New Species of Hemigrammocharax (Characiformes: Distichodontidae) from
the Northwestern Congo River Basin
Fernando C. Jerep1 and Richard P. Vari1
Copeia 2013, No. 1, 31–37
New Species of Hemigrammocharax (Characiformes: Distichodontidae) from
the Northwestern Congo River Basin
Fernando C. Jerep1 and Richard P. Vari1
Hemigrammocharax rubensteini is described from the northwestern Congo River basin in the Lékoli River of the Likouala
River system, Republic of the Congo. The species differs from congeners in the combination of pigmentation pattern,
meristics, and position of the dorsal fin. The presence of a well-developed pseudotympanum, a feature not previously
reported in the Distichodontidae, is reported for H. rubensteini and other members of Hemigrammocharax and
Nannocharax.
S
PECIES of the genus Hemigrammocharax inhabit the
streams and rivers of major portions of sub-Saharan
Africa including the Congo, Cunene, Okavango, and
Zambezi river basins (Poll, 1973; Daget and Gosse, 1984).
Hemigrammocharax was proposed by Pellegrin (1923) to host
Nannocharax ocellicauda, a species which although sharing
the other diagnostic features of the species then assigned to
Nannocharax, nonetheless lacked poring of the lateral line
along the posterior portion of the body contra the
completely pored lateral line in the other species of
Nannocharax. As now defined, Hemigrammocharax consists
of nine species diagnosed among the genera of the
Distichodontidae by the presence of a single row of teeth
on the dentary and premaxilla and incomplete poring of the
lateral line. Hemigrammocharax is hypothesized to form a
natural lineage with Nannocharax inside the family Distichodontidae based on the common possession in those
genera of a series of characters involving multiple body
systems (Vari, 1979:331); however, the monophyly of
Hemigrammocharax has yet to be assessed and indeed has
been questioned (Vari and Géry, 1981:1082). We describe a
new species from the Likouala River basin, a tributary on the
Congo River in the northwestern portion of the basin that
conforms to the present concept of Hemigrammocharax and
is assigned to that genus pending the results of ongoing
phylogenetic analyses.
MATERIALS AND METHODS
Measurements and counts conform to those used by Vari
and Ferraris (2004) and Dunz and Schliewen (2009).
Measurements were taken point-to-point on the left side of
the specimens under a microscope using a digital caliper
with a precision of 0.1 mm. Body measurements are given as
a percentage of the standard length (SL), except for subunits
of the head which are presented as percentages of the head
length (HL). In the description, the frequency of a value is
indicated by the number of specimens in parentheses. The
value for the holotype is presented in square brackets.
Details of dentition, infraorbitals, gill rakers, supraneurals,
and scale ctenii were taken from two cleared-and-stained
specimens prepared according to Taylor and Van Dyke
(1985). Vertebral count was taken from cleared-and-stained
specimens and radiographs and includes the four vertebrae
of the Weberian apparatus and the terminal centrum.
Presence of the pseudotympanum was verified on alcoholpreserved material by incident and/or transmitted light
1
under a microscope with pseudotympanum morphology
assessed by removal of the overlying skin and adipose tissue
of an alcohol-preserved specimen. Muscular nomenclature
follows Winterbottom (1974). Museum abbreviations follow
the listing in http://www.asih.org/node/204.
Hemigrammocharax rubensteini, new species
Figure 1, Table 1
Holotype.—CU 89502, 23.6 mm SL, Republic of the Congo,
Cuvette-Ouest, Likouala River Drainage, Lékoli River at
Mboko dock, Odzala National Park, 0u37930N, 14u549270E,
J. P. Friel, S. Lavoué, J. P. Sullivan, 6 August 2002.
Paratypes.—All Republic of the Congo, Cuvette-Ouest: CU
97248, 8, 6 alc, 20.0–25.8 mm SL, 2 CS, 21.3–21.9 mm SL;
AMNH 255311, 2, 20.1–21.9 mm SL; MRAC B2-05-P-1-2, 2,
20.0–22.8 mm SL; USNM 405306, 2, 20.3–22.1 mm SL,
captured with holotype. CU 89498, 6, 20.8–24.6 mm SL,
Likouala River drainage, small creek draining into Mambili
River, Odzala National Park, J. P. Friel, S. Lavoué, J. P.
Sullivan, 27 August 2002. CU 89499, 2, 19.7–19.8 mm SL,
Likouala River drainage, Lékoli River, Odzala National Park,
0u369510N, 14u569110E, J. P. Friel, S. Lavoué, J. P. Sullivan, 8
August 2002. CU 89500, 4, 19.8–25.4 mm SL, Likouala River
drainage, Pandaka River, Odzala National Park, 0u379260N,
14u559350E, J. P. Friel, S. Lavoué, J. P. Sullivan, 12 August
2002. CU 89501, 6, 20.1–24.1 mm SL, Likouala River
Drainage, Lékoli River at Mboko dock, Odzala National
Park, 0u37930N, 14u549270E, J. P. Friel, S. Lavoué, J. P.
