verified that
there are, at least, three very similar undescribed species erroneously recognized under this name. We herein
describe these species and propose a definition for the Moenkhausia jamesi species complex.
Material and methods
Osteological analysis was carried out on cleared and counterstained (c&s) specimens prepared according to Taylor
& van Dyke (1985) and dissected following Weitzman (1974). Osteological terminology follows Weitzman (1962)
and Zanata & Vari (2005). Counts and measurements follow Benine et al. (2015), except for the number of scale
rows between lateral line and the midventral scale series, which were counted at the vertical just ahead of the
pelvic-fin insertion, and head depth, which was measured at the vertical through the tip of supraoccipital spine.
Measurements were taken point to point, on the left side of specimens whenever possible, with a Mitutoyo digital
caliper (0.01 mm).
In the diagnosis and descriptions, principal dentary teeth are the anteriormost similar teeth that decrease
gradually followed by a series of distinctly smaller teeth. Gill rakers were counted from the lower and upper limbs
of the first branchial arch. Lower limb rakers include those on the ceratobranchial and hypobranchial. Upper limb
rakers include those of the epibranchial bone and the raker at the junction between the epibranchial and
ceratobranchial. Unbranched fin rays are indicated by lower case roman numerals, and branched fin rays by arabic
numerals (e.g. iv,38). Counts are followed by their frequency in parentheses. Counts of supraneurals and total
vertebrae were obtained from cleared and stained (c&s) specimens, with the four vertebrae of the Weberian
apparatus and the fused PU1+U1 considered as single bones. Asterisks indicate values for the holotype. Institutions
abbreviations are as listed by Sabaj-Pérez (2014). The examined comparative material is listed separately in the
Comparative Material section.
Results
The Moenkhausia jamesi species complex. The analysis of the type and comparative material showed that a group
of species of Moenkhausia sensu Eigenmann (1917), very similar phenotypically, formed by M. jamesi Eigenmann,
M. justae Eigenmann, M. ischyognatha sp. n., M. sthenosthoma sp. n., and M. alesis sp. n., can be diagnosed from
all congeners by the combination of the presence of a short maxilla, with its distal margin not exceeding anterior
third of second infraorbital (Fig. 1a–b) (vs. maxilla exceeding anterior third of orbit); first (symphyseal) and second
or third teeth of the inner row of premaxilla, and first (symphyseal) and second dentary teeth with cusps arranged in
a pronounced arch when examined in ventral view (Fig. 2a–b), in which the most lateral cusps are positioned in
402 · Zootaxa 3986 (4) © 2015 Magnolia Press PETROLLI & BENINE
parallel or even closer to each other than to the central cusp, conferring a distinctive C-shape to these teeth (vs.
slightly arched, with most lateral cusps never positioned in parallel, and closer to the central cusp than to each
other); humeral spot positioned between the fourth and sixth or seventh scales in the lateral line and extending up to
four scale rows above the lateral line and one scale row below the lateral line; and a vertically oval to round spot at
the base of the caudal-fin rays (see variation of these feature in the Color in alcohol section for each species). This
combination of features is not observed in other known species of Moenkhausia, including its type species,
Moenkhausia xinguensis, except for isolated diagnostic features that are present in the closely related, according to
Mariguela et al. (2013), M. dichroura, M. intermedia, M. costae (tooth morphology), and M. lata (tooth
morphology and humeral blotch color pattern) (see Discussion). Thus, we consider that these very similar species
form a complex within Moenkhausia that is herein defined as the Moenkhausia jamesi species complex, whose
components are described below.
FIGURE 1. Computed tomography images of two syntypes of Moenkhausia jamesi, showing details of the maxilla: (a) MCZ
20742, 55.7mm SL; (b) MCZ 20816, 53.1mm SL.
FIGURE 2. Moenkhausia ischyognatha, LBP 15948, 45.7 mm SL. (a) Left premaxilla (ventral view). (b) Left dentary (medial
view). Scale bars: 0.5 mm.
THREE NEW SPECIES OF MOENKHAUSIA Zootaxa 3986 (4) © 2015 Magnolia Press · 403
Moenkhausia ischyognatha, new species
(Figs. 2a–b, 3a–b, 9c, Table 1)
Holotype. MZUSP 117140, 45.5 mm SL, Mato Grosso, Gaúcha do Norte; Rio Curisevo, afluente rio Xingu, Porto
Vitório, next ribeirão Kevuaieli, 13°02’05”S 53°25’19” W; C. Moreira, I. Landin, A. Datovo; 19 Oct 2004.
Paratypes. MZUSP 91182, 1, 47.2 mm SL, same data as holotype. LBP 15948, 37 (2 c&s), 39.8–56.9 mm SL,
Mato Grosso, Canamara, Rio Culuene, 13°29’42”S, 53°04’58”W; C. Oliveira, M. Taylor, G.H.C. Silva, J.H.M.
Martinez, 02 Aug 2012.
Diagnosis. Moenkhausia ischyognatha is readly distinguished from M. jamesi, M. justae, M. alesis, and M.
sthenosthoma by its lower head depth (29.3–32.6% in SL vs. 34.0–43.2% in SL in the last four species).
Additionally, it differs from M. jamesi and M. alesis by its lower body depth (37.0–43.0% in SL vs 44.3–52.3% in
SL in M. jamesi, and 43.5–54.6% in SL in M. alesis). It is also distinguished from M. alesis by the number of scale
rows between lateral line and dorsal-fin origin (seven vs eight to nine in M. alesis) and by the caudal-peduncle
length (8.3–11.2% in SL vs. 3.8–7.5% in SL in M. alesis). It is further distinguished from M. justae by the number
of teeth in the maxilla (edentulous vs one tooth with three to five cusps in M. justae), and by the number of cusps on
the fourth dentary tooth (three to five vs six in M. justae) (Fig. 9b, c). Moenkhausia ischyognatha is further
distinguished from M. jamesi by the number of principal dentary teeth (four vs five in M. jamesi) (Figs. 9a, c).
FIGURE 3. Moenkhausia ischyognatha. (a) MZUSP 117140, holotype, 45.5 mm SL. (b) LBP 15948, 45.43 mm SL.
-
TABLE 1. Morphometric data for Moenkhausia ischyognatha sp. n., Moenkhausia alesis sp. n., and Moenkhausia sthenosthoma sp. n. N includes holotype. Range includes
holotype and paratypes.
M. ischyognatha (n=38) M. alesis (n= 32) M. sthenosthoma (n=97)
Holotype Range Mean Holotype Range Mean Holotype Range Mean
Standard Length (mm) 45.5 39.8–56.9 52.6 44.7–57.4 56.0 22.3–63.4
Percentage of Standard Length
Greatest depth 38.5 37.0–43.0 40.5 45.8 43.0–54.6 48.1 49.6 38.7–51.3 44.4
Snout to dorsal-fin origin 50.6 48.7–52.8 50.3 51.1 47.5–54.1 51.5 52.3 48.7–54.0 51.2
THREE NEW SPECIES OF MOENKHAUSIA

