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A New Species of Hasemania from the Upper Rio Tapajo´s Drainage, Brazil
(Teleostei: Characiformes: Characidae)
VINICIUS A. BERTACO AND LUIZ R. MALABARBA
A new characid species, Hasemania nambiquara, is described from the upper Rio
Tapajo´s, Mato Grosso, Brazil. The new species is distinguished from all other
Hasemania by the unique presence of one vertically-elongate black humeral spot and
a broad horizontal black band ventral to midlateral line between humeral region and
caudal fin.
Uma nova espe´ cie de caracı´deo, Hasemania nambiquara e´ descrita para o alto Rio
Tapajo´s, Mato Grosso, Brasil. A nova espe´cie distingue-se das demais espe´cies de
Hasemania pela presenc¸ a de uma mancha umeral preta verticalmente alongada, e de
uma larga faixa longitudinal preta, ventral em relac¸a˜o a` linha me´dio-lateral do corpo,
da regia˜o umeral ate´ o pedu´ nculo caudal.
THE genus Hasemania was proposed by Ellis
(1911) to include three species differing
from other characids mainly in the absence of an
adipose fin. The presence of two series of teeth in
the premaxilla, the maxilla with few or no teeth
along its exposed anterior margin, the incom-
plete lateral line, and naked caudal fin led Ellis
(1911:148) to consider the new genus ‘‘like
Hyphessobrycon, but without an adipose’’.
Five valid species have been recognized in
Hasemania, four with a limited distribution in the
Rio Parana´(H.crenuchoides, upper Rio Parana´; H.
maxilaris and H. melanura, Rio Iguac¸u) and Rio
Sa˜o Francisco (H. nana) drainages, and one
species (H. hanseni) from an unknown locality in
the state of Goia´s, Brazil (Lima et al., 2003). We
herein describe a new species of Hasemania from
the upper Rio Juruena, a tributary of the upper
Rio Tapajo´s drainage, thereby extending the
distribution of the genus into the Amazon basin.
MATERIALS AND METHODS
Counts and measurements follow Fink and
Weitzman (1974), with the exception of the
number of scale rows below the lateral line,
which were counted from the scale row ventral to
lateral line to the scale row nearest the base of
the first pelvic-fin ray. Vertebrae, supraneurals,
gill-rakers, and procurrent caudal-fin ray counts
were taken from one cleared-and-stained speci-
men (CS) prepared according to the method of
Taylor and Van Dyke (1985). Values for the
holotype are indicated by an asterisk. Vertebral
counts include the four vertebrae integrated in
the Weberian apparatus and the terminal cen-
trum was counted as one vertebra. Scanning
electron micrographs (SEM) photos were taken
from the cleared-and-stained dissected specimen.
Measurements were taken point-to-point with
electronic calipers on the left side of specimens.
All measurements other than standard length
(SL) are expressed as a percentage of SL except
for subunits of the head, which are presented as
a percentage of head length (HL). Institutional
abbreviations follow Leviton et al. (1985). Com-
parisons to Hasemania maxilaris,H. melanura, and
H. crenuchoides were based on data presented in
the literature.
Hasemania nambiquara, new species
Figures 1–3, Table 1
Holotype.—MCP 38390, 27.0 mm SL, Brazil, Mato
Grosso, Comodoro, Rio Doze de Outubro on the
highway BR 364 between Comodoro and Vil-
hena, tributary of Rio Juruena, upper Rio
Tapajo´s drainage, 12u589390S60u009300W, 14 July
2004, R. E. Reis, P. A. Buckup, A. R. Cardoso, and
E. H. L. Pereira.
Paratypes.—ANSP 185229, 2, 19.1–21.5 mm SL;
MCP 38038, 4, 1 CS, 10.5–23.4 mm SL; MNRJ
29862, 2, 17.5–19.9 mm SL, collected with the
holotype.
