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Apistogramma eleutheria and Apistogramma flavipedunculata are described from the upper portion of two adjacent tributaries of the rio Curuá on the Serra do Cachimbo. Apistogramma eleutheria, new species, is isolated from the lower portion of rio Treze de Maio (a tributary of the middle rio Curuá), by a waterfall with same name. It is similar to several species of the A. regani species group, but differs by the coloration of brooding females. Brooding females of A. eleutheria have a very dark, blackish pigmentation on the median area of gular region occupying the ventral portion of opercular series and branchiostegal membranes, which is most similar to that found in females of A. kullanderi. In A. kullanderi, however, the dark gular pigmentation is more expansive, extending onto the ventral half of the head, and continuing posteriorly onto belly and along anal-fin base. Apistogramma flavipedunculata, new species, is only known from the rio Escorpião, and is probably isolated in the upper rio Curuá by a series of three waterfalls. It is distinguished by the unique color pattern of adult males, which have the posterior part of the flank and caudal peduncle dominated by a light yellow pigmentation, which results in a dark midlateral band fragmented posteriorly and dark bars masked or absent (though present in juveniles); also differing from all congeners except A. luelingi and A. eleutheria by having exclusively four anal-fin spines instead of three.
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Ichthyological Exploration of Freshwaters
An international journal for field-orientated ichthyology
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Ichthyol. Explor. Freshwaters, Vol. 27, No. 1
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Copyright © Verlag Dr. Friedrich Pfeil
Ichthyol. Explor. Freshwaters, Vol. 27, No. 1, pp. 81-95, 5 figs., 1 tab., April 2016
© 2016 by Verlag Dr. Friedrich Pfeil, München, Germany – ISSN 0936-9902
Apistogramma eleutheria and A. flavipedunculata,
two new species of dwarf cichlids
from the rio Curuá on Serra do Cachimbo, Brazil
(Teleostei: Cichlidae)
Henrique R. Varella* and Ricardo Britzke**
Apistogramma eleutheria and Apistogramma flavipedunculata are described from the upper portion of two adjacent
tributaries of the rio Curuá on the Serra do Cachimbo. Apistogramma eleutheria, new species, is isolated from the
lower portion of rio Treze de Maio (a tributary of the middle rio Curuá), by a waterfall with same name. It is
similar to several species of the A. regani species group, but differs by the coloration of brooding females. Brood-
ing females of A. eleutheria have a very dark, blackish pigmentation on the median area of gular region occupying
the ventral portion of opercular series and branchiostegal membranes, which is most similar to that found in
females of A. kullanderi. In A. kullanderi, however, the dark gular pigmentation is more expansive, extending onto
the ventral half of the head, and continuing posteriorly onto belly and along anal-fin base. Apistogramma flavipe-
dunculata, new species, is only known from the rio Escorpião, and is probably isolated in the upper rio Curuá by
a series of three waterfalls. It is distinguished by the unique color pattern of adult males, which have the poste-
rior part of the flank and caudal peduncle dominated by a light yellow pigmentation, which results in a dark
midlateral band fragmented posteriorly and dark bars masked or absent (though present in juveniles); also dif-
fering from all congeners except A. luelingi and A. eleutheria by having exclusively four anal-fin spines instead of
three.
* Instituto de Biociências da Universidade de São Paulo, Museu de Zoologia da Universidade de São Paulo,
Caixa Postal 42494, 04218-970 São Paulo, SP, Brazil. E-mail: hrvarella@usp.br
** Universidade Estadual Paulista, Instituto de Biociências, Departamento de Morfologia, Rubião Jr. s/n., 18618-
970 Botucatu, SP, Brazil; Universidad Técnica de Machala, Facultad de Ciencias Agropecuarias, Laboratorio
de Acuacultura, Km. 41
/
2 Vía Machala-Pasaje, E583, Machala, El Oro, Ecuador. E-mail: britzker@gmail.com
Introduction
Apistogramma Regan, 1913 and Crenicichla Heckel,
1840 are the most species-rich genera among
Neotropical cichlids, the former with 89 species
and the latter with 90 species (Kullander, 2003;
Kullander & Varella, 2015; Römer et al., 2015). The
geographic distribution of Apistogramma includes
much of tropical and subtropical South America,
but most of the species of the genus are thought
to have distributions restricted to a particular
region or hydrographic basin (Kullander, 1986).