Snout to pectoral-fin origin 27.2 25.0–29.1 26.8 26.7 24.4–29.9 27.2 27.3 25.5–34.3 29.2
Snout to pelvic-fin origin 49.4 47.2–51.9 49.2 49.8 45.8–53.9 50.0 51.3 46.8–53.1 50.0
Snout to anal-fin origin 67 64.3–69.6 66.5 67.2 61.9–70.9 67.3 67.7 61.5–69.6 65.9
Caudal-peduncle depth 11 10.4–12.0 11.2 11.8 10.6–13.0 11.9 12.1 9.2–12.8 11.4
Caudal-peduncle length 8.5 8.3–11.2 9.6 7.3 3.8–7.5 5.6 7.7 6.3–10.2 8.1
Pectoral-fin length 22.9 17.8–23.5 21.3 23.6 17.6–25.1 21.8 22 16.1–24.7 21.7
Pelvic-fin length 19.9 16.0–19.9 17.8 20.6 16.7–20.6 18.2 19.3 14.7–20.9 18.2
Dorsal-fin length 31.2 25.6–32.4 29.8 34 28.1–35.6 33.0 34.4 27.7–37.5 34.2
Dorsal-fin base 15.8 13.9–16.3 14.9 16.3 13.5–17.6 15.9 15.9 14.4–18.6 16.5
Anal-fin length 17.8 12.1–18.2 15.9 17.2 13.4–19.4 16.9 20.1 14.9–23.1 20.4
Anal-fin base 33 29.9–34.3 31.8 35.4 32.1–39.8 36.2 36.6 33.1–38.4 35.6
Eye to dorsal-fin origin 36.2 34.7–39.1 36.8 38.1 35.8–40.8 38.8 39.6 33.9–51.2 37.1
Dorsal-fin origin to caudal-fin 55.4 53.2–59.0 56.0 56.7 49.2–58.6 58.6 55.4 51.5–58.6 54.9
origin
Head length 26.5 24.4–27.4 25.7 25.2 23.2–27.0 25.0 25 22.9–29.5 26.2
Head depth 30.4 29.3–33.3 31.2 36.9 35.9–43.2 39.1 38.0 33.5–40.1 36.2
Percentage of Head length
Snout length 30.1 25.5–33.5 28.6 29.6 22.6–34.1 29.4 25.1 20.2–31.2 25.6
Maxillary length 34 31.8–37.6 34.4 35.8 33.7–41.3 36.9 37.1 31.1–39.8 35.6
Horizontal orbital diameter 41.9 39.9–47.0 43.2 43.2 36.9–45.3 41.7 44.4 41.7–49.6 46.3
Least interorbital width 36.8 35.3–39.3 37.3 39.9 37.6–48.6 42.1 39.2 33.3–49.2 39.6
Zootaxa 3986 (4) © 2015 Magnolia Press ·

405
FIGURE 4. Partial map of South America indicating the distribuition of Moenkhausia jamesi species complex and M. justae.
Red circle = M. jamesi; Blue circle = M.ischyognatha; Orange circle = M.alesis; Light blue circle = M.sthenosthoma; Yellow
circle = M.justae.
Description. Morphometric data summarized in Table 1. Largest specimen examined 55.9 mm SL. Body
compressed and somewhat elongate. Greatest body depth at dorsal-fin origin. Dorsal profile of head straight to
slightly convex; straight to slightly concave along the supraoccipital spine; slightly convex from tip of
supraoccipital spine to end of dorsal-fin base; straight to slightly convex from end of dorsal fin up to end of adipose
fin; caudal peduncle concave in dorsal and ventral margins; ventral profile slightly convex from tip of snout to end
of anal fin.
Mouth terminal. Maxilla only reaching the vertical through anterior margin, and not trespassing anterior third
of second infraorbital. Premaxillary teeth in two rows. Inner row with five tetracuspidate (symphyseal) or
pentacuspidate (Fig. 2a) teeth with median cusp pronounced, the first two teeth from the symphysis with cusps
arranged in a pronounced arch when examined from a ventral view; outer row with four to five pentacuspidate
teeth; edentulous maxilla. Dentary bearing four pentacuspidate teeth with central cusp longest followed by three to
five distinctly small conical or tricuspidate teeth. First two or three dentary teeth from the symphysis with cusps
arranged in a pronounced arch when examined from a dorsal view.
Dorsal-fin rays ii,9. Pectoral-fin rays i,12. Tip of pectoral fin extending slightly beyond anterior insertion of
pelvic fin. Pelvic-fin rays i,7, tip of adpressed pelvic fin not reaching anal fin. Anal-fin rays iv,26(4), 27(10),
28(15), 29*(7), 30(2); caudal fin forked with i,9,8,i.
Scales cycloid. Lateral line with 35(1), 36*(17), 37(18), 38(1) perforated scales; Scale rows between lateral
line and dorsal-fin origin 7. Scale rows between lateral line and midventral scale series 6(30) to 7*(8).
Circumpeduncular scale rows 14*(2) and 15(36); Scale sheath along anal-fin base 7–15*, in single series, covering
base of anteriormost rays. Small scales covering proximal two-third of caudal-fin lobes.
First gill arch with 12(1), 13(33)*, 14(2) gill rakers on lower limb and 8(1), *9(31), 10(5) on upper limb. Total
vertebrae 31. Supraneurals 4.
Color in alcohol. Overall coloration silvery or yellow tan. Field of few dark chromatophores on upper lip and
maxilla. Infraorbital and opercular series silvery due to the presence of guanine pigmentation. Dark
chromatophores more densely concentrated along entire dorsal midline. Sparsely spread dark chromatophores
dorsal of horizontal skeletogenous septum. Dark line over horizontal skeletogenous septum. Conspicuous silver
midlateral stripe extending from posterior humeral mark to base of median caudal fin-rays. Irregularly shaped,
humeral mark located over fourth to sixth lateral-line scales and extending vertically over four horizontal scale
rows above and up to one horizontal scale row below lateral line. Paired fins and anal fin hyaline. Inconspicuous
round dark spot at the base of the caudal-fin rays formed by few chromatophores. More well-preserved specimens
may present a more conspicuous round caudal spot. Adipose with very few dark chromatophores (Fig. 3a–b).
Distribution. Moenkhausia ischyognatha occurs in the Rio Xingu basin, in Rio Culuene and Rio Curisevo
(Fig. 4).
Etymology. The species name ischyognatha is from Greek ischyo meaning strong, and gnatha meaning jaw, in
reference to the strong musculature associated to the dentary, and robust teeth of the premaxilla and dentary, a
diagnostic feature of the Moenkhausia jamesi species complex.
Moenkhausia alesis, new species