Diagnosis.—Hasemania nambiquara is distin-
guished from all congeners by the unique
presence of a broad black band, located ventral
to midlateral line, and extending from the
humeral region to the caudal-fin, and by the
presence of a black vertically-elongate humeral
spot (Figs. 1, 2). Additional characters to distin-
guish H. nambiquara include a higher number of
maxillary teeth (Fig. 3) with more cusps (two or
three tri- to pentacuspid teeth vs. none, one, or
two conical or bicuspid teeth in H. hanseni,H.
maxilaris,H. melanura, and H. nana) and more
Copeia, 2007(2), pp. 350–354
#2007 by the American Society of Ichthyologists and Herpetologists
branched anal-fin rays (16–19 vs. 11–16 in H.
crenuchoides,H. melanura, and H. nana).
Description.—Morphometric data summarized in
Table 1. Body compressed and moderately deep;
greatest body depth located between tip of
supraoccipital spine and dorsal-fin origin. Dorsal
profile of body convex from snout to dorsal-fin
origin; posteroventrally slanted at dorsal-fin base;
straight from base of last dorsal-fin ray to caudal
peduncle. Ventral profile of head convex. Ven-
tral body profile slightly convex from pectoral-fin
insertion to anal-fin origin. Profile of body along
anal-fin base posterodorsally slanted. Caudal
peduncle deep, profile nearly straight to slightly
concave along dorsal and ventral margins.
Snout rounded from margin of upper lip to
vertical through anterior nostrils. Mouth termi-
nal. Posterior tip of maxilla at vertical through
middle of orbit; maxilla oriented at angle of
approximately 45uto longitudinal body axis.
Maxilla nearly straight on dorsal border; ante-
roventral border concave and posteroventral
border convex. Maxilla slightly deeper poster-
iorly.
Premaxilla with two tooth rows; outer row with
2–3 tricuspid teeth, central cusp largest (Fig. 3).
Inner row with five teeth, teeth gradually de-
creasing in length posteriorly, last tooth thinner
than other teeth. Teeth with 3–5 cusps and
central cusp longer and broader than lateral
cusps. Maxilla with two or three teeth, each with
3 to 5 cusps with central cusp largest. Five
anteriormost dentary teeth largest, with five
cusps, and followed first by somewhat smaller
tooth with 3–5 cusps, and then four or five
distinctly smaller teeth with 1–3 cusps; central
cusp in all teeth two or three times longer and
broader than remaining cusps. Cusp tips slightly
curved posteriorly and lingually.
Scales cycloid. Lateral line incomplete, pored
scales 6–7 (7*, mode 57, n58). Longitudinal
scale series including lateral line with 33–35 (35*,
mode 534, n58) scales. Scale rows between
Fig. 1. Hasemania nambiquara, MCP 38390, holotype, 27.0 mm SL.
Fig. 2. Hasemania nambiquara, MCP 38038, paratype, 17.4 mm SL.
BERTACO AND MALABARBA—NEW HASEMANIA FROM TAPAJO
´S DRAINAGE 351
dorsal-fin origin and lateral line 5–7 (7*, mode 5
5, n58); scale rows between lateral line and
pelvic-fin insertion 3–5 (4*, mode 54, n58).
Predorsal scales 11–12, arranged in regular series
(11*, mode 511, n57). Scales rows around
caudal peduncle 14. Scale sheath along anal-fin
base with 4–6 scales in single series extending
from base of first unbranched ray to base of
fourth to sixth branched rays.
Dorsal-fin rays ii,9 (n59); first unbranched ray
approximately one-half length of second un-
branched ray. Dorsal-fin origin posterior to mid-
dle of SL and posterior to vertical through pelvic-
fin insertion. Adipose fin absent. Anal-fin rays
iv,16–19 (18*, mode 517, n59). First un-
branched ray only apparent in cleared-and-stained
specimen. Longest branched anal-fin ray not
reaching origin of last anal-fin ray when fin is
depressed. Anal-fin origin at vertical between base
of seventh through ninth dorsal-fin rays. Pectoral-
fin rays i,9–10 (10*, mode 510, n58), tip of fin
almost reaching pelvic-fin insertion. Pelvic-fin rays
i,6–7 (7*, mode 57, n59); longest ray reaching
just to anal-fin origin. Pelvic-fin insertion located
anterior to vertical through dorsal-fin origin.