Only few species of the genus have wide distri-
butions, as Apistogramma agassizi (Steindachner,
1875), A. bitaeniata Pellegrin, 1936, A. cacatuoides
Hoedeman, 1951, A. commbrae (Regan, 1906) and
82 Copyright © Verlag Dr. Friedrich Pfeil
Varella & Britzke: Apistogramma eleutheria and A. flavipedunculata
A. borellii (Regan, 1906). The highest diversity and
degree of endemism of Apistogramma is found
in the Western Amazon, notably in Peru (with
24 valid species, 21 of them endemic), in the río
Orinoco basin (19 endemic species) and in the rio
Negro basin (13 endemic species). The diversity
of Apistogramma in the Brazilian Shield remains
little studied with many putative undescribed
species indicated for some drainages (e. g. Kos-
lowski, 2002).
Apistogramma kullanderi Varella & Sabaj Pérez,
2014, from the upper rio Curuá on the Serra do
Cachimbo, is the only nominal species described
for the rio Xingu drainage. The Serra do Cachimbo
is the highest portion of the Brazilian Shield in
the Amazon basin that includes headwaters from
both middle Xingu (i. e., rio Curuá) and rio Tapajós
(rio Jamanxim and rio Teles Pires) river basins
(Birindelli et al., 2009). Inventories and taxonomic
studies of the region have shown high degrees
of endemism for various fishes. For instance, the
upper rio Curuá presents six endemic species de-
scribed only recently: Leporinus guttatus Birindelli
& Britski, 2009, Jupiaba kurua Birindelli et al., 2009,
Moenkhausia petymbuaba Lima & Birindelli, 2006,
Lebiasina marilynae Netto-Ferreira, 2012, L. melano-
guttata Netto-Ferreira, 2012 and Apistogramma
kullanderi Varella & Sabaj Pérez, 2014.
Varella & Sabaj Pérez (2014) mentioned two
undescribed species of Apistogramma, one syntopic
with A. kullanderi in the upper rio Curuá and
another from the rio Treze de Maio, a tributary
that joins the rio Curuá downstream of the major
waterfalls (Salto do Curuá) that mark its depar-
ture from the Serra do Cachimbo. The aim of this
paper is to provide formal descriptions of those
species and to discuss aspects of the diversity of
Apistogramma recently discovered on the Serra do
Cachimbo.
Material and methods
Measurements, counts and color pattern termi-
nology follow Kullander (1980, 1986); counts for
holotype are denoted by asterisks. Scale rows
are numbered according to Kullander (1990).
A half-scale row (1
/
2 scale) typically occurs adjacent
to a fin base and is formed by scales approxi-
mately half the size of a normal flank scale. The
correspondence between coloration and sex was
determined by examination of gonads or genital
papillae under stereomicroscope. Vertebral counts
include the last half-centrum and were taken from
radiographs made with the system Faxitron LX-60
(DC 12, version 1.0).
Institutional abbreviations are: ANSP, Aca-
demy of Natural Sciences of Philadelphia; LBP,
Laboratório de Biologia e Genética de Peixes,
Botucatu, São Paulo; MZUSP, Museu de Zoolo-
gia da Universidade de São Paulo; MCP, Museu
de Ciências e Tecnologia PUCRS, Porto Alegre;
MUSM, Museo de Historia Natural, Universidad
Nacional Mayor de San Marcos, Lima; and ROM,
Royal Ontario Museum, Toronto. Comparative
material examined is as listed by Varella & Sabaj
Pérez (2014) and Britzke et al. (2014), with addition
of the material listed at the end of this paper.
Results
Apistogramma eleutheria, new species
(Fig. 1)
Apistogramma sp. upper Treze de Maio: Varella
& Sabaj Pérez, 2014: 255.
Holotype. MZUSP 115293, 39.5 mm SL, adult
male; Brazil: Pará: Altamira: rio Xingu basin: rio
Iriri drainage: rio Treze de Maio, a tributary of
rio Curuá, upstream of PCH Salto Treze de Maio
dam; 8°45'06" S 55°02'05" W; A. Netto-Ferreira, J.
Birindelli, L. Sousa & P. Hollanda-Carvalho, 23
Jan 2009.
Paratypes. MZUSP 97095, 1 unsexed, 25.1 mm SL,
1 female, 38.9 mm SL; same locality as holotype;
J. Birindelli, A. Netto-Ferreira, L. Sousa, M. Sabaj
Pérez & N. Lujan, 22 Oct 2007. – MZUSP 101422, 2
unsexed or juveniles, 23.2-29.0 mm SL, 2 females,
31.3-40.1 mm SL; collected with holotype.