Sullivan, 13 August 2002. CU 97249, 1 (meristics and
morphometrics not taken), 18.2 mm SL, Likouala River
Drainage, Lékoli River, Odzala National Park, 0u369510N,
14u569110E, J. P. Friel, S. Lavoué, J. P. Sullivan, 8 August
2002.
Diagnosis.—Hemigrammocharax rubensteini is distinguished
from all congeners, except H. lineostriatus, by the presence of
a dark, typically continuous, longitudinal band extending
from the posterodorsal margin of the opercle to the dark
spot overlying the caudal peduncle (versus a body pigmentation pattern consisting of several vertical bars in H.
angolensis, H. machadoi, H. minutus, H. monardi, H. multifasciatus, and H. wittei or lacking a dark longitudinal band in
H. ocellicauda and H. uniocellatus). In specimens of H.
rubensteini in which the longitudinal band is variably
Division of Fishes, Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C.; E-mail:
(FCJ) fjerep@gmail.com; and (RPV) varir@si.edu. Send reprint requests to RPV.
Submitted: 8 May 2012. Accepted: 24 September 2012. Associate Editor: D. Buth.
F 2013 by the American Society of Ichthyologists and Herpetologists
DOI: 10.1643/CI-12-059
32
Copeia 2013, No. 1
Fig. 1. Hemigrammocharax rubensteini, CU 89502, 23.6 mm SL, holotype; Africa, Republic of the Congo, Cuvette-Ouest, Likouala River drainage,
Lékoli River at Mboko dock, Odzala National Park.
interrupted, the pigmentation pattern somewhat resembles
that of H. lineostriatus, in which the continuous dark
longitudinal line has several short vertical bars along its
length. Hemigrammocharax rubensteini differs from H. lineostriatus in having the dorsal-fin origin located anterior to
the vertical through the pelvic-fin origin (versus the dorsalfin origin located posterior to the vertical through the
pelvic-fin insertion), 18 to 30 perforated scales along the
lateral line (versus 6 to 9), and 12 to 14 branched dorsal-fin
rays (versus 9 to 11). Hemigrammocharax rubensteini has a
variably discontinuous lateral line which in some instances
has only a few unpored scales and thus approximates the
completely pored lateral line characteristic of the species of
Nannocharax. The new species differs from the member of
that genus with a continuous dark midlateral longitudinal
band both in the incompletely pored lateral line (versus
completely pored) and the lower number of total scales
along that series (35–38 versus 39 or greater in N. ansorgii, N.
latifasciatus, N. lineomaculatus, N. parvus, and N. usongo).
Description.—Morphometric data presented in Table 1. Small
body size with maximum length 25.8 mm SL. Body slightly
Table 1. Morphometrics of Hemigrammocharax rubensteini, New Species. Range, mean, and standard deviation (SD) include values for
the holotype.
Holotype
n
Range
Mean
SD
Standard length
23.6
33
19.7–25.8
21.7
1.7
Percentage of SL
Head length
Predorsal length
Prepectoral length
Prepelvic length
Preanal length
Head width
Body width
Head depth
Body depth
Dorsal-fin length
Pectoral-fin length
Pelvic-fin length
Anal-fin length
Caudal-peduncle length
Caudal-peduncle depth
32.3
48.5
31.8
51.4
78.2
16.3
13.1
21.7
29.3
24.9
17.9
19.4
15.1
14.1
12.3
33
33
33
33
33
33
33
33
33
33
33
33
33
33
33
30.0–34.4
47.5–51.4
30.1–34.9
49.1–54.3
76.1–80.5
14.4–22.2
9.9–14.6
20.9–23.3
24.5–29.5
16.4–29.4
13.6–18.6
18.0–21.5
12.3–17.9
11.2–15.3
11.8–13.1
32.1
49.6
32.5
52.1
78.4
15.6
12.2
22.0
27.2
22.0
16.0
19.6
14.6
13.4
12.6
1.0
0.9
1.0
1.0
1.1
1.6
1.1
0.6
1.1
2.9
1.3
1.0
1.2
0.9
0.3
Percentage of HL
Eye horizontal diameter
Snout length
Upper-jaw length
Interorbital width
40.0
26.6
18.7
23.9
33
33
33
33
34.3–41.6
24.0–32.0
17.8–23.4
20.5–25.8
38.3
28.5
19.9
23.7
1.7
1.8
1.2
1.2
Jerep and Vari—Hemigrammocharax from Congo
elongated and laterally compressed (Fig. 1). Dorsal profile
convex from tip of snout to vertical situated slightly anterior
to anterior nares, straight to slightly convex from that point
to tip of supraoccipital spine. Dorsal profile convex from tip
of supraoccipital spine to dorsal-fin origin; straight to
slightly convex along dorsal-fin base; straight from insertion
of last dorsal-fin ray to posterior terminus of adipose-fin
base; and straight to slightly concave along caudal peduncle.