(Figs. 5, 9d, Table 1)
Holotype. MZUSP 117139, 52.6 mm SL, Tocantins, Conceição de Tocantins; rio Palmas at TO 050, near to Chuva
de Manga locality, 12°25’7”S, 47°11’52”W, J.L. Birindelli; F.C. Dagosta; M.V. Loeb; C. Santos; 01 Dec 2012.
FIGURE 5. Moenkhausia alesis, MZUSP 117139, holotype, 52.6 mm SL.
Paratypes. MZUSP 113845, 2, 45.1–55.6 mm SL, same data as holotype. INPA 48444, 6 (1 c&s), 44.7–55.1
mm SL, Caseara, Parque Estadual do Cantão, E.G. Ferreira, J. A. Zuanon & G. M. Santos, May 2000. MZUSP
52354, 10 of 20, 53.9–57.7 mm SL, Goiás; Rio Araguaia, at beach between Bandeirantes and Luis Alves;
13°40’00”S, 50°48’00”W, Coleção Rio Araguaia, Jul/Aug 1997. LBP 18905, 1, 58.4 mm SL, Goiás, Aragarças,
beach of rio Araguaia; 15°53'30.09"S, 52°14'59.70" W, P.C. Vênere, no date. MZUSP 52347, 12, 52.0–57.4 mm
SL, Mato Grosso; Rio Araguaia, Fio Velasco, 12°52’58”S, 50°30’07”W, R. S.A. Matias, Jul 1997.
Diagnosis. Moenkhausia alesis is distinguished from M. jamesi, M. ischyognatha, and M. sthenosthoma by the
number of scale rows between lateral line and dorsal-fin origin (eight to nine vs seven in the last three). Differs
from M. ischyognatha by its greater body depth (43.5–54.6% in SL vs 37.0–43.0% in SL in M. ischyognatha), by
its greater head depth (35.9–43.2% in SL vs 29.3–32.6% in SL in M. ischyognatha), and by its lower caudal
peduncle length (3.8–7.5% in SL vs 8.3–11.2% in SL in M. ischyognatha). Moenkhausia alesis is additionaly
distinguished from M. jamesi by the number of scale rows between lateral line and midventral scale series (seven to
eight vs six in M. jamesi) and by the number of principal dentary teeth (four vs five in M. jamesi) (Fig. 9a, d). It is
distinguished from M. justae by the number of teeth in the maxilla (edentulous vs one tooth with five cusps in M.
justae), and by the number of cusps on the fourth dentary tooth (three to five vs six in M. justae) (Figs. 9b, d).
compressed and deep. Greatest body depth slightly anterior to dorsal-fin origin. Dorsal profile of head slightly
convex; straight to slightly concave along the occipital spine; slightly convex to convex from tip of supraoccipital
spine to end of dorsal-fin base; straight to slightly convex from end of dorsal fin up to end of adipose fin; caudal
peduncle concave in dorsal and ventral margins; ventral profile slightly convex from tip of snout to end of anal fin.
pentacuspidate teeth with median cusp pronounced, the first two teeth from the symphysis with cusps arranged in a
pronounced arch when examined from a ventral view; outer row with four to five pentacuspid teeth; edentulous
maxilla. Dentary bearing four pentacuspidate teeth with central cusp longest followed by three to four distinctly
small conical or tricuspidate teeth. First two or three dentary teeth from the symphysis with cusps arranged in a
pronounced arch when examined from a dorsal view.
Dorsal-fin rays ii,9. Pectoral-fin rays i,11(2), 12*(23), 13(7). Tip of pectoral fin extends slightly beyond
anterior insertion of pelvic fin. Pelvic-fin rays i,7. Tip of adpressed pelvic fin not reaching anal fin. Anal-fin rays
iv,28(5), 29*(11), 30(8), 31(6), 32(2) rays; caudal fin forked with i,9,8,i.
line and dorsal-fin origin 8*(30), 9(2). Scale rows between lateral line and midventral scale series 7*(23) to 8*(9).
Circumpeduncular scale rows 13(1), 14*(22), 15(2), 16(2), 17(1), 18(4). Scale sheath along anal-fin base 7–15*, in
1–3* series, covering base of anteriormost rays. Small scales covering proximal two-third of caudal-fin lobes.
First gill arch with 13(9), 14(9)*,15(1) gill rakers on lower limb and 9(5), 10(11)*, 11(3) on upper limb. Total
Color in alcohol. Overall coloration slightly silvery or pale yellow. Field of few dark chromatophores on
upper lip and maxilla. Infraorbital and opercular series silvery due to the presence of guanine pigmentation. Dark
midlateral stripe extending from posterior margin of opercle to base of median caudal fin-rays. Irregularly shaped,
humeral mark located over fourth to seventh lateral-line scales and extending vertically over four-five horizontal
scale rows above and up to one horizontal scale row below lateral line. Paired fins and anal fin hyaline. Round dark
spot at the base of the caudal-fin rays formed by few chromatophores. Adipose with very few dark chromatophores
(Fig. 5).
Distribution. Moenkhausia alesis occurs in the upper and middle portions of Rio Araguaia and Rio Tocantins
(Fig. 4).
Etymology. The species name alesis is from the Greek meaning grinder, in reference to the robust teeth of the
premaxilla and dentary, a diagnostic feature of the Moenkhausia jamesi species complex.
Moenkhausia sthenosthoma, new species