Caudal fin forked, with 19 principal rays (n5
9). Dorsal procurrent rays 10, and ventral procur-
rent rays 8 (n51). Precaudal vertebrae 15; caudal
vertebrae 18. Supraneurals 4. Gill-rakers on first
arch 6/10 (n51).
Coloration in alcohol.—Area immediately proxi-
mate to mid-dorsal line of head and body densely
pigmented except for light mid-dorsal line along
predorsal region. In small specimens (Fig. 2),
lower lip, upper portion of maxilla, first and
fourth infraorbitals, and upper portion of oper-
cle covered with dark chromatophores forming
a longitudinal black line across eye continuous
with longitudinal body stripe. Pigmentation
diffuse on head of adults (Fig. 1), not forming
a conspicuous dark line crossing eye. Scales on
midlateral surface of body with scattered dark
brown chromatophores. Black humeral spot
rounded in small specimens; vertically-elongate
and narrowing ventrally in larger specimens.
Spot located over second to fourth lateral-line
scales and extending over two horizontal series of
scales below lateral line. Wide, densely pigmen-
ted horizontal stripe located largely ventral to
midlateral line. Lateral body stripe extending
from humeral region to caudal-fin in small
specimens, but limited to the lateral area of
body above pelvic and anal fins in large speci-
mens. Region immediately above pelvic and anal
fins lighter; with few dark chromatophores on
abdomen. Middle caudal-fin rays densely pig-
mented, continuous with midlateral body stripe.
All fins dusky.
Sexual dimorphism.—The cleared-and-stained dis-
sected specimen (20.8 mm SL) was a female with
mature ovaries. The larger specimens in the type
series are consequently presumably mature;
however, no secondary sexual characters were
observed.
Ecological notes.—All specimens of Hasemania
nambiquara were collected along the margins of
Fig. 3. Scanning electronic micrograph of upper
and lower jaws (right side, lateral view) of Hasemania
nambiquara, MCP 38038, paratype, 20.8 mm SL.
Scale bar 0.5 mm.
TABLE 1. MORPHOMETRIC DATA OF HOLOTYPE (MCP
38390, H) AND PARATYPES (ANSP 185229, 2; MCP 38038,
2 of 4; MNRJ 29862, 2) OF Hasemania nambiquara. The
range includes the holotype.
Character H Range Mean
Standard length (mm) 27.0 17.5–27.0 21.3
Percents of standard length
Predorsal distance 56.7 54.3–57.4 55.9
Prepelvic distance 47.0 46.6–49.6 48.0
Prepectoral distance 27.3 27.2–29.3 28.1
Preanal distance 64.3 61.5–66.9 64.0
Depth at dorsal-fin origin 32.6 27.4–32.6 30.0
Caudal-peduncle depth 13.5 10.3–13.5 11.8
Caudal-peduncle length 13.0 12.3–14.4 12.8
Anal-fin base 26.7 24.1–28.0 25.7
Dorsal-fin length 34.4 22.9–34.4 28.0
Pelvic-fin length 17.0 13.6–17.2 15.5
Pectoral-fin length 20.7 17.4–20.7 18.6
Head length 26.9 27.9–28.7 27.6
Percents of head length
Snout length 22.8 19.3–22.8 20.9
Upper-jaw length 41.9 35.7–41.9 38.4
Orbital diameter 37.2 36.7–44.6 39.4
Interorbital width 29.7 26.0–31.6 29.3
352 COPEIA, 2007, NO. 2
semilentic stretches of a shallow, clear water
river, with sand and leaves on the bottom, and
a moderate amount of riparian vegetation.
Distribution.—Hasemania nambiquara is known
only from its type locality in the upper Rio
Tapajo´s drainage, Mato Grosso, Brazil.