Ventral profile of head convex along anterior portion of
dentary; slightly convex from that point to isthmus. Ventral
profile of body slightly convex from isthmus to pelvic-fin
insertion; straight to slightly concave from pelvic-fin
insertion to vertical situated anterior to urogenital papillae;
concave from that point to anal-fin origin; slightly convex
to straight along anal-fin base; and straight to slightly
concave along caudal peduncle. Ventral surface of body
flattened between pelvic-fin insertions.
Head pointed in lateral profile with snout rounded.
Mouth subterminal; located posteroventral of tip of snout.
Premaxilla bearing one tooth row consisting of 6 elongate,
bicuspid, slightly distally expanded teeth. Teeth on premaxilla gradually decreasing in size posteriorly; posteriormost
tooth approximately one-half size of tooth proximate to
symphysis. Contralateral premaxillae attached medially
through syndesmotic articulation. Dentary bearing one
tooth row consisting of 6 elongate, bicuspid, slightly distally
expanded teeth gradually decreasing in size posteriorly, with
posteriormost tooth approximately one-half size of symphyseal tooth. Contralateral dentaries medially attached
through syndesmotic immovable articulation. Dentary
movably articulated posteriorly with anterodorsal region of
anguloarticular and lacking laterosensory canal segment.
Maxilla toothless, flat and expanded posteriorly. Ventral
margin of maxilla straight and posterior border rounded.
Maxilla extending posteriorly to vertical through anterior
margin of anterior nares. Posterior portion of maxilla lying
medial to anterior portion of first infraorbital when mouth
closed. Anterior naris rounded, separated posteriorly from
triangular posterior naris by skin flap. Eye laterally situated
with pupil ovoid and slightly extended anteriorly. Infraorbital series with infraorbitals 1 to 3 well developed,
infraorbitals 4 and 5 absent, and infraorbital 6 limited to
ossified laterosensory canal segment.
Dorsal-fin origin located slightly anterior to middle of SL
and anterior to vertical through pelvic-fin insertion. Distal
profile of fin straight to slightly convex. Most branched
dorsal-fin rays of same length other than few posteriormost
rays progressively decreasing in size. Unbranched dorsal-fin
rays ii (1), iii (17), iv (14), or v (1); branched rays 12(1), 13
(11), or 14 (21) [iii,14]. Pectoral fin pointed in profile. Tip of
pectoral fin falling short of pelvic-fin insertion. Unbranched
pectoral-fin rays i (33); branched rays 8 (18), 9 (14), or 10 (1)
[i,9]. Distal profile of pelvic fin straight to slightly convex.
Tip of fin falling short of anal fin. Pelvic-fin rays i,7,i (33)
[i,7,i]. Adipose fin well developed with origin situated along
vertical slightly anterior to base of last anal-fin ray. Distal
profile of anal fin straight to slightly convex. Unbranched
anal-fin rays i (1), ii (21), or iii (11); branched rays 7 (7) or 8
(26) [iii,8]. Forked caudal-fin lobes short, distally rounded,
and of similar size. Caudal-fin rays i,17,i (33) [i,17,i].
Scales ctenoid (sensu Roberts, 1993); covering all of body.
Lateral-line scale at vertical through dorsal-fin origin
bearing 24 ctenii formed of independent ossifications along
posterior margin of scale. Scales on humeral region and
33
Fig. 2. Pseudotympanum of Hemigrammocharax rubensteini, CU
89498, 23.7 mm SL, left side, anterior to left. Overlying skin and
adipose tissue removed. Narrow white band horizontally crossing
pseudotympanum is lateral-line nerve. Arrow indicates first pleural rib.
dorsal portion of body immediately posterior to head
smaller than remaining body scales. Lateral line straight,
but incompletely pored with 35 (6), 36 (11), 37 (13), or 38
(3) [36] scales; last one or two scales extending beyond point
of flexure along posterior margin of hypural plate. Pored
scales 18 (1), 19 (1), 20 (1), 21 (4), 22 (3), 23 (8), 24 (5), 25
(2), 26 (3), 27 (4), or 30 (1) [23]. Pored portion of lateral line
discontinuous in five specimens with 1 (2), 2 (2), or 3 (1)
sequential non-pored scales at positions 16, 17, 18, 19, 20,
23, 24, or 26. Scale rows between dorsal-fin origin and lateral
line 5 (33) [5]. Scale rows between pelvic-fin origin and
lateral line 4 (15), 4.5 (17), or 5 (1) [4]. Middorsal scales to
dorsal-fin origin 8 (2), 9 (8), 10 (16), 11 (6), or 12 (1) [10].
Scales from posterior terminus of dorsal-fin base to adipose
fin 8 (15), 9 (16), or 10 (2) [9]. Scales between anus and analfin origin 2 (11) or 3 (22) [3]. Scale rows around caudal
peduncle 12 (33) [12]. First gill arch with 11 or 12 gill rakers;
branchiostegal rays 4; supraneurals 5; vertebrae 35(1), 36
(15), or 37(2).