(Figs. 6a–c, 9e–f, Table 1)
Moenkhausia jamesi: Queiroz et al. (2013): 312–313. Mariguela et al. (2013): 4–5.
Holotype. MCP 48789, 56.0 mm SL, Rondônia, Madeira, Rio Jaci-Paraná between Porto Velho and Jaci-Paraná;
9º15’23”S, 64º23’13”W, R. E. Reis, P. A. Buckup, F. Langeani, V. Bertaco, P. Lehmann., 18 Jul 2004.
Paratypes. All from Brazil. Rondônia. MCP 39421, 20, 40.5–56.0 mm SL, Rio Jaci-Paraná, between Porto
Velho and Jaci-Paraná, rio Madeira, 9º15’23”S, 64º23’13”W, R. E. Reis et al., 18-Jul-2004. MCP 39417, 30, 29.6–
41.4 mm SL, Rio Madeira, Rio Mamoré, in front of Cristo Rei neighborhood, Gujará-Mirim, 10°47’03”S
65°20’58”W, R. E. Reis et al., 25-Jul-2004. ANSP 200003, 10, 32.3–45.6mm SL, same data as MCP 39417. MNRJ
43441, 10, 33.7–46.7, same data as MCP 39417. UFRO 300, 1, 40.3 mm SL, Igarapé Arara, Porto Velho;
10º0’45”S, 65º19’7”W; L. Queiroz; 08-Apr-2004. UFRO 317, 1, 52.7 mm SL, Fortaleza do Abuña, Rio Abuña
waterfall, rio Madeira; 09º47’S, 65º31’W; M.A.L, Lima; 09-Jun-2004. UFRO 3785, 2, 41.5–46.1 mm SL, Jaci-
Paraná, mouth of Rio Jaci-Paraná, rio Madeira; 09º13’01,5”S, 64º22’58,4”W; T. Fernandes; 07-Nov-2009. INPA
48440, 2, 22.3–31.4, Fortaleza do Abuña, Rio Abuña waterfall, Rio Madeira; 09º04’S, 65º31’W; G. Vilara; 07-Apr-
2004. INPA48441, 1, 40.8 mm SL, Guajará-Mirim, rio Pacaás Novos, near to the confluence with rio Mamoré;
10º51’31,6”S, 65º16’39,1”W; A. Ribeiro, 30-Sep-2009. INPA 48442, 3, 44.1–56.6 mm SL, Pimenteiras do Oeste,
Rio Guaporé, mouth of Rio Cautário; 12º12’4989”S, 64º35’09,41”W; D. Hungria; 10-Jan-2012.MZUSP 117143,1,
49.7 mm SL, Jaci-Paraná, Lago Madalena, rio Jaci-Paraná; 9º13’00” S, 64º00’70’’W; L.C.R Melo; 10-Jul-2009.
MZUSP 117144, 1, 35.4 mm SL, mouth of Igarapé Belmont, Rio Madeira; 8º37’56,56”S, 63º50’20,51”W; G.
Vilara; Feb-2006. MZUSP 117145, 2, 57.5–61.4 mm SL, Jaci-Paraná, upstream of rio Jaci- Paraná, left margin
shore; 09º27’16,5”S, 64º24’51,9”W; T. Fernandes; 03-Aug-2011. MZUSP 117142,1, 39.0 mm SL, Guajará-Mirim,
rio Soterio, shore at mouth of Rio Negro; 11º35’51,9”S, 65º 13’ 49,1”W; LIP/UNIR team, 04-Jul-2011. MZUSP
117141, 3, 49.3–54.6 mm SL, Pimenteiras do Oeste, rio Guaporé, mouth of rio Cautário; 12º12’49,89”S,
64º35’09,41”W; D. Hungria; 10-Jan-2012. LBP 19956, 1 (c&s), 46.0 mm SL, Jaci-Paraná, mouth of Igarapé
Karipuna, rio Madeira; 09º11’27,7”S, 64º37’04”W; Talles Fernandes; 04-Sep-2009. LBP 19957, 1, 57.2 mm SL,
Jaci-Paraná, mouth of Rio Jaci-Paraná, rio Madeira; 09º17’29”S, 64º23’56,67”W; G. Vilara; 01-Apr-2006. LBP
19958, 2, 1 (c&s), 61.9–63.4 mm SL, Jaci-Paraná, Três Praias, rio Jaci-Paraná, rio Madeira. 9º27’9.9”S,
64º24’56,7”W; C. Röpke; 11-Dec-2009. LBP 19959, 3, 54.9–63.0 mm SL, Guajará-Mirim, Surpresa, rio Guaporé;
11º53’S; 65º01’W; LIP/UNIR team; 11-Jan-2012. 08-Apr-2004. Mato Grosso. LBP 19960, 3, 52.1–53.4 mm SL,
Vila Bela da Santíssima Trindade, rio Guaporé, ponte para Vila Bela da Santíssima Trindade; 15ºS, 59,95ºW; W. M.
Ohara; 13-Dec-2011.
Diagnosis. Moenkhausia sthenosthoma is readly distinguished from M. jamesi by the number of scale rows
between lateral line and the midventral scale series (seven vs six in M. jamesi). It differs from M. ischyognatha by
its greater head depth (33.9–37.8% in SL vs 29.3–32.6 in SL in M. ischyognatha). It is distinguished from M. alesis
by the number of scale rows between lateral line and dorsal-fin origin (seven vs eight to nine in M. alesis).
Moenkhausia sthenosthoma is distinguished from M. justae by the number of teeth in maxilla (edentulous vs one
tooth with 5 cusps in M. justae), and by the number of cusps on the fourth dentary tooth (three to five vs six in M.
justae) (Fig. 9b, 9e–f).
compressed and deep. Greatest body depth slightly anterior to or at dorsal-fin origin. Dorsal profile of head slightly
pentacuspidate teeth with median cusp pronounced, the first two or three teeth from the symphysis with cusps
arranged in a pronounced arch when examined from a ventral view; outer row with four to five pentacuspid teeth;
edentulous maxilla. Dentary bearing four to five pentacuspidate teeth with central cusp longest followed by three to
four distinctly small conical or tricuspidate teeth. First two or three dentary teeth from the symphysis with cusps
anterior insertion of pelvic fin. Pelvic-fin rays i,7, tip of adpressed pelvic fin not reaching anal fin. Anal-fin rays
iv,28(3), 29(14), 30(27), 31*(34), 32(15), 33(3). Caudal fin forked with i,9,8,i.
Scales cycloid. Lateral line with 34(2), 35(10), 36(43), 37*(36), 38(6) perforated scales; scale rows between
lateral line and dorsal-fin origin 7*. Scale rows between lateral line and midventral scale series 7*.
Circumpeduncular scale rows 13(4), 14(61), 15*(28), 16(3). Scale sheath along anal-fin base 7–15*, in 1–3* series,
covering base of anteriormost rays. Small scales covering proximal two-third of caudal-fin lobes.
First gill arch with 11(3), 12(35), 13*(45), 14(2) gill rakers on lower limb and 8*(25), 9(57), 10(3) on upper
limb. Total vertebrae 31–32. Supraneurals 4.
FIGURE 6. Moenkhausia sthenosthoma. (a) MCP 48789, holotype, 56.0 mm SL. (b) UFRO 311, 57.21 mm SL. (c)UFRO
13621, 50.31 mm SL.
Color in alcohol. Overall coloration slightly silvery or yellow tan. Field of few dark chromatophores on upper
lip and maxilla. Infraorbital and opercular series silvery due to the presence of guanine pigmentation. Dark
midlateral stripe extending from posterior margin of opercle to base of median caudal fin-rays. In some individuals,
silver stripe is not preserved. Irregularly shaped humeral mark located over fourth to eight lateral line scales and
extending vertically over four-five horizontal scale rows above and over one-two horizontal scale rows below
lateral line. Paired fins and anal fin hyaline. Round dark spot at the base of the caudal-fin rays formed by few
chromatophores. Adipose with very few dark chromatophores (Fig. 5).
Distribution. Moenkhausia sthenosthoma occurs in the Rio Madeira basin (Fig. 4).
Etymology. The species name sthenosthoma is from the Greek sthenos meaning strong, and sthoma meaning
mouth, in reference to the strong musculature associated to the dentary, and robust teeth of the premaxilla and
dentary, a characteristic feature of the Moenkhausia jamesi species complex.
Moenkhausia jamesi Eigenmann, 1908