Diagnosis. Apistogramma eleutheria is similar in
general counts and body shape to several species
of the A. regani species group sensu Kullander
(1980), but differs from them by details of col-
oration of mature males and females. The color
pattern of mature females includes a very dark,
blackish pigmentation on the median area of gular
region, occupying the ventral portion of opercular
region and branchiostegal membranes, expressed
as a ventral continuation of the suborbital stripe
(Fig. 5a). Among species of the A. regani group,
only females of A. kullanderi show similar gular
pigmentation, but it differs from A. eleutheria by
Ichthyol. Explor. Freshwaters, Vol. 27, No. 1
83
Copyright © Verlag Dr. Friedrich Pfeil
having this dark pigmentation more expansive,
extending onto the ventral half of the head, and
continuing posteriorly onto the belly and along
the anal-fin base (Fig. 5b); females of A. eleutheria
have dark pigments on basal portion of the dorsal
fin discontinuous, forming blotches with paler
interspaces whereas females of A. kullanderi have
the basal portion of the dorsal fin intensely and
continuously dark pigmented. Besides, males of
A. eleutheria do not have the reticulate pattern
of narrow dark stripes on flank and the coarse
pattern of dark vermiculations on head present
Fig. 1. Apistogramma eleutheria: a, MZUSP 115293, holotype, 39.5 mm SL, adult brooding male; b, MZUSP 97095,
paratype, 38.9 mm SL, adult brooding female; c, MZUSP 101422, paratype, 31.3 mm SL, unsexed subadult; Brazil,
Pará, rio Xingu basin, rio Curuá drainage, rio Treze de Maio above the waterfalls.
a
b
c
84 Copyright © Verlag Dr. Friedrich Pfeil
in males of A. kullanderi. Apistogramma eleutheria
apparently does not reach a large size (around
40 mm SL) while A. kullanderi reaches 79.7 mm SL.
Additionally, A. eleutheria differs from oth-
ers species of the A. regani species group, like
A. cinilabra, A. commbrae, A. gosssei, A. inconspicua,
A. ortegai, A. ortmanni, A. paulmuelleri, A. regani
and A. rubrolineata, by the presence of two ab-
dominal stripes only in juveniles (vs. three or
four abdominal stripes invariably present). Apisto-
gramma eleutheria differs from A. eunotus, A. gossei
and A. ortegai by the presence of small dark dots
forming narrow vertical bands on caudal fin (vs.
caudal fin immaculate). It differs from A. kullan-
deri, A. rubrolineata and A. tucurui by the absence
of the reticulate pattern with narrows dark stripes
on flanks in males (vs. present) and the presence
of abdominal stripes in juveniles (vs. abdominal
stripes invariably absent).
Apistogramma eleutheria may also be com-
pared to A. hongsloi, in the A. macmasteri species
group, by having a similar pattern of blotches on
abdominal area. The pattern of irregular dark ab-
dominal blotches in A. eleutheria, however, starts
at pectoral fin and ends at the 7th bar whereas
in A. hongsloi this pattern of blotches starts more
posteriorly (at the 3rd bar) and ends at the 7th
bar. Also, females of A. eleutheria do not have the
dark blotch on chest (present in mature females
of A. hongsloi, as in all other A. macmasteri-group
species) and adult males do not present the first
dorsal-fin lappets prolonged (vs. first dorsal-fin
lappets prolonged in A. hongsloi, as in the other
A. macmasteri-group species).
Description. Largest male 39.5 mm SL, largest
female 40.1 mm SL. Body deep (35.1-37.4 % SL,
mean 36.4), greatest depth between verticals
through dorsal-fin spines V and VII. Dorsal and
ventral head profiles convex in juveniles and
adults, except one male with a soft nuchal hump.
Snout relatively short (10.7-12.6 % SL, mean
11.5); posterior border of maxilla reaching verti-
cal through anterior margin of orbit or slightly
beyond it. Orbit dorsolateral and entirely situated
in anterior half of head. Posterior margins of pre-
opercle and supracleithrum entire; posttemporal
serrations lacking.