Well developed pseudotympanum composed of three
muscular hiatuses delimited medially by anterior swimbladder chamber and laterally by adipose tissue and skin
(Fig. 2). Anterior muscular hiatus triangular; bordered
dorsally by lateralis superficialis muscle, anteroventrally by
obliquus superioris muscle, and posteriorly by anterior margin
of first pleural rib. Middle hiatus pentagonal and margined
anteriorly and posteriorly by first and second pleural ribs,
respectively, dorsally by lateralis superficialis muscle, anteroventrally by obliquus superioris muscle, and posteroventrally
by obliquus inferioris muscle. Posterior hiatus overall ovoid;
delimited dorsally by lateralis superficialis muscle, anteriorly
by posterior surface of second pleural rib, anteroventrally by
obliquus superioris muscle, posteroventrally by obliquus
inferioris muscle, and posterodorsally by obliquus superioris
muscle.
Coloration in alcohol.—Overall color of head and body
yellowish; darker dorsally (Fig. 1). Head with darker pigmentation most intense over fontanels, posterior portion of
cranium and opercular region due to higher concentration
of melanophores. Band of dark pigmentation extending
34
Copeia 2013, No. 1
Fig. 3. Life coloration of Hemigrammocharax rubensteini, Likouala River drainage, Republic of the Congo. Photo: J. P. Sullivan.
from tip of snout and upper jaw to anterior margin of eye.
Scale pigmentation more intense over distal region of
exposed portion of scales resulting in overall reticulated
pattern. Region of anus and urogenital papillae dark brown
with pigmentation sometimes extending over anterior
portion of anal-fin base. Dark brown longitudinal band
extending along lateral surface of body from posterodorsal
margin of opercle to dark caudal-peduncle spot. Anterior
portion of longitudinal band darker and narrower than
generally fainter posterior portion. Posterior portion approximately two scales high. Longitudinal band divided in
some individuals to varying degrees into seven or eight dark
blotches. Spot on caudal peduncle dark brown, rounded and
extending onto bases of middle caudal-fin rays but falling
short of dorsal and ventral margins of caudal peduncle.
Dorsal, pectoral, and caudal fins with dark pigmentation
along margins of rays but otherwise hyaline. Pelvic and anal
fins with dark pigmentation densely scattered over interradial membrane.
Coloration in life.—Melanophores on head, body, and fins as
in alcohol-preserved specimens (Fig. 3). Dorsal portion of
head olive brown and ventral region white. Eye and
opercular region brownish with iridescent copper overlay.
Dorsal and posterior regions of body olive brown. Abdominal region white with few melanophores and fainter
reticulation pattern than dorsal portion of body. Caudal
spot round, black, and extending slightly onto bases of
middle caudal-fin rays. Base of dorsal and ventral caudal-fin
lobes white with yellowish tint. Dorsal fin slightly yellow.
Pelvic, anal, and caudal fins hyaline overall other than for
scattered melanophores. Pectoral fin hyaline.
Distribution.—Hemigrammocharax rubensteini is currently
known from small tributaries of the Likouala River within
the Odzala–Kokoua National Park, Republic of the Congo, in
the northwestern portion of the Congo River basin (Fig. 4).
Etymology.—The species name, rubensteini, is in recognition
of David Rubenstein who generously funded the Rubenstein
Fellowships of the Encyclopedia of Life at the National
Museum of Natural History.
Fig. 4. Map of central and northwestern portion of Congo River basin
showing type locality of Hemigrammocharax rubensteini in Likouala
River drainage. Dashed line represents borders of Republic of the Congo.
Remarks.—Hemigrammocharax rubensteini fits the definition
of the genus within the Distichodontidae in the combination of the presence of a single row of teeth on the jaws and
an incomplete lateral line. Monophyly of the Hemigrammocharax–Nannocharax lineage is well supported by multiple
morphological synapomorphies and retrieved in the molecular analysis by Calcagnotto et al. (2005). The monophyly of
each genus is, however, questionable (Vari, 1979; Vari and
Géry, 1981; Coenen and Teugels, 1989; Vari and Ferraris,
2004). Hemigrammocharax is now discriminated from Nannocharax in the incomplete versus complete poring of the
lateral line, respectively; however, as discussed by Vari and
Géry (1981), the distribution of these conditions is incongruent with that of several other synapomorphies proposed
by Vari (1979). Degree of lateral-line poring is also variable
Jerep and Vari—Hemigrammocharax from Congo
35
in some species of those genera (e.g., Hemigrammocharax
multifasciatus, Nannocharax ansorgii, and N. maculicauda) and
other species in the Characiformes (Roberts, 1967; Jubb and
Gaigher, 1971; Vari and Géry, 1981), highlighting the
possibility of independent reduction in the extent of poring.
