Zootaxa 3994 (1): 133141

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Copyright 2015 Magnolia Press

Article

ISSN 1175-5326 (print edition)

ZOOTAXA

ISSN 1175-5334 (online edition)

http://dx.doi.org/10.11646/zootaxa.3994.1.7
http://zoobank.org/urn:lsid:zoobank.org:pub:D9D78466-2E20-4BAA-8520-E0F2FBBDDB1D

Bryconops munduruku (Characiformes: Characidae), a new species of fish from

the lower Tapajs River basin, Brazil
CRLISON SILVA-OLIVEIRA1, ANDR LUIZ C. CANTO2 & FRANK RAYNNER V. RIBEIRO1,2
1

Programa de Ps-Graduao em Recursos Aquticos Continentais Amaznicos, Instituto de Cincias e Tecnologia das guas, Universidade Federal do Oeste do Par. Rua Vera Paz, s/n Sal, CEP 68035-110, Santarm, Par, Brazil.
E-mail: carlison3@gmail.com; fraynner@yahoo.com.br
2
Coleo Ictiolgica da Universidade Federal do Oeste do Par. Campus Amaznia. Avenida Mendona Furtado, 2.946 - Ftima,
CEP 68040-470, Santarm, Par, Brazil. E-mail: cantoandre@gmail.com

Abstract
A new species of Bryconops is described from a right tributary of the lower Tapajs River, State of Par, Brazil. Bryconops
munduruku, sp. nov., differs from its congeners, except B. inpai and B. piracolina, by having a black adipose fin (vs. adipose fin hyaline in alcohol) and, except B. inpai, by possessing two humeral blotches (vs. lack of humeral blotch or only
one humeral blotch). Bryconops munduruku differs from B. inpai by having a uniform color pattern on the posterior portion of the side of the body (vs. a dark stripe extending posteriorly from the half of the anal-fin base onto the base of the
middle caudal-fin rays). It differs further from B. inpai and B. piracolina by the presence of a black adipose fin that is
hyaline along its base (vs. entirely black adipose fin in B. inpai and B. piracolina). The new species is allocated in the
subgenus Creatochanes by having a maxillary bone with one to three teeth on both sides, and its posterior extension reaching the junction of second and third infraorbital bones.
Key words: subgenus Creatochanes, taxonomy, Tapajs National Forest, Amazon basin

Resumo
Uma espcie nova de Bryconops descrita de um afluente da margem direita do baixo rio Tapajs, Par, Brasil. Bryconops
munduruku difere de seus congneres, exceto B. inpai e B. piracolina, pela nadadeira adiposa preta (vs. nadadeira adiposa
hialina em lcool) e, exceto B. inpai, por apresentar duas manchas umerais. Bryconops munduruku difere de B. inpai pelo
padro uniforme de colorao na poro posterior do corpo (vs. uma faixa lateral escura que se estende posteriormente da
metade da base da nadadeira anal at a base dos raios medianos da nadadeira caudal). Difere ainda de B. inpai e B. piracolina pela nadadeira adiposa escura distalmente, mas hialina ao longo de sua base (vs. nadadeira adiposa completamente
escura em B. piracolina e B. inpai). A espcie nova alocada no subgnero Creatochanes por apresentar de um a trs dentes
em cada osso maxilar e a margem posterior da maxila alcanando a margem posterior do segundo infraorbital.

Introduction
Bryconops was proposed by Kner (1858) to allocate two new species, Bryconops alburnoides Kner and Bryconops
lucidus Kner (= Bryconops alburnoides Kner), described based on specimens from the Guapor and Branco rivers,
Brazil, respectively. Currently 19 valid species are recognized in the genus (Lima et al. 2003; Chernoff &
Machado-Allison 2005; Wingert & Malabarba 2011; Eschmeyer 2015). The genus is restricted to freshwater
drainages of South America, occurring in the Amazon, Orinoco, Tocantins, So Francisco, and Paraguay river
basins, and in coastal rivers of northern Brazil and the Guianas (Lima et al. 2003; Lima & Caires 2011; Polaz et al.
2014; Ramos et al. 2014; Eschmeyer 2015).
Chernoff & Machado-Allison (1999, 2005) divided the species of Bryconops into two subgenera, Bryconops
and Creatochanes Gnther 1864, based on the number of maxillary teeth and position of the posterior margin of the
maxillary bone in relation to the contact between the second and third infraorbital bones.
Accepted by M.R. de Carvalho: 18 May 2015; published: 30 Jul. 2015