(Figs. 7a–c, 9a, Table 2)
Moenkhausia jamesi Eigenmann, 1908: 102. Type Locality: “Iça; Obidos; Lago do Máximo; Tajapuru”.
Diagnosis. Moenkhausia jamesi is distinguished from M. justae, M. ischyognatha, and M. alesis, by the number of
teeth on dentary (five vs. four in the last three). It is further distinguished from M. justae, M. alesis, and M.
sthenosthoma by the number of scale rows between the lateral line and midventral scale series (six vs seven in M.
sthenosthoma, and seven to eight in M. justae and M. alesis). Additionally, it is distinguished from M. justae by the
number of teeth in maxilla (edentulous vs one tooth with 5 cusps in M. justae), and by the number of cusps on the
fourth dentary tooth (three to five vs six in M. justae). Moenkhausia jamesi is additionally distinguished from M.
ischyognatha by its greater body depth (45.0–51.3% in SL vs 38.2–43.9% in SL in M. ischyognatha) and by its
greater head depth (34.4–39.2% in SL vs 29.3–32.6 in SL in M. ischyognatha).
arranged in a pronounced arch when examined in a ventral view; outer row with four to five pentacuspid teeth;
edentulous maxilla. Dentary bearing four to five pentacuspidate teeth with central cusp longest followed by three to
four distinctly small conical or tricuspidate teeth. First two or three dentary teeth from the symphysis with cusps
Dorsal-fin rays ii,9. Pectoral-fin rays i,11(3), 12*(37), 13(20), 14(4). Tip of pectoral fin extends slightly
beyond anterior insertion of pelvic fin. Pelvic-fin rays i,7, Tip of adpressed pelvic fin not reaching anal fin. Anal-
fin rays iv,28(11), 29(15), 30(18), 31(15), 32*(5). Caudal fin forked with i,9,8,i.
Scales cycloid. Lateral line with 34*(3), 35(11), 36(24), 37(22), 38(4) perforated scales; Scale rows between
lateral line and dorsal-fin origin seven. Scale rows between lateral line and midventral scale series six.
Circumpeduncular scale rows 13(8), 14(37), 15(13), 16(1). Scale sheath along anal-fin base 7–20, in one series,
covering base of anteriormost rays. Small scales covering proximal two-third of caudal-fin lobes.
First gill arch with 12(18), 13(23) gill rakers on lower limb and 8(10), 9(25), 10(6) on upper limb. Total
FIGURE 7. Moenkhausia jamesi. (a) MCZ 20734, syntype, 51.3 mm SL. (b) MZUSP 105860, 33.4 mm SL. (c) MCP 28197,
50.84 mm SL.
silver stripe is not preserved. Irregularly shaped humeral mark located over fourth to eight lateral line scales and
chromatophores. Adipose with very few dark chromatophores (Figs. 7a–c).
Distribution. Moenkhausia jamesi occurs along Rio Amazonas basin and appears to be restricted to the
Amazon lowlands area, as defined by Lima & Ribeiro (2011) (Fig. 4).
Material examined. Types: MCZ 20734, 1, syntype, 51.3 mm SL, Amazonas, Lagoa do Máximo (lake near to
Parintins), 2°40" S, 56°45". MCZ 20742, 1, syntype, 53.6 mm SL, Pará, Tajapuru (Ilha de Marajó, Furo Tajapuru),
1° 50" S, 50° 25". MCZ 20816, 2, syntypes, 47.0–53.1 mm SL, Amazonas, Rio Putomajo, Rio Iça (tributary of Rio
Solimões), near to Brazil/Colombia border, 3°7" 0' S, 67° 58" 0' W. MCZ 20827, 1, paralectotype (by present
designation), 38.5 mm SL, Pará, Obidos, (Rio Amazonas), 1°52"0' S, 55° 30" W. Non Types: USNM 307025, 2,
49.8–52,9 mm SL, Amazonas, Ressaca da Ilha de Marchantaria. USNM 307035, 2, 37.1–46,9 mm SL, Amazonas,
Paraná de Janauacá, entrance of Lago do Castanho. USNM 307323, 2, 58.6–59.4 mm SL, Amazonas. USNM
307331, 3, 31.6–40.5 mm SL, Amazonas, Lago Murumuru, curral de gado, Janauacá. USNM 307362, 1, 42.5 mm
SL, Amazonas, Ressaca da Ilha de Marchantaria. USNM 308062, 1, 36.9 mm SL, Amazonas, Paraná de Janaucá,
entrance of Lago do Castanho. USNM 308197, 2, 23.8–24.9 mm SL, Amazonas. MZUSP17583, 2, 47.9–52.4 mm
SL, Amazonas, Jutaí; Ilha Xibeco, rio Solimões, upstream of mouth of Rio Jutaí; 02°45’00”S, 66°45’00”W. LIRP
5010, 1, 35.9 mm SL, Amazonas, Manaus; Rio Solimões, Paraná de Janauacá, entrance of Lago do Castanho;
3°33’28”S,59°11’38”W. MZUSP 105860, 1, 33.6 mm SL, Amazonas, Manaus; Rio Solimões, Paraná de Janauacá,
entrance of Lago do Castanho; 3°33’28”S, 59°11’38”W. MZUSP17430, 4, 47.8–52.2 mm SL, Amazonas; Rio
Solimões, in front of Jacaré, near to Fonte Boa; 3°36’10”S, 61°19’02”W. MZUSP 49507, 54 (5 c&s), 44.9–56,9
mm SL, Acre; Rio Acre, Bôca do Acre, Rio Purus; 8°43’48”S, 67°23’55”W. MCP 47788, 4, 49.9–55.7 mm SL,
Brasil, Pará, Rio Tapajós, Alter do Chão, 02°31’00”S, 54°57’00”W, R.B. Oliveira, 03 Dec 1997.
Moenkhausia justae Eigenmann, 1908