Hemigrammocharax rubensteini possesses 11 of the 12
synapomorphies proposed by Vari (1979:331) for the
Nannocharax–Hemigrammocharax clade, lacking only the
vertical expansion of the posterior strut of the lateral
ethmoid contacting the orbitosphenoid (Vari, 1979:fig. 7,
synapomorphy 101). Hemigrammocharax rubensteini largely
lacks any ossified process or expansion on the posterior
surface of the lateral ethmoid with a resultant lack of an
articulation with the orbitosphenoid, although a narrow,
poorly developed, posterior bony projection occurs on the
left lateral ethmoid of one cleared-and-stained specimen in a
location homologous with the posterior process found in
most of the Distichodontidae. The lack of direct contact
between the lateral ethmoid and the orbitosphenoid was
observed by Vari (1979) in the distichodontid genera
Nannaethiops and Neolebias, a hypothesized secondary loss
within the context of the hypothesis of phylogenetic
relationships within the Distichodontidae. We similarly
hypothesize that the absence of the same posterior process
in Hemigrammocharax rubensteini is a secondary loss under
the phylogenetic hypothesis proposed by Vari (1979).
The description of Hemigrammocharax rubensteini increases
to ten the total number of valid species in the genus (Daget
and Gosse, 1984; Coenen and Teugels, 1989). Members of
Hemigrammocharax are widely distributed in the Congo,
Cunene, Okavango, and Zambezi river basins but with 80%
of the species occurring in the Congo basin: H. angolensis,
H. lineostriatus, H. minutus, H. multifasciatus (also inhabiting
the Zambezi System), H. ocellicauda, H. rubensteini, H.
uniocellatus, and H. wittei (Daget and Gosse, 1984; Poll,
1973). Among these, only H. uniocellatus is sympatric and
syntopic with H. rubensteini in the Lékoli River.
evolutionary lineages (Malabarba, 1998) among South
American characiforms; however, very little is known about
the presence and morphology of this structure among
African characiforms. Within the African family Alestidae,
a pseudotympanum occurs only in Lepidarchus, which has a
triangular hiatus in the musculature anterior to the first
pleural rib (Zanata and Vari, 2005:88). The more complex
pseudotympanum in Hemigrammocharax rubensteini spans
more than two ribs with only the relatively small component anterior to the first pleural rib positionally comparable
to that of Lepidarchus. That opening is complemented by a
second portion between the first and second pleural ribs and
a third component posterior to the second pleural rib. An
extension of the hiatus to between the first and second ribs
comparable to that in Hemigrammocharax rubensteini was
described by Weitzman and Malabarba (1999:16) as a
synapomorphy for Spintherobolus in the Neotropical characiform subfamily Cheirodontinae. This is an obvious convergence given the phylogenetic distance between these
genera. The species of Spintherobolus additionally lack the
muscular hiatus posterior to the second pleural rib found in
H. rubensteini.
We observed forms of the pseudotympanum similar to
that occurring in Hemigrammocharax rubensteini in H.
machadoi, H. multifasciatus, H. ocellicauda, and H. uniocellatus. Among members of Nannocharax, a pseudotympanum
was observed in N. ansorgii, N. brevis, N. elongatus, N.
fasciatus, N. intermedius, N. latifasciatus, N. lineomaculatus,
N. macropterus, N. maculicauda, N. niloticus, N. occidentalis, N.
parvus, N. procatopus, N. pteron, N. reidi, N. rubrolabiatus, N.
schoutedeni, N. seyboldi, N. taenia, and N. usongo. Although
not assessed in this study, it is probable that the remaining
species of both genera possess a reduction or gap in the
musculature in the pseudotympanum area. An evaluation of
the morphological variation of the pseudotympanum and
its phylogenetic significance in the Hemigrammocharax–
Nannocharax clade is beyond the scope of this study.
Pseudotympanum.—As mentioned above, Hemigrammocharax
rubensteini is characterized by a well developed muscular
hiatus in the humeral region, an opening referred to as the
pseudotympanum. Such reductions or eliminations of
portions of the body musculature lateral to the anterior
portion of the swim bladder likely facilitate sound transmission to the anterior chamber of the swimbladder with sound
vibrations then conveyed to the inner ear via the Weberian
apparatus (Malabarba, 1998). Within the Characiformes, a
pseudotympanum has been documented in multiple Neotropical taxa including some species of Characidium, family
Crenuchidae (da Graça and Pavanelli, 2008; Silveira et al.,
2008), and within the Characidae in some members of the
subfamilies Aphyoditeinae (Aphyocharacidium, Microschemobrycon, and Tyttobrycon; Géry, 1973), Characinae (Acestrocephalus, Charax, Galeocharax, Phenacogaster, and Roeboides;
Malabarba and Lucena, 1995; Lucena, 2003, 2007; Menezes,
2006; Lucena and Malabarba, 2010), across the subfamily
Cheirodontinae (Malabarba, 1998; Weitzman and Malabarba,
1999; Bührnheim et al., 2008), and various incertae sedis
species (Hyphessobrycon elachys, Leptagoniates pi, Paracheirodon
axelrodi, Psellogrammus kennedyi, Rhoadsia altipinna, Tetragonopterus chalceus, and some representatives of the rosy tetra
clade; Malabarba, 1998; Mirande, 2010).