133

Among the valid species, eight exhibit characters proposed for the subgenus Bryconops: B. alburnoides, B.
caudomaculatus (Gnther 1864), B. collettei Chernoff & Machado-Allison 2005, B. disruptus Machado-Allison &
Chernoff 1997, B. durbini (Eigenmann 1908), B. gracilis (Eigenmann 1908), B. magoi Chernoff & MachadoAllison 2005, and B. piracolina Wingert & Malabarba 2011. Eleven species exhibit characters proposed for the
subgenus Creatochanes: B. affinis (Gnther 1864), B. colanegra Chernoff & Machado-Allison 1999, B. colaroja
Chernoff & Machado-Allison 1999, B. cyrtogaster (Norman 1926), B. giacopinii (Fernndez-Ypez 1950), B.
humeralis Machado-Allison, Chernoff & Buckup 1996, B. imitator Chernoff & Machado-Allison 2002, B. inpai
Knppel, Junk & Gry 1968, B. melanurus (Bloch 1794), B. transitoria Steindachner 1915, and B. vibex MachadoAllison, Chernoff & Buckup 1996.
In this paper, a new species of Bryconops is described from the igarap Au, a tributary of the lower Tapajs
River basin in Brazil.

Material and methods

Methods for taking morphometric and meristic data followed Fink & Weitzman (1974) and Machado-Allison et al.
(1993, 1996), except for counts of horizontal scale rows below the lateral line that were taken following Malabarba
& Bertaco (1999). All measurements were taken point to point, on the left side of the body, with digital caliper to
the nearest 0.1 mm, and are summarized in the species accounts and table as percentages of Standard Length (SL)
or Head Length (HL). Drawings and counts of vertebrae, supraneurals, pterygiophores, procurrent caudal-fin rays,
gill rakers on the first branchial arch, and branchiostegal rays were taken from cleared and stained (c&s) paratypes,
prepared following the clearing and staining methods of Taylor & van Dyke (1985).
Institutional abbreviations follow Ferraris Jr. (2007), with the addition of the Fish Collection of Universidade
Federal do Oeste do Par (UFOPA-I). Comparisons to species not listed in comparative material were based on the
literature.
In the description, counts are followed by number of frequencies, corresponding to the number of specimens
analyzed; asterisk within parentheses (*) indicates the holotype count. In the list of comparative material examined,
museum abbreviations and catalog numbers are followed by the total number of specimens in that lot, range of
standard length, and collecting data.

Bryconops munduruku, new species

(Figures 13; Table 1)
Holotype. INPA 46510, male, 76.6 mm SL, Brazil, Par State, Aveiro, igarap Au, tributary of the right margin of
the Tapajs River, about 10 km of the city of Aveiro, 03360.99S 551459.14W, 23 May 2013, C. Silva-Oliveira
& S. R. Oliveira.
Paratypes. All collected with holotype, Brazil, Par State, Tapajs River basin: MCP 48315 (5, 34.079.6 mm
SL); INPA 46511 (11, 33.591.7 mm SL; 2 c&s, 31.874.1 mm SL); UFOPA-I-00655 (7, 33.496.4 mm SL; 5
c&s, 47.663.3 mm SL).
Diagnosis. Bryconops munduruku differs from its congeners, except B. inpai and B. piracolina, by possessing
a black adipose fin (vs. adipose fin hyaline in alcohol). It differs from B. inpai and B. piracolina by possessing a
hyaline band on the black adipose-fin base (vs. entirely black adipose fin in B. piracolina and B. inpai). Bryconops
munduruku differs further from B. piracolina by having a hyaline dorsal fin (vs. presence of a large black blotch on
the dorsal-fin base). Bryconops munduruku differs from other species of the subgenus Creatochanes, except B.
inpai, by possessing two humeral blotches (vs. lack of humeral blotch or humeral region with a single humeral
blotch in B. humeralis and B. vibex). It differs further from B. inpai due to the uniform color pattern on the
posterior portion of the side of the body (vs. presence of a dark stripe extending posteriorly from the half of the
anal-fin base onto the base of the middle caudal-fin rays).
Description. Morphometric data presented in Table 1. Body shape compressed, greatest body depth located
just anterior to dorsal-fin origin. Dorsal profile of body slightly convex from snout tip to dorsal-fin origin; straight
to slightly concave from dorsal insertion to adipose-fin origin, gently sloping to beginning of caudal peduncle.

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Caudal peduncle concave along dorsal and ventral margins. Ventral profile of head and body convex from lower lip
to pelvic-fin origin, and straight from this point to anal-fin origin. Ventral profile straight and posterodorsally
inclined along anal-fin base.
TABLE 1. Morphometric data of holotype and 30 paratypes of Bryconops munduruku. Holotype is included in the
ranges and means. SD = standard deviation.