(Figs. 8a–b, 9b, Table 2)
Moenkhausia justae Eigenmann 1908: 102. Type locality: “Uncertain in Amazon, Brazil”.
Diagnosis. Moenkhausia justae is distinguished from M. jamesi, M. ischyognatha, M. alesis and M. sthenosthoma
by the number of teeth in the maxilla (one tooth with five cusps vs an edentulous maxilla in the last four) and by the
number of cusps on the fourth dentary tooth (six vs three to five in the last four) (Fig. 9b). Moenkhausia justae is
further distinguished from M. ischyognatha and M. sthenosthoma by the number of scale rows between lateral line
and dorsal-fin origin (eight vs seven in the last two). Additionally, it differs from M. ischyognatha by its greater
head depth (34.9–40.2% in SL vs 29.3–32.6 in SL in the last three). It also differs from M. jamesi by the number of
scale rows between lateral line and the midventral scale series (seven to eight vs six scale rows in M. jamesi).
arranged in a pronounced arch when examined from a ventral view; outer row with four to five pentacuspidate
teeth; one pentacuspidate tooth on maxilla. Dentary bearing four*(52) or five (15) penta to heptacuspidate teeth
with central cusp longest followed by two to four distinctly small conical or tricuspidate teeth. Fourth dentary tooth
with six to seven cusps. First two or three dentary teeth from the symphysis with cusps arranged in a pronounced
arch when examined from a dorsal view.
TABLE 2. Morphometric data for Moenkhausia jamesi and Moenkhausia justae. Range includes the listed examined
material.
Moenkhausia jamesi (n=73) Moenkhausia justae (n=75)
N Syntypes Mean Range Mean Holotype Range Mean
Standard Length (mm) 4 38.5–54.6 33.4–56.9 45.2 32.1–63.4
Percentage of Standard Length
Greatest depth 3 45.1–48.5 47.0 44.3–52.3 47.9 45.0 41.4–51.9 47.0
Snout to dorsal-fin origin 3 20.8–52.3 51.5 49.2–54.5 51.9 51.2 47.5–54.0 51.2
Snout to pectoral-fin origin 3 26.1–28.3 26.9 25.9–30.3 27.7 27.6 25.7–31.2 27.9
Snout to pelvic-fin origin 3 45.8–47.8 46.5 47.4–52.9 49.5 49.7 48.6–53.8 51.3
Snout to anal-fin origin 3 64.3–65.5 65.0 63.2–71.7 68.0 66.9 65.1–74.2 67.8
Caudal-peduncle depth 3 10.5–11.6 11.3 10.2–13.3 12.0 12.7 10.4–13.4 11.8
Caudal-peduncle length - - - 4.0–8.4 5.7 10.2 4.2–9.4 5.9
Pectoral-fin length 3 21.4–22.8 22.0 21.1–24.9 22.7 21.0 14.1–23.0 21.0
Pelvic-fin length 3 17.0–19.4 18.1 17.9–20.8 19.3 16.9 14.3–19.3 17.5
Dorsal-fin length 2 32.4–34.3 33.3 33.1–39.3 35.7 27.5 28.4–35.7 31.9
Dorsal-fin base - - - 14.7–18.0 16.3 - 14.7–17.6 15.9
Anal-fin length 3 16.2–19.7 17.9 16.6–22.6 19.4 13.1 14.5–23.2 19.5
Anal-fin base - - - 34.0–40.2 36.8 - 32.4–37.7 35.4
Eye to dorsal-fin origin 3 36.8–38.8 37.6 37.0–41.9 39.1 37.2 34.5–41.0 38.3
Dorsal-fin origin to caudal-fin - - - 52.5–58.4 55.9 - 52.0–58.5 55.7
origin
Head length 3 24.5–26.1 25.1 23.3–29.0 25.3 25.5 22.8–28.2 25.0
Head depth - - - 34.4–39.2 37.2 - 34.9–40.2 37.4
Percentage of Head length
Snout length 3 25.6–27.8 26.9 21.2–31.0 26.3 27.5 22.3–31.8 26.6
Maxillary length 3 33.6–36.8 35.3 30.6–37.7 34.5 34.6 30.5–41.7 36.8
Horizontal orbital diameter 3 40.6–43.3 41.8 39.3–49.8 44.7 43.6 38.9–48.9 44.1
Least interorbital width 3 36.9–39.7 38.8 35.0–49.3 39.7 37.6 36.6–47.2 41.3
anterior insertion of pelvic fin. Pelvic-fin rays i,7, tip of adpressed pelvic fin not reaching anal fin. Anal-fin rays
iv,28*(12), 29(25), 30(21), 31(9). Caudal fin forked with i,9,8,i.
line and dorsal-fin origin 7(21) 8*(45). Scale rows between lateral line and midventral scale series 7(60), 8*(7).
Circumpeduncular scale rows 13(2), 14*(8), 15(21), 16(25), 17(6), 18(3). Scale sheath along anal-fin base 7–21, in
one series, covering base of anteriormost rays. Small scales covering proximal two-third of caudal-fin lobes.
First gill arch with 11(6), 12(40), 13(10) gill rakers on lower limb and 8(20), 9(34), 10(6) on upper limb. Total
silver stripe is not preserved. Irregularly shaped humeral mark located over fourth to eight lateral-line scales and
chromatophores. Adipose with very few dark chromatophores (Figs. 8a–b).
FIGURE 8. Moenkhausia justae. (a) MCZ 21014, holotype, 45.2 mm SL. (b) MCP 20546, 58.9 mm SL.
Distribution. Moenkhausia justae occurs along Rio Amazonas basin and the middle and lower Rio Araguaia,
and appears to be restricted in Amazon lowlands area, as defined by Lima & Ribeiro (2011) (Fig. 4).
Remarks. Lima et al. (2003) briefly discussed that the proposition of Eschmeyer et al. (1998) indicating that
the type locality of M. justae could be Rio Paraiba do Norte, João Pessoa, Paraíba was undisclosed and that recent
collections at João Pessoa and other sites in northeastern Brazil did not reveal specimens that could be assigned to
this species. Although the type locality of M. justae was not provided at the original description (Eigenmann 1908),
this author, in 1917, stated that the single (type) specimen of M. justae “came with others from Dr. Justa through
Major Coutinho and was probably found in the neighborhood of Manaos”. In fact, an image of the catalogue book
of the Museum of Comparative Zoology (available at: http://ids.lib.harvard.edu/ids/view/36195367?buttons=y)
indicates that the locality of M. justae is “Prob. Manaos” (= probably Manaus). According to Lima et al. (2003),
this should be considered as uncertain, but with strong evidence it is around Manaus, AM, Brazil. We identified M.
justae from two major tributaries of the Rio Solimões, Rio Japurá and Rio Branco (tributary of Rio Negro),
reinforcing the idea of Manaus as its type locality rather than João Pessoa, PB.
FIGURE 9. Dentary of Moenkhausia jamesi species complex. (a) M. jamesi, MZUSP 49507, 54.0mm SL; (b) M. justae, INPA
21489, 47.0mm SL; (c) M. ischyognatha, LBP 15948, 48.2mm SL; (d) M. alesis, INPA 48444, 50.4mm SL; (e) M.
sthenosthoma, LBP 19956, 46.0 mm SL; and (f) M. sthenosthoma, LBP 19958, 63.4mm SL. Scale bars= 1mm.
Material examined. Type: MCZ 21014, holotype, 45.2 mm SL, Brasil, J. M. S. Coutinho, 02-Ago1865.
MZUSP 55752,12, 38.7–49,5 mm SL, Brasil, Roraima, Rio Branco, near to Viruá; 1°14’59”S, 61°50’22”W, J.N.
Baskin 8 Dec 1993. Non-types: MZUSP 103127, 18, 42.8–49.7 mm SL, Amazonas, Japurá, Rio Japurá, Acanauaí,
01°50’00”S, 66°36’00”W, Expedição Permanente à Amazônia, 30 Nov–02 Dec 1997. MZUSP 55752, 12, 39.4–
49.2 mm SL, Roraima, Rio Branco, near to Viruá, J. N. Baskin, 08 Dec 1993. MCP 19661,6, 53.2–60.0 mm SL,
Brasil, Pará Santarém, Rio Curuatinga, R.B. Oliveira, 01 Nov 1996. MCP 20546, 16, 51.9–63.4 mm SL, Brasil,