Details of pseudotympanum form and position have been
used to diagnose species (Menezes, 2006) and define
MATERIAL EXAMINED
Hemigrammocharax angolensis: MRAC P.158822, paratypes, 3,
25.1–31.9 mm SL, Angola, Luita River, Xa-Ua, poste de Cuilo.
Hemigrammocharax lineostriatus: MRAC P.159945–159949,
paratypes, 5, 17.6–19.4 mm SL, Angola, mare Tchifuca.
Hemigrammocharax machadoi: AMNH 215607, 10, 20.5–
21.8 mm SL, Zambia, Western Province, Kataba stream on
Mongu-Senanga highway; MRAC P.159969, paratypes, 2,
20.5–20.7 mm SL, Angola, Longa River, tributary of Luena
River, Nharicumbi village.
Hemigrammocharax multifasciatus: AMNH 215614, 2, 31.2–
31.5 mm SL, Zambia, Western Province, Litoya stream on
Mongu-Senanga highway; AMNH 217445, 4, 26.5–33.0 mm
SL, Botswana, Ngamiland, Okavango River delta, Thamalakane River at Matlapenang bridge.
Hemigrammocharax ocellicauda: CU 87637, 30, 22.3–30.9 mm
SL, Republic of the Congo, Cuvette-Ouest, Likouala River
Drainage, Lékoli River, Odzala National Park; CU 89163, 1,
24.2 mm SL, Republic of the Congo, Cuvette-Ouest,
Likouala River Drainage, Lékoli River, Odzala National Park.
Hemigrammocharax uniocellatus: CU 89494, 7, 23.6–26.1 mm
SL, Republic of the Congo, Cuvette-Ouest, Likouala River
36
Drainage, Lékoli River, Odzala National Park; CU 89495, 7,
20.2–26.9 mm SL, Republic of the Congo, Cuvette-Ouest,
Likouala River Drainage, small canal around island in Lékoli
River, Odzala National Park.
Hemigrammocharax wittei: MRAC P.36247–36311, cotypes,
65, 27.1–34.9 mm SL, Kando River de Tenke.
Nannocharax ansorgii: AMNH 228534, 3, 23.6–25.4 mm SL,
Central African Republic, Chaine Des Bongos, control zone
of Sangba, Bamingui river confluence with Ivingou River,
60 km downstream of Sangba base; NRM 38317, 9, 16.2–
21.2 mm SL, Gambia, MacCarthy Island, Gambia River
drainage, ca 2 km NW Bansang, Farida Creek.
Nannocharax brevis: CU 91688, 1, 38.3 mm SL, Central
African Republic, Basee-Kotto, Oubangui River drainage,
Oubangi River downriver from Mobaye; CU 91690, 3, 39.1–
50.7 mm SL, Central African Republic, Basee-Kotto, Oubangui River drainage, Oubangi River shoreline at uncompleted
bridge at Mobaye.
Nannocharax elongatus: CU 91667, 1, 50.6 mm SL, Central
African Republic, Basee-Kotto, Oubangui River drainage,
Oubangi River shoreline at uncompleted bridge at Mohaye.
Nannocharax fasciatus: USNM 303754, 5, 26.2–32.3 mm SL,
Cameroon, Southwest Province, Manyu, Cross System, main
Cross River downstream of Mamfe, Mamfe River junction
with Cross River; USNM 303756, 2, 36.0–41.3 mm SL,
Cameroon, Southwest Province, Manyu, Cross System, main
Cross River downstream of Mamfe, main Cross River ca
23 km below Mamfe.
Nannocharax intermedius: MCZ 32508, 1, 47.9 mm SL, Ja
River at Bitye; SU 15579, 4, 39.0–45.5 mm SL, Cameroon,
Menyoo River, Ntem River Drainage, Ambam.
Nannocharax latifasciatus: USNM 303996, 3, 32.8–49.5 mm
SL, Cameroon, Southwest Province, Manyu, Cross System,
main Cross River downstream of Mamfe, Northern Munaya
River junction with Cross River.
Nannocharax lineomaculatus: MHNG 2433.045, 10 of 16,
21.9–32.6 mm SL, Central African Republic, BaminguiBangoran, river in Gounda, along route Ndele-Birao.
Nannocharax macropterus: AMNH 247845, 1, 65.9 mm SL,
Democratic Republic of Congo, Kasai Occidental, Lulua,
Nsanga Nyembo; AMNH 247846, 2, 43.9–56.5 mm SL,
Democratic Republic of Congo, Kasai Occidental, Lulua,
Ntumba Shambuyi; MCZ 48055, 2, 35.9–37.6 mm SL,
Ghana, Ongwa, and Prah rivers at or near Twifu Prasu.
Nannocharax maculicauda: CU 80621, 1, 30.6 mm SL, Gabon,
Woleu-Ntem, Ngomo River where it crosses road from Oyem to
Bitam; USNM 224524, 3, 23.5–28.7 mm SL (1 CS), Gabon,
Ogooue Ivindo, still arm of upper Ivindo River opposite
Bourassie, Gabon, at juncture of Djouah and Karouaga rivers.