Standard Length (mm)

Holotype

Range

Mean

SD

76.6

31.896.4

61.4

Percents of Standard Length

Predorsal length

41.4

41.450.6

48.8

1.6

Preanal length

66.7

60.467.3

65.3

1.4

Prepelvic length

49.0

47.452.6

49.8

1.3

Prepectoral length

26.4

25.529.1

27.1

0.8

Dorsal-fin base length

13.1

9.817.9

14.5

1.4

Dorsal origin to pectoral origin

37.3

31.538.5

35.8

1.7

Dorsal origin to pelvic origin

33.7

27.135.3

31.9

2.0

Dorsal terminus to anal origin

35.1

27.637.8

34.1

2.5

Dorsal terminus to anal origin

28.1

21.530.8

27.4

2.2

Dorsal terminus to pelvic origin

33.4

27.735.4

32.1

1.9

Dorsal terminus to anal terminus

31.9

26.335.3

31.0

1.7

Dorsal terminus to adipose origin

22.1

21.326.6

22.9

1.1

Adipose terminus to hypural plate

14.6

11.716.0

14.4

1.1

Pectoral origin to pelvic origin

23.8

20.625.6

23.2

1.1

Anal-fin base length

25.7

24.629.5

27.2

1.2

Anal origin to adipose origin

30.4

27.133.0

30.1

1.6

Length of caudal peduncle

12.4

9.614.3

12.5

1.0

Adipose origin to base of last anal-fin ray

10.8

10.813.5

12.4

0.7

Maxillary length

14.0

13.115.7

14.1

0.7

Snout length

8.0

5.98.2

6.9

0.6

Horizontal orbit diameter

8.4

7.112.0

9.7

1.0

Head length

26.0

22.229.2

26.1

1.4

86.4

81.7102.7

89.1

4.6

Percents of Head Length

Tip of snout to tip supraoccipital spine
Maxilla to pectoral origin

45.7

44.659.8

50.8

3.9

Maxilla to posterior margin of opercle

55.8

51.769.3

57.8

3.7

Anterior margin of orbit to maxilla

38.2

32.646.7

39.2

2.7

Posterior margin of orbit to opercle

41.0

31.348.7

36.7

3.9

Maxillary length

53.8

48.560.7

54.0

2.1

Snout length

30.7

21.532.0

26.3

2.2

Horizontal orbit diameter

32.2

32.244.7

7.5

2.8

Mouth terminal. Posterior extension of maxilla reaching junction between second and third infraorbital bones
(Fig. 2); third infraorbital moderately developed, reaching preopercle ventrally. Supraorbital bone present.
Premaxillary teeth in two rows, with midcentral cusp more developed than remaining cusps; outer teeth row with
5* (9) or 6 (3) tricuspid teeth; inner tooth row with 5* (10) pentacuspid teeth. Maxillary with 1 (2), 2* (15), or 3 (1)
tri- or tetracuspid teeth on contralateral parts (Fig. 3). Dentary with 5 (2) or 6* (6) penta- or hexacuspid teeth,
followed by 6 smaller conical teeth.

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FIGURE 1. Bryconops munduruku, INPA 46510, holotype, male, 76.6 mm SL, Brazil, Par State, Aveiro, Igarap Au,
tributary of the lower Tapajs River basin.

FIGURE 2. Bryconops munduruku, UFOPA-I-00655, paratype, 71.5 mm SL (c&s). Left side of the premaxilla, maxilla, and
dentary. Lateral view. Scale bar = 1 mm.

Dorsal-fin rays ii,8 (3) or ii,9* (28); first unbranched ray about one-half length of second unbranched ray.
Dorsal-fin origin slightly ahead of mid-body, positioned vertically through anterior third of pelvic-fin base.
Posterior margin of dorsal fin straight to slightly concave. Adipose-fin origin approximately at vertical through
base of 18th to 19th anal fin branched rays. Principal caudal-fin rays i,8,8,i (7) or i,8,9,i* (23). Lobes of caudal fin
unequal, lower lobe slightly longer than upper lobe. Dorsal procurrent caudal-fin rays 14 (5), ventral procurrent
caudal-fin rays 13 (4) or 14 (1). Pectoral-fin rays i,10 (15) or i,11* (15). Tip of pectoral fin reaching slightly
beyond half of distance between pectoral and pelvic fins. Pelvic-fin rays i,7*(30). Pelvic fin originating at vertical
through middle of dorsal-fin base, reaching slightly beyond half the distance between pelvic and anal fins. Anal-fin

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rays iii, 23* (16), iii, 24 (6), iii, 25 (6), or iii, 26 (2). Anal-fin origin located posterior to vertical through base of last
dorsal-fin ray. Last unbranched anal-fin ray and first to fourth branched rays slightly longer than remaining anal-fin
rays. Distal margin of anal fin concave.