Almost all body scales ctenoid, except for
cycloid scales on anterior portion of cheek, pre-
opercle, interopercle, subopercle, anteromedian
portion of predorsal and interpelvic area and
eventually around urogenital papilla. Scales
in E1 row 22* (5) or 23 (2). Cheek fully scaled,
4 horizontal scale rows. Predorsal scales 8-9. Pre-
pelvic area scaled; scales anterior to tip of cleithra
absent or 2-3, and 6-8 scales from tip of cleithra to
pelvic fin. Transverse row with 1 1
/
2 scales above
and 1
/
2 scales below anterior ramus of lateral line.
Circumpeduncular scale rows 16; 2 lateral line
Table 1. Proportional measurements of holotype and six paratypes of Apistogramma eleutheria and A. flavipedun-
culata. Pectoral-fin length of A. flavipedunculata obtained from 5 paratypes only. Holotype measurements included
in sample range. SD, standard deviation.
A. eleutheria A. flavipedunculata
holotype range mean SD holotype range mean SD
Standard length (mm) 39.5 23.2-40.1 45.5 26.3-45.5 – –
Percents of standard length
Body depth 37.4 35.1-37.4 36.4 1.0 37.9 35.9-38.5 37.3 1.0
Head length 35.2 35.2-36.7 36.0 0.6 34.3 34.3-35.7 34.8 0.5
Head depth 25.8 25.5-27.4 26.4 0.8 28.6 26.5-28.6 27.4 0.7
Head width 15.1 15.1-16.1 15.6 0.4 15.8 15.2-16.6 15.8 0.4
Orbital diameter 9.9 9.9-11.8 10.7 0.7 10.1 10.1-11.5 10.6 0.5
Interorbital width 8.9 8.7-9.5 9.1 0.4 9.0 8.5-9.9 9.1 0.4
Snout length 11.8 10.7-12.6 11.5 0.6 11.0 10.0-11.4 10.9 0.5
Preorbital depth 4.0 3.3-4.6 4.1 0.4 4.4 3.4-4.4 3.9 0.3
Caudal peduncle depth 16.9 15.8-16.9 16.4 0.4 16.1 16.1-18.0 16.7 0.6
Caudal peduncle length 10.5 10.0-11.0 10.5 0.3 9.8 9.3-10.4 9.8 0.4
Pectoral-fin length 26.3 26.3-28.7 27.1 0.9 29.0 28.9-30.7 29.8 0.8
Pelvic-fin length 26.2 25.9-29.2 27.6 1.2 32.6 29.8-34.1 32.3 1.7
Upper jaw length 10.6 9.2-10.9 10.0 0.7 11.8 10.0-11.8 10.7 0.6
Lower jaw length 14.3 13.4-14.3 13.9 0.3 14.1 13.6-14.4 14.0 0.3
Varella & Britzke: Apistogramma eleutheria and A. flavipedunculata
Ichthyol. Explor. Freshwaters, Vol. 27, No. 1
85
Copyright © Verlag Dr. Friedrich Pfeil
scales plus 7 dorsally and 7 ventrally. Lateral-line
scales range from 14-17/6-8. Tube-bearing scales
on upper branch of lateral line 11 (2), 12 (2), 13 (1),
14 (1), 15* (1), plus 2-5 pore-bearing scales. Tube-
bearing scales on lower branch of lateral line 3 (3),
5 (2), 6* (2), plus 1-4 pore-bearing scales. Scales
between upper lateral line and first dorsal-fin
spine 4. Only 1
/
2 scale between anterior ramus of
lateral line and last dorsal-fin spine. Fins naked
except caudal fin, which is scaled basally for one-
third of its length or slightly beyond.
Infraorbital series composed of three bony
elements: lachrymal, infraorbital one and infraor-
bital 2 + 3 (co-ossification interpreted by presence
of median pore). Infraorbital canal with four
pores on lachrymal, ventro-posterior one shared
by anterior opening of canal through infraorbi-
tal one; postlachrymal infraorbital pores three:
anteriormost pore shared between infraorbital
one and infraorbital 2 + 3, second pore median,
and posteriormost pore close to posterior end
of infraorbital 2 + 3. Dentary with five pores;
anguloarticular canal present with simple pore.
Preopercular pores six; nasal with pores at each
end, posterior one shared with anterior opening
of frontal canal. Frontal with four pores plus
coronalis pore.
Dorsal fin XVI.6 (3), XVII.5 (1), XVII.6* (3).