Etymology.—The species name, nambiquara,isin
reference to the Nambiquara, an indigenous
group from the region of the upper Rio Tapajo´s
drainage, near the type locality. A noun in
apposition.
DISCUSSION
The new species is described herein in
Hasemania because it fits best in the traditional
diagnosis of the genus given by Ellis (1911) and
subsequent authors. The lack of an adipose fin is
unusual in the Characidae and was hypothesized
by Weitzman and Malabarba (1999) as indepen-
dently derived in Spintherobolus (Cheirodonti-
nae), Priocharax (Characinae), Iotabrycon,Tytto-
charax, and Xenurobrycon (Xenurobryconini), and
Coptobrycon,Grundulus,Hasemania, and Nemato-
brycon (incertae sedis characid genera). Hasemania
nambiquara lacks the synapomorphies proposed
for the Cheirodontinae or Xenurobryconini by
Malabarba (1998) and Weitzman and Fink
(1985), respectively. It similarly lacks the synapo-
morphies used by Lucena (1998) to group
Charax with eleven other genera in the Characi-
nae. The lack of an adipose fin in these three
groups may be correlated with miniaturization
(Weitzman and Malabarba, 1998), but Hasemania
nambiquara exceeds the size limits for miniatures
proposed by Weitzman and Vari (1988).
Regarding the remaining genera, Hasemania
nambiquara does not fit in the traditional di-
agnosis of Coptobrycon by lacking the very large,
pedunculate and distally expanded teeth of the
dentary and inner premaxillary tooth series; of
Grundulus bythepresenceoftwoseriesof
multicuspidate teeth in the premaxilla instead
of a single series of conical teeth; and of
Nematobrycon by the incompletely toothed maxilla
with a small number of maxillary teeth (2–3 vs. 7–
16; Weitzman and Fink, 1971).
Relationships of Hasemania nambiquara with
other Hasemania species remain to be investigat-
ed. The monophyly of Hasemania based solely on
the lack of an adipose fin has been questioned by
several authors (Bo¨hlke, 1958; Ge´ry, 1972; Weitz-
man and Malabarba, 1999) and more recently by
Lima and Gerhard (2001), who included a newly
described characid species lacking an adipose fin
in Hyphessobrycon instead of Hasemania due to an
overall body resemblance of the new species,
Hyphessobrycon negodagua, to several species of the
Hyphessobrycon. A similar procedure was adopted
previously by Ellis (1911), in the description of
Hyphessobrycon taurocephalus that lacks an adipose
fin. Hasemania nambiquara differs from H. taur-
ocephalus and H. negodagua in its color pattern,
and additionally from H. taurocephalus in the
number of branched anal-fin rays (16–19 vs. 12–
14, respectively).
MATERIAL EXAMINED
Coptobrycon bilineatus: MCP 39051, 10, 2 CS,
23.5–34.0 mm SL, Brazil, Sa˜o Paulo, Rio Itatinga,
Parque das Neblinas, Bertioga, 23u449580S,
46u099510W. Hasemania hanseni: MCP 23420, 8,
21.0–24.8 mm SL; MCP 27367, 6, 19.8–22.3 mm
SL, Brazil, Distrito Federal, stream on highway
BR 020 between the highway BR 251 and
Planaltina, 15u419060S, 47u399080W; MZUSP
35676, 10 of 268, 27.0–30.8 mm SL, Rio Pipiripau
near to Planaltina, 15u419130S, 47u399070W.