FIGURE 3. Bryconops munduruku,UFOPA-I-00655, paratype,63.3 mm SL(c&s). Maxilla (MA), infraorbitals (IO) 1-5 and
antobital (AO). Scale bar = 1 mm.

Lateral line reaching base of caudal-fin rays. Longitudinal scales in lateral line 44 (6), 45* (18), or 46 (8). All
scales of lateral line pored. Scale rows between lateral line and dorsal-fin origin 7* (22) or 8 (8); scale rows
between lateral line and pelvic-fin origin 3* (18) or 4 (12). Predorsal scales 10 (9) or 11* (5), arranged in a regular
series. Scale rows around caudal peduncle 15 (5) or 16* (3). Precaudal vertebrae 19 (5); caudal vertebrae 22 (1) or
23 (4); total vertebrae 41 (1) or 42 (4). Supraneurals 7 (4). First dorsal fin pterygiophore located between 11th and
12th vertebrae. Gill rakers of first gill arch 16 (5): 2 (5) hypobranchial, 7 (5) ceratobranchial, 1 (5) on cartilage
between ceratobranchial and epibranchial, 6 (5) epibranchial. Gill rakers setiform. Branchiostegal rays 5: 4 (5) on
ceratohyal and 1 (2) on epihyal.
Color in alcohol. Overall ground coloration of body dusky brown to yellowish. Dorsal profile of head and
snout dark. Infraorbitals, maxillary, and ventral region of opercle silver. Infraorbital 5 and dorsal portion of opercle
with scattered chromatophores. Lips and anterior portion of maxilla with dense concentration of chromatophores.
Gular region light. Dorsolateral region of body darkened. Scales of lateral region of body with chromatophores
concentrated on posterior borders. Two humeral spots present; first humeral spot conspicuous, vertically elongated,

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extending over three rows of scales above lateral line and horizontally over two rows of scales; second humeral
spot inconspicuous, vertically elongated; its form resembling the number 3. Dorsal-fin rays darkened from base to
mid-section, light at distal portion. Black adipose fin, but hyaline along its base. Caudal fin with concentrated
chromatophores on half nearest most external rays of ventral and dorsal lobes. Pectoral, pelvic, and anal fins
hyaline, with few scattered chromatophores.
Color in life. General body color reddish on dorsolateral region above lateral line; light grey from lateral line
to ventral scale series. Upper portion of head, opercle, intraorbital bones, and maxilla dark grey. Upper margin of
orbit red; anterior and posterior margins yellowish. Dorsal fin red from base to mid-length of rays; distally hyaline.
Black adipose fin, with narrow hyaline band at base. Caudal fin red at base to mid-length of dorsal and ventral rays;
distal region of lobes hyaline. Pectoral and pelvic fins, as well as first anal-fin rays, yellow.
Sexual dimorphism. Mature males bear hooks on the anal, pelvic, and dorsal-fins rays. Dorsal and pelvic-fin
hooks are smaller, fewer in number and spine-like. Anal-fin hooks larger and more numerous, present from distal
half of the third unbranched ray, decreasing in number and size to15th branched ray. Hooks absent in females.
Distribution. Bryconops munduruku is known from the igarap Au, a tributary on the right margin of the
Tapajs River, about 10 km from Aveiro, Par State, Brazil (Fig. 4).
Etymology. The specific epithet is given in allusion to a tribe of Munduruku Indians denominated TapajsTapera, who settled on the right margin of the Tapajs River, giving rise to what today is the city of Aveiro, the type
locality of Bryconops munduruku. A noun in apposition.
Ecological notes. igarap Au presents clear water running over sandy beds. Its headwaters are located within
the limits of the Tapajs National Forest and are densely covered by marginal vegetation. Representatives of
Bryconops munduruku were collected in rapid flowing waters among stretches, where it occurs syntopically with
Bryconops cf. giacopinii, Hyphessobrycon cf. agulha, Hemigrammus sp., and Moenkhausia comma.

FIGURE 4. Map of part of Central and Northern South America, showing the type locality of Bryconops munduruku (black
square) in the lower Tapajs River basin.

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FIGURE 5. Igarap Au, type locality of Bryconops munduruku, lower Tapajs River, Aveiro, Par State, Brazil.