Lappets of dorsal-fin spines not conspicuously
prolonged in mature males. Posterior margin
of soft dorsal fin pointed with 3rd ray longest;
tip of 3rd ray reaching approximately one-third
length of caudal fin. Anal fin III.6 (2), IV.5* (4),
IV.6 (1). Posterior margin of soft anal fin pointed
in males and females; third ray longest, reach-
ing approximately one-third length of caudal
fin. Caudal fin rounded with 3 procurrent and 8
principal each lobe. Pectoral fin rounded, nearly
symmetrical, with 13* (7) rays; 6th ray longest,
tip reaching vertical through genital papilla.
Pelvic fin pointed, first ray elongated; tip of first
pelvic-fin ray usually reaching base of 1st anal-fin
spine, extending beyond urogenital papilla but
not reaching anal-fin origin in few specimens.
Jaw teeth unicuspid, erect, cusp slightly ret-
rorse; outer row teeth similar in size or slightly
larger than inner row teeth; 24-25 teeth in upper
right jaw outer hemi-series, 23-24 in upper left,
26-28 in lower right jaw outer hemi-series, 26-27
in lower left. Outer row of teeth occupying almost
entire premaxilla and dentary margins; inner row
of premaxilla and dentary extending to approxi-
mately 1
/
3 of jaw margin; few teeth forming one
additional, short and irregular, symphyseal row
on upper and lower jaws.
Gill rakers on first epibranchial lobe 2 (1),
3* (4); one gill raker in angle and 4 (5) gill rakers
on ceratobranchial. Gill rakers laterally on lower
pharyngeal toothplate 9 (1), 10 (2), 11 (2).
Vertebrae 11 + 13 = 24 (3), 11 + 14 = 25 (2), 12 + 12
= 24 (1) or 12 + 13 = 25* (1), including last half-
centrum. Hypurals 1-5 separate.
Color pattern in alcohol. Juveniles and imma-
ture specimens (three specimens 23.2-29.0 mm
SL) with tan ground colour, dorsal half of head
and body darker than ventral half; markings
dark brown or blackish. Brown preorbital stripe
running from anterior margin of orbit to post-
labial skin fold. Postorbital stripe aligned with
preorbital stripe, running from posterior margin
of orbit to posterior edge of opercle, divided in
a blotch just posterior to orbit and a stripe on
dorsal border of opercle. Dark supraorbital stripe
conspicuous only closer to dorsoposterior mar-
gin of orbit, running more faintly onto median
area of nape. Suborbital stripe evident, oblique
from just beneath ventral margin of orbit to
end of preopercle, with another dark marking
expressed as a posteroventral continuation of
suborbital stripe occupying area of articulation
between subopercle, interopercle and opercle.
About seven to eight irregularly pigmented dark
brown blotches from nape, along dorsal-fin base,
to caudal peduncle. Anteriormost blotch on nape,
followed by one blotch at dorsal-fin origin, four or
five blotches along dorsal fin and one on caudal
peduncle. Dorsal blotches corresponding in po-
sition to six or seven vertical bars on flank, only
nuchal blotch not corresponding to any bar. Bars
1-2 fainter or almost indistinct, situated anterior
to vertical through dorsal-fin spine V. Bars 3-7
more evident; bar 3 between dorsal-fin spines
VII and IX, bars 4-6 between verticals through
anal-fin origin and end of dorsal fin, and bar 7
on middle of caudal peduncle. Midlateral dark
band aligned with postorbital stripe, finishing just
before caudal flexure and separate from darker
caudal-fin blotch. Band occupying entire depth
of E1 scale row plus adjacent half of scales in row
E0, but fragmented between bars 5-6, between
bars 6-7, and less frequently between bars 2-3.
Additional two narrow abdominal stripes formed
by a series of dark spots, fainter than midlateral
band, running from end of bar 1 to end of bar 5
and occupying two horizontal scale rows below
86 Copyright © Verlag Dr. Friedrich Pfeil
E0 row. Caudal fin with a large blotch some-
what rectangular but with poorly defined edges,
tangent to caudal flexure and situated between
caudal-fin rays D4-V4; two or three faint dark,
narrow bands posterior to caudal blotch. Dorsal
fin dusky, with more concentration of dark pig-
ment in basal portion of spines, corresponding
with position of flank bars, and with two or
three irregular lighter spaces on soft portion of
fin. Anal fin dusky, with two paler spaces on soft
portion. A specimen measuring 31.3 mm SL, ten-
tatively considered an immature female, may be
interpreted as showing intermediate colouration
between juvenile and adult patterns, with anal fin
dark margined. Pectoral and pelvic fins hyaline,
with no markings except pelvic fin with a very