Hasemania nana: MZUSP 84034, 6, 13.4–
17.8 mm SL, Brazil, Bahia, Formosa do Rio
Preto, Rio Preto at Cacimbinhas, 11u089280S,
46u049010W; MZUSP 84072, 1, 16.5 mm SL,
Bahia, Formosa do Rio Preto, Rio Sapa˜o at
Breja˜o, Rio Preto drainage, 10u559390S,
45u419540W. Hasemania sp.: MZUSP 73136, 1,
30.7 mm SL, Brazil, Minas Gerais, Dores de
Ganha˜es, Rio Guanha˜es, Rio Doce drainage,
approximately 19u039S, 42u579W. Hyphessobrycon
negodagua: MCP 23432, 5 paratypes, 22.8–
28.2 mm SL, Brazil, Bahia, Iraquara, Rio Pra-
tinha, 12u219130S, 41u329510W. Hyphessobrycon
heterorhabdus: MCP 25577, 2, 30.4–35.6 mm SL,
Brazil, Mato Grosso, Aripuana˜, stream on road
between Vilhena and Aripuana˜, approximately
09u599S, 59u219W. Hyphessobrycon cf. herbertaxel-
rodi: MCP 30821, 26, 17.3–24.8 mm SL, Brazil,
Mato Grosso, Simione, stream on road MT 338,
Rio Xingu drainage, 11u479400S, 56u449090W.
Hyphessobrycon taurocephalus: CAS 22778, 5 para-
types, 29.9–42.7 mm SL, Brazil, Parana´, rio
Iguac¸u. Hyphessobrycon vilmae: MCP 30823, 9,
19.9–24.1 mm SL, Brazil, Mato Grosso, Sinop,
co´rrego Etne´ia at the road MT 423, 11u429210S,
55u179320W. Nematobrycon palmeri: CAS 70883, 43,
2 CS, 11.2–29.3 mm SL, Colombia, Departa-
mento Choco, Rı´o Condoto, tributary of Rı´o
San Juan; MHNG 2182.86, 7, 2 CS, 25.8–31.2 mm
SL, Colombia, aquarium center.
ACKNOWLEDGMENTS
We are grateful to M. Toledo-Piza, J. Serra, and
S. Weitzman for comments and suggestions on
the manuscript; to the Centro de Microscopia e
BERTACO AND MALABARBA—NEW HASEMANIA FROM TAPAJO
´S DRAINAGE 353
Microana´lises (CEMM), of the Pontifı´cia Uni-
versidade Cato´lica do Rio Grande do Sul
(PUCRS) for the SEM preparations, and to A.
Charcansky for preparing Figure 3. Thanks to D.
Catania, O. Oyakawa, and S. Fish-Muller for loan
of specimens. The specimens were collected
during an expedition organized by R. Reis and
sponsored by the All Catfish Species Inventory
(NSF-DEB 0315963). VAB is grateful for the
visiting support offered by the California Acade-
my of Sciences and is financially supported by the
Coordenac¸a˜o de Aperfeic¸oamento de Pessoal de
Nı´vel Superior (CAPES). Research was supported
by the Conselho Nacional de Desenvolvimento
Cientı´fico e Tecnolo´ gico (CNPq Proc. 476821/
2003-7; Proc. 478002/2006-8). Collecting permits
were provided by Instituto Brasileiro do Meio
Ambiente e dos Recursos Naturais Renova´veis—
IBAMA, Portaria 054/2004.
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(VAB, LRM) MUSEU DE CIE
ˆNCIAS E TECNOLOGIA,
PONTIFI
´CIA UNIVERSIDADE CATO
´LICA DO RIO
GRANDE DO SUL,AV.IPIRANGA 6681, CAIXA
POSTAL 1429, 90619-900 PORTO ALEGRE, RS,
BRAZIL;AND (LRM) DEPARTAMENTO DE ZOOLO-
GIA, IB, UNIVERSIDADE FEDERAL DO RIO GRANDE
DO SUL,AV.BENTO GONC¸ ALVES, 9500, 91501-970
PORTO ALEGRE, RS, BRAZIL. E-mail: (VAB)
ubertaco@pucrs.br; and (LRM) malabarb@
pucrs.br. Send reprint requests to VAB.
Submitted: 28 Dec. 2005. Accepted: 3 Dec.
2006. Section editor: J. W. Armbruster.
354 COPEIA, 2007, NO. 2