Discussion
Bryconops munduruku shares with its congeners characters proposed by Chernoff & Machado-Allison (1999) as
synapomorphies for the genus, as follows: (1) ventral margin of maxillary bone curved at an angle of
approximately 90; (2) infraorbital sensory canal well developed, reaching the antorbital bone; and (3) sensory
canal extends onto nuchal scales. Within the genus Bryconops, Chernoff & Machado-Allison (1999) proposed the
monophyly of each subgenera, Bryconops and Creatochanes, based on two putative synapomorphies. Thus,
subgenus Bryconops is defined by the absence, or sporadic presence of a sole conical tooth located in a single
maxillary bone, and the posterior end of the maxilla not reaching the junction of second and third infraorbital
bones; and subgenus Creatochanes is recognized by the presence of one to three teeth on each maxilla, and the
posterior end of the maxilla reaching the region of contact between the second and third infraorbital bones. As for
Bryconops munduruku, it possesses the characters proposed by Chernoff & Machado-Allison (1999, 2005) that
define the subgenus Creatochanes.
Among its congeners, B. munduruku resembles B. inpai by having a deep body, presence of two humeral
blotches, and a black adipose fin. However, B. munduruku can be readily distinguished from B. inpai by the uniform
color pattern on the posterior portion of the body side (vs. a dark stripe on the posterior portion of the body side).
Although Bryconops inpai was described from igarap Barro Branco in the lower portion of the Negro River,
this species also occur along the middle and upper Negro River in Brazil, and in the Casiquiare River in Venezuela
(Lima et al. 2003). Analysis of Bryconops inpai holdings deposited at the Instituto Nacional de Pesquisas da
Amaznia (INPA 31573) allowed us to extend the distributional range of this species to the Trombetas River, near
Oriximin, Par State, Brazil.
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Bryconops munduruku is the fourth species of the genus described from the Tapajs River. Its congeners
described from the Tapajs River are: B. durbini, B. gracilis, and B. transitoria. Even though the new species can
be readily diagnosed among it congeners in the Tapajs River, the validity of B. transitoria and B. gracilis needs
further investigation. Steindachner (1915) described B. transitoria and stressed its similarity with B. melanurus,
but distinguished them based on morphometric accounts, and by the number of anal-fin rays (2327 vs. 2829,
respectively). Bryconops gracilis was described based on a sole specimen, which was also recognized as similar to
B. melanurus. We were not able to find any record of these two species in the Tapajs River.

Comparative material examined

Bryconops alburnoides: Brazil. INPA 12316 (1, 107.5 mm SL), Tocantins River; INPA 30701 (1, 145.7 mm SL),
Ja River; INPA 37794 (1, 81.1 mm SL), Negro River; INPA 36079 (6, 112.3115.1 mm SL), Branco River.
Bryconops caudomaculatus. Brazil. INPA 12397 (4, 69.377.1 mm SL), Trombetas River; INPA 12446 (2, 70.0
73.0 mm SL), Jamari River. Bryconops durbini. Brazil. UFOPA-I-00337 (109, 18.261.0 mm SL), Lago Verde,
Tapajs River. Bryconops aff. magoi: INPA 35801 (28, 27.138.0 mm SL), Purus River. Bryconops inpai: INPA
13249 (3, 63.272.9 mm SL), Urubu River; INPA 29524 (1, 66.7 mm SL), Trombetas River; INPA 31573 (1, 59.3
mm SL), Aneb River. Bryconops cf. vibex: INPA19638 (3, 49.866.3 mm SL), Negro River; INPA 19634 (6,
53.883.5 mm SL), Negro River; INPA 19636 (7, 31.379.6 mm SL), Negro River. Bryconops giacopinii: INPA
32636 (7, 57.167.7 mm SL), Negro River. Bryconops melanurus: Brazil. INPA 12443 (3, 91.191.9 mm SL),
Caraip River; INPA 26656 (1, 109.0 mm SL), Madeira River; INPA 38579 (3, 67.381.4 mm SL), Madeira River;
UFOPA-I-00336 (16, 84.1109.7 mm SL), Moju River; UFOPA-I-00338 (11, 49.178.0 mm SL), Curu River.

Acknowledgements
The authors are indebted to the following: Instituto Chico Mendes de Conservao da Biodiversidade (ICMBio) for
authorization to collect fishes (number 35649-2) in the Floresta Nacional do Tapajs; Programa de Pesquisas em
Biodiversidade (PPBio/Amaznia Oriental), and Universidade Federal do Oeste do Par (UFOPA) for financial
and logistical support. We thank David de Santana for suggestions to the manuscript; Srgio R. Oliveira for helping
in the field and to Coordenao de Aperfeioamento de Pessoal de Nvel Superior (CAPES) for the fellowship
granted to Crlison Silva-Oliveira.

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