Pará, Rio Tapajós, Alter do Chão, 02°31’00”S, 54°57’00”W, R.B. Oliveira, 03 Dec 1997. NUP 8106, 3 of 5, 54.5–
56.0 mm SL, Tocantins, Xambioá; Ribeirão Xambioá, tributary of Rio Araguaia, Rio Tocantins-Araguaia system,
06°24’27”S, 48°36’54”W, Gerpel, 22 Mar 2009. NUP 8136, 2 of 3, 47.7–55.1 mm SL, Tocantins, Xambioá, Rio
Araguaia, tributary of Rio Tocantins, Rio Tocantins-Araguaia system; 06°11’11”S, 48°26’20”W; Gerpel, 09 Jul
2009. INPA 21489, 9, 32.1–51.5 mm SL, Caseara, Parque Estadual do Cantão, E.G. Ferreira, J. A. Zuanon & G. M.
Santos, May 2000.
Discussion
Mariguela et al. (2013) performed a phylogenetic analysis based on molecular characters encompassing 29 from
the 76 valid species of Moenkhausia. Their results showed that Moenkhausia jamesi (=M. sthenosthoma sp. n.) is
more closely related to M. lata. This clade (M. jamesi + M. lata) is, in turn, the sister group of the clade formed by
M. dichroura, M. intermedia, and M. costae. Among all the examined species of Moenkhausia, only the
aforementioned species have premaxillary and dentary teeth with cusps arranged in a pronounced arch (C-shaped).
However, the short and curved maxilla, as described for the M. jamesi species complex, was not observed in any
other species of Moenkhausia.
It is noteworthy that M. lata shares with the M. jamesi complex the characteristic humeral blotch, which can
indicate that this feature may actually be a synapomorphy of this clade rather than for the M. jamesi species
complex, which would be expected considering the hypothesis of Mariguela et al. (2013). A similar humeral mark
is also present in Moenkhausia hysterosticta, differing only by the fact it is located on fifth through eighth lateral
line scales and extending two rows above and one row below lateral line (vs. humeral blotch located on fourth
through seven lateral line scales and extending up to four rows above and two below the lateral line in the M.
jamesi species complex and M. lata). Although M. hysterosticta also shares with M. lata a black mark on dorsal
lobe of caudal fin, it lacks the C-shaped premaxillary and dentary teeth. Indeed, Géry (1992) affirmed that this
caudal mark seems to have no phylogenetic implication and it is probably used as a recognition signal among
unrelated species for defensive associations.
The characteristic morphology of the mouth (teeth, premaxilla, maxilla, and dentary) that defines the
Moenkhausia jamesi species complex is comparable to that observed in the genus Ctenobrycon, and in Astyanax
aleni, A. pellegrinni, A. correntinus, and A. erythropterus, in which it consists of premaxillary and dentary teeth
with cusps arranged in a pronounced arch, posterior margin of maxilla not exceeding the anterior third of second
infraorbital, and very similar general morphology of premaxilla and dentary. Nevertheless, according to Mariguela
et al. (2013), there is no close relationship between M. jamesi (=M. sthenosthoma) and Ctenobrycon hauxwellianus
(=C. spilurus), or with any analyzed species of Astyanax (i.e. A. aeneus, A. jordani, and A. mexicanus), which is
partially congruent with Mirande's (2010) conclusions. According to this author, premaxillary teeth with cusps
aligned to form a concave semicircle in ventral view (his character 127) is one of the two synapomorphies of his
clade 262, formed by Ctenobrycon? (not examined by that author), Markiana, Psellogrammus, Astyanax cf.
abramis, A. assuncionensis, A. correntinus, A. lineatus, A. pellegrini, and other Astyanax? (e.g. Astyanax
magdalenae). The presence of this condition in M. dichroura was considered homoplastic by Mirande (2010),
which is also congruent to Mariguela et al. (2013). The presence of such a modified mouth in these non-related
groups may represent an interesting case of adaptive convergence in Characidae.
Comparative material examined. Astyanax assuncionensis: LBP 19923, 2, 45.0–50.7 mm SL, Argentina,
Arroyo Leyes, Santa Fé. Astyanax correntinus: LBP 19925, 4, 64.7–74.3 mm SL, Argentina, Arroyo Leyes, Santa
Fé. Astyanax erythropterus: LBP 19924, 2, 70.2–74.3 mm SL, Argentina, Arroyo Leyes, Santa Fé. Astyanax
pellegrini: MZUSP 54023, 33, 46.6–74.4 mm SL) Astyanax magdalenae: LBP 6108, 7, 66.9–74.8 mm SL,
Venezuela, Machiques de Perijá/Zulia, Lago Maracaibo, Rio Santa Rosa. Ctenobrycon alleni: LBP 13532, 2, 31.2–
42.0 mm SL, ot measured, Ctenobrycon spilurus: MZUSP 5156,28, not measured, Roraima, Surumu, rio Surumu.
Markiana nigripinnis: LBP 7586, 84.4 mm SL. Brasil, Mato Grosso, Barão de Melgaço, shore lake of rio Cuiabá.
Moenkhausia affinis: NMW 56322, syntypes, 35.7–64.8 mm SL, Brazil, foz do Rio Negro. Moenkhausia agnesae:
ZMA 113863, paratype, 63.5 mm SL, Brazil, Igarapé Preto, upper Amazonas. Moenkhausia aurantia: MZUSP
107827, paratype, 1 of 8, Brazil, Goiás, upper Tocantins, córrego Kavanca. Moenkhausia australe: CAS 70818-19,
Syntypes, 31.2–38.9 mm SL, Paraguai, Anistis. Moenkhausia barbouri: MCZ 20708, Syntypes, 47.4–50.9 mm SL,
Brazil, Amazonas, Parintins. Moenkhausia bonita: MCP 30686, paratypes, 3, 30.7–35.4 mm SL, Brazil, Mato
Grosso do Sul, upper rio Miranda. Moenkhausia browni: MZUSP 17140, 2, 30.48–37.66 mm SL, Brazil, Pará,
Castanhal, Igarapé Apeu. Moenkhausia celibela: MZUSP 93197, paratype, 1 of 51, Brazil, Pará, Pimental, rio
Tapajós. Moenkhausia ceros: MZUSP 30275, 2, 31.3–38.7 mm SL, Brazil, Pará, Pederneiras, rio Tapajós.
Moenkhausia chlorophthalma: MZUSP 97092, paratype, 1 of 28, Brazil, Pará, Altamira, rio Xingu. Moenkhausia
chrysargyrea: USNM 226155, 2, 52.32–49.31 mm SL, Suriname, Nickerie District. Moenkhausia collettii: LBP
4153, 1, 33.1 mm SL, Brazil, Acre, Mâncio Lima, rio Moa. LBP 6882, 1, 32.1 mm SL, Brazil, Amazonas, São
Gabriel da Cachoeira. Moenkhausia comma: MCZ 20972, syntype, 60,1 mm SL, Brazil, Amazonas, Lago Cudajas.
Moenkhausia copei: MZUSP 30263, 2, 30.91–35.2 mm SL, Brazil, Roraima, Maracá, rio Uraricoera. Moenkhausia
cosmops: MZUSP 93556, 3, 28.6–31.0 mm SL, Brazil, Tapajós drainage, rio Juruena. Moenkhausia costae: LBP
8271, 5, 36.6–43.5 mm SL, Brazil, rio Verde, São Franscisco drainage. Moenkhausia cotinho: MCZ 21070,
syntypes, 2, 51.0–54.5 mm SL, Brazil, Pará, Belém. Moenkhausia crisnejas: CAS 60477, 3, 30.24–35.14 mm SL,
Peru, Paipay, rio Crisnejas. Moenkhausia diamantina: MNRJ 30168, holotype, 57.3 mm SL, Brazil, Bahia,
Lençois, Foz do rio Toalha. Moenkhausia dichroura: LIRP 250, 1, 47.38 mm SL, Brazil, São Paulo, Ribeirão Preto,
rio Pardo basin. Moenkhausia diktyota: LBP 7074, 10 of 91, 34.9–46.1 mm SL, Brazil, rio Negro e rio Amazonas