Nannocharax niloticus: AMNH 50826, 4, 29.3–37.7 mm SL,
Egypt, Nile River; FMNH 54288, 2, 36.8–36.9 mm SL, Egypt,
Nile at Luxor.
Nannocharax occidentalis: CU 91678, 2, 26.9–38.5 mm SL,
Central African Republic, Nana-Grébiz, Gribingui/Chari
drainage, Grinbingui River at Kanga Bandoro.
Copeia 2013, No. 1
Nannocharax parvus: MRAC 90-57-P-2159-161, 3, 31.8–
40.4 mm SL, Congo Brazzaville, Ngoombi River, tributary
of Dola River, Loc Dinga basin.
Nannocharax procatopus: MHNG 1248.26, 1, 58.2 mm SL,
Congo, Kalubadi, Lulua River.
Nannocharax pteron: ANSP 65560, holotype, 45.5 mm SL,
Central African Republic, Tomi River, tributary of Ubangi, at
Fort Sibut, Ubangi-Shari, French Equatorial Africa.
Nannocharax reidi: USNM 304046, holotype, 62.7 mm SL,
Cameroon, Cross River system, southern Munaya River draining
northern Korup, on Basep River at junction with Munaya River;
USNM 375193, paratypes, 16, 34.3–59.0 mm SL, 2 CS,
Cameroon, Cross River System, southern Munaya River draining
northern Korup, on Basep River at junction with Munaya River.
Nannocharax rubrolabiatus: MRAC 95-22-P-1-7, 7, 45.4–
59.8 mm SL, Cameroon, Mi River, tributary of Lom River,
Sanaga River basin, igoum.
Nannocharax schoutedeni: AMNH 241889, 5, 22.9–29.7 mm
SL, Democratic Republic of Congo, Salonga National Park,
Ifumu, Luilaka River just downstream from Monkoto;
AMNH 241891, 14, 14.4–39.3 mm SL, Democratic Republic
of Congo, Salonga National Park, Luilaka River, Monkoto.
Nannocharax seyboldi: USNM 118757, holotype, 32.9 mm SL,
Liberia, Bellyella; USNM 118758, paratypes, 3, 34.8–41.3 mm
SL, Liberia, Bromley.
Nannocharax taenia: CU 91682, 3, 29.6–31.2 mm SL, Central
African Republic, Ouaka, Ouaka/Oubangui drainage,
Mbourou River at AXMIN Ndassima mine camp; CU
91684, 1, 35.0 mm SL, Central African Republic, Ouaka,
Ouaka/Oubangui drainage, Baı̈dou River, above and below
pontoon crossing.
Nannocharax usongo: USNM 303795, 1, 27.7 mm SL,
Cameroon, Southwest Province, Manyu, Cross System, main
Cross River downstream of Mamfe, Mam River junction with
Cross River.
ACKNOWLEDGMENTS
This project was made possible by support from the National
Museum of Natural History, Smithsonian Institution and
the Herbert R. and Evelyn Axelrod Chair in the Division of
Fishes of the National Museum of Natural History. We thank
B. Brown (AMNH), J. Friel (CU), and E. Vreven and M.
Parrent (MRAC) for the loan of specimens. S. Raredon, J.
Clayton, and L. Palmer provided assistance at USNM.
Figure 1 was prepared by T. Griswold, Figure 2 by D.
Johnson, Figure 3 by J. Sullivan, and Figure 4 by M.
Tsuchiya.
LITERATURE CITED
Bührnheim, C. M., T. P. Carvalho, L. R. Malabarba, and
S. H. Weitzman. 2008. A new genus and species of
characid fish from the Amazon basin—the recognition of
a relictual lineage of characid fishes (Ostariophysi:
Cheirodontinae: Cheirodontini). Neotropical Ichthyology
6:663–678.
Calcagnotto, D., S. A. Schaefer, and R. DeSalle. 2005.
Relationships among characiform fishes inferred from
Jerep and Vari—Hemigrammocharax from Congo
analysis of nuclear and mitochondrial gene sequences.
Molecular Phylogenetics and Evolution 36:135–153.
Coenen, E. J., and G. G. Teugels. 1989. A new species of
Nannocharax (Pisces, Distichodontidae) from south-east
Nigeria and west Cameroun, with comments on the
taxonomic status of Hemigrammocharax polli Roman,
1966. Cybium 13:311–318.
da Graça, W. J., and C. S. Pavanelli. 2008. Characidium
heirmostigmata, a new characidiin fish (Characiformes:
Crenuchidae) from the upper rio Paraná basin, Brazil.
Neotropical Ichthyology 6:53–56.
Daget, J., and J. P. Gosse. 1984. Distichodontidae,
p. 124–211. In: Check-list of the Freshwater Fishes of
Africa. J. Daget, J. P. Gosse, and D. F. E. Thys van den
Audenarde (eds.). Museé Royal de l’Afrique Centrale,
Tervuren, Belgium and Office de la Recerche Scientifique
et Technique Outre-Mer, Paris.
Dunz, A. R., and U. K. Schliewen. 2009. Description of two
new species of Nannocharax Günther, 1867 (Teleostei:
Characiformes: Distichodontidae) from the Cross River,
Cameroon. Zootaxa 2028:1–19.
Géry, J. 1973. New and little-known Aphyoditeina (Pisces,
Characoidei) from the Amazon basin. Studies on Neotropical Fauna 8:81–137.
Jubb, R. A., and I. G. Gaigher. 1971. Check list of the fishes
of Botswana. Arnoldia 5:1–22.
Lucena, C. A. S. 2003. Revisão taxonômica e relações
filogenéticas das espécies de Roeboides grupo-microlepis
(Ostariophysi, Characiformes, Characidae). Iheringia,
Série Zoologia 93:283–308.
Lucena, C. A. S. 2007. Revisão taxonômica das espécies do
gênero Roeboides grupo-affinis (Ostariophysi, Characiformes, Characidae). Iheringia, Série Zoologia 97:117–136.
Lucena, Z. M. S., and L. R. Malabarba. 2010. Descrição de
nove espécies novas de Phenacogaster (Ostariophysi:
Characiformes: Characidae) e comentários sobre as demais
espécies do gênero. Zoologia 27:263–304.
Malabarba, L. R. 1998. Monophyly of the Cheirodontinae,
characters and major clades (Ostariophysi, Characidae),
p. 193–234. In: Phylogeny and Classification of Neotropical
Fishes. L. R. Malabarba, R. E. Reis, R. P. Vari, Z. M. S. Lucena,
and C. A. S. Lucena (eds.). Edipucrs, Porto Alegre, Brazil.
Malabarba, L. R., and Z. M. S. Lucena. 1995. Phenacogaster
jancupa, new species, with comments on the relationships
and a new diagnosis of the genus (Ostariophysi: Characidae). Ichthyological Exploration of Freshwaters 6:337–
344.
Menezes, N. A. 2006. Description of five new species of
Acestrocephalus Eigenmann and redescription of A. sardina
and A. boehlkei (Characiformes: Characidae). Neotropical
Ichthyology 4:385–400.
37
Mirande, J. M. 2010. Phylogeny of the family Characidae
(Teleostei: Characiformes): from characters to taxonomy.
Neotropical Ichthyology 8:385–568.
Pellegrin, J. 1923. Les poissons des eaux douces de l’Afrique
occidentale (du Sénégal au Niger)a. Gouvernement Général de l’Afrique Occidentale Française, Publications Du
Comité D’Etudes Historiques Et Scientifiques, Paris.1–373.
Poll, M. 1973. Nombre et distribution géographique des
poisons d’eau douce Africans. Bulletin du Muséum
National d’Histoire Naturelle 150:113–128.
Roberts, C. D. 1993. Comparative morphology of spined
scales and their phylogenetic significance in the Teleostei.
Bulletin of Marine Sciences 52:60–113.
Roberts, T. R. 1967. Virilia, a new genus of sexually
dimorphic characid fish from West Africa with comments
on characoids having an incomplete line. Stanford
Ichthyological Bulletin 18:251–258.
Silveira, L. G. G., F. Langeani, W. J. da Graça, C. S.
Pavanelli, and P. A. Buckup. 2008. Characidium xanthopterum (Ostariophysi: Characiformes: Crenuchidae): a new
species from the Central Brazilian Plateau. Neotropical
Ichthyology 6:169–174.
Taylor, W. R., and G. C. Van Dyke. 1985. Revised
procedures for staining and clearing small fishes and
other vertebrates for bone and cartilage study. Cybium
9:107–119.
Vari, R. 1979. Anatomy, relationships and classification of
the families Citharinidae and Distichodontidae (Pisces,
Characoidea). Bulletin of the British Museum (Natural
History), Zoology series 36:261–344.
Vari, R. P., and C. J. Ferraris, Jr. 2004. A new species of
Nannocharax (Characiformes: Distichodontidae) from Cameroon, with descriptions of contact organs and breeding
tubercles in the genus. Proceedings of the Biological
Society of Washington 117:551–563.
Vari, R. P., and J. Géry. 1981. Nannocharax maculicauda, a
new species of African characoid fish (Characoidea,
Distichodontidae) with comments on the genus Hemigrammocharax. Proceedings of the Biological Society of
Washington 94:1076–1084.
Weitzman, S. H., and L. R. Malabarba. 1999. Systematics of
Spintherobolus (Teleostei: Characidae: Cheirodontinae)
from eastern Brazil. Ichthyological Exploration of Freshwaters 10:1–43.
Winterbottom, R. 1974. A descriptive synonymy of the
striated muscles of the Teleostei. Proceedings of the
Academy of Natural Sciences of Philadelphia 125:
225–317.
Zanata, A. M., and R. P. Vari. 2005. The family Alestidae
(Ostariophysi, Characiformes): a phylogenetic analysis of
a trans-Atlantic clade. Zoological Journal of the Linnean
Society 145:1–144.