drainage. Moenkhausia doceana: MZUSP 54777, 2, 43.99–47.3 mm SL, Brazil, Espírito Santo, São Mateus, rio
Bamburral. Moenkhausia dorsinuda: MTD F 24833, holotype, 70.9mm SL, Bolívia, Beni, rio Ipurupuru.
Moenkhausia dosalmas: MZUSP 106076, paratype, 28.9mm SL, Brazil, Goiás, Cavalcante. Moenkhausia
eigenmanni: USNM 167811, holotype, 49.0mm SL, Colômbia, rio Manacacias. Moenkhausia eurystaenia:
MZUSP 91404, paratype, Brazil, Mato Grosso, Nova Ubiratã, Xingu drainage, rio Von Den Stein. Moenkhausia
forestii: MZUSP 97827, holotype, 34.0 mm SL, Brazil, Mato Grosso, Cáceres, rio Sepotuba, rio Paraguay basin.
Moenkhausia georgiae: ANSP 177000, 1, 48.48 mm SL, Guiana, Essequibo, rio Siparuni. Moenkhausia gracilima:
USNM 310357, 2, 29.84–30.42 mm SL, Brazil, Amazonas, rio Solimões. Moenkhausia grandisquamis: MZUSP
101474, 9, 42.5–60.8 mm SL, Brazil, Pará, Monte Dourado, rio Jari. Moenkhausia hemigrammoides: MZUSP
30531, 1, 35.8 mm SL, Brazil, Roraima, rio Branco. Moenkhausia heikoi: MTD F 27909, paratype, 43.4mm SL,
Brazil, Pará, rio Xingu basin, rio Novo. Moenkhausia hysterosticta: MCP 32559, holotype, Brazil, Tocantins,
Ipueiras,rio Tocantins. Moenkhausia inrai: MNHN 1992-0943, holotype, 64.3mm SL, Guiana Francesa, riacho
Rock. Moenkhausia intermédia: LIRP 346, 2, 46.3–50.4 mm SL, Brazil, São Paulo, Luís Antônio, lagoa do Diogo.
Moenkhausia lata: MCZ 20860, syntype, 77 mm SL, Brazil, Pará, rio Tapajós. Moenkhausia latíssima: MCZ
20769, syntypes, 3, 42.69–66.65 mm SL, Brazil, Amazonas, Tabatinga. Moenkhausia lepidura: MZUSP 34627, 2,
60.88–64.68 mm SL, Brazil, Amazonas, rio Negro basin. Moenkhausia levidorsa: INPA 16774, holotype, 59.2 mm
SL, Brazil, Mato Grosso, Furo Bahia. Moenkhausia lopesi: LIRP 4642, 1, 37.8mm SL, Brazil, Mato Grosso do Sul,
upper rio Piquiri. Moenkhausia loweae: MTD F 27524, paratype, 49.4mm SL, Brazil, Xavantina, rio das Mortes.
Moenkhausia margitae: MTD F 17256, holotype, 64.1mm SL, Peru, Ucayali, Quebrada Campo Verde.
Moenkhausia megalops: INPA 7031, 1, 70.0 mm SL, Brasil, Pará, Itaituba, Rio Tapajós. Moenkhausia metae: MTD
F 17987, 1, 42.7mm SL, Colômbia, Restrepo, upper rio Meta. Moenkhausia melogramma: USNM 310776, 2, 25–
28.46 mm SL, Brazil, Amazonas, rio Madeira basin. Moenkhausia miangi: NMW 56297-300, Syntypes, 29.7–88.9
mm SL, Brazil, rio Miang. Moenkhausia mikia: MZUSP 81219, paratype, 1 of 52, Brazil, Amazonas, Rio Negro
drainage, rio Tiquié. Moenkhausia moisae: MNHN 1995-1073, holotype, 44.0 mm SL, Guiana Francesa.
Moenkhausia naponis: USNM 164067 holotype, 52.9 mm SL, Equador. Moenkhausia nigromarginata: LBP 9033,
13 of 33, 40.08–49.9 mm SL, Brazil, Amazonas, rio Juruena, rio Verde drainage. Moenkhausia oligolepis: LBP
5676, 5, 29.0–79.6 mm SL, Brazil, Mato Grosso, Barra dos Garças, Amazonas Araguaia drainage. Moenkhausia
orteguasae: ANSP 70496, holotype, 57.9 mm SL, Colômbia, Caquetá, rio Orteguasa. Moenkhausia ovalis: BMNH
1867.6.13.87, holotype, 75.6 mm SL, Peru, Xeberus. Moenkhausia pankilopteryx: MZUSP 30289, paratype,
Brazil, Pará, Tocantins drainage, rio Itacaiunas. Moenkhausia petymbuaba: MZUSP 30231, 49.5 mm SL, Brazil,
Pará, rio Curuá, Serra do Cachimbo. Moenkhausia phaeonota: LBP 15857, 3, 25.3–28.2 mm SL, Brazil, Mato
Grosso, Querência, rio Xingu drainage. Moenkhausia pirauba: LBP 5689, 1, 76.0 mm SL, Brazil, Barra do Garça.
Moenkhausia pittieri: MCNG 140301, 1, 42.2mm SL, Venezuela. Moenkhausia plúmbea: MZUSP 101435,
paratype, 1 of 24, Brazil, Pará, Novo Progresso, rio Tapajós drainage. Moenkhausia robertsi: USNM 200427,
holotype, 47.7 mm SL, Peru, Iquitos, upper rio Amazonas. Moenkhausia sanctaefilomenae: LBP 5541, 49, 27.4–
46.1 mm SL, Brazil, Piauí, Santa Filomena. Barra do Garça, Mato Grosso, rio Araguaia drainage. Moenkhausia
shideleri: MCNG 16904, 2, 26.8–36.7 mm SL, Venezuela, Bolivar, rio Cuyuni. Moenkhausia simulata: USNM
167811, 2, 47.45–50.54mm SL, Peru, rio Pichis. Moenkhausia surinamensis: ZMA 104.221, holotype, 48.8 mm
SL, Suriname, Brownscreek. Moenkhausia takasei: ZMA 113.849, paratype, 27.3 mm SL, Brazil, Pará, rio Guama
basin. Moenkhausia tergimacula: MCP 20287, 1, 34.7 mm SL, Brazil, Goiás, Minaçu, rio Tocantins. Moenkhausia
tridentata: NMW 8947, holotype, 118.6 mm SL, Brazil, Pará, io Amazonas, Piquirão. Moenkhausia xinguensis:
LBP 16745, 10, 38.5–53.4 mm SL, Brazil, Pará, rio Xingu, Rio Amazonas drainage.
Acknowlegdments
We thank Aléssio Datovo and Osvaldo T. Oyakawa (MZUSP), Carla S. Pavanelli (NUPELIA), Lúcia H. Rapp Py-
Daniel and Renildo R. Oliveira (INPA), Angela Araújo and Mariluce Fernandes (UFRO), Carlos A. S. de Lucena
(MCP), Claudio Oliveira (LBP), Karsten E. Hartel (MCZ), Mark H. Sabaj Pérez (ANSP), and Richard P. Vari
(USNM) for loan of comparative material and curatorial assistance. Figure 1 was modified from the original
prepared by Andrew Williston (MCZ). Figures 7a and 8a were provided by the Museum of Comparative Zoology
(MCZ), Harvard University. Valter M. Azevedo-Santos assisted with the preparation of Figure 4 and in clearing
and staining material. Cibele B. Raio examined and provided information on type material deposited at MTD,
NMW, and ZMA. The authors were funded by FAPESP proc. 2013/24729-0, 2015/01972-1 (MGP), and 2000/
001920-6, 2006/005453 (RCB). This study was also part of the FAPESP Thematic Project “Phylogenetic
relationships in the Characidae (Ostariophysi: Characiformes)” (FAPESP grant # 04/09219-6), and benefited from
the FAPESP Thematic Project “South American Characiformes Inventory” (FAPESP grant # 11/50282-7).
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Three new species of Moenkhausia (Teleostei, Characiformes, Characidae)

Article in Zootaxa · July 2015

Marina Petrolli Ricardo C Benine

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Description of three new species of Moenkhausia (Teleostei, Characiformes,

MARINA G. PETROLLI & RICARDO C. BENINE

Key words: Moenkhausia justae, taxonomic revision, Systematics, Neotropical region

Palavras-chave: Moenkhausia justae, revisão taxonômica, Sistemática, região Neotropical

Accepted by M.R. de Carvalho: 18 Jun. 2015; published: 20 Jul. 2015 401

Material and methods

THREE NEW SPECIES OF MOENKHAUSIA

Zootaxa 3986 (4) © 2015 Magnolia Press ·

Moenkhausia alesis, new species

FIGURE 5. Moenkhausia alesis, MZUSP 117139, holotype, 52.6 mm SL.

Moenkhausia sthenosthoma, new species

Moenkhausia jamesi Eigenmann, 1908

Moenkhausia justae Eigenmann, 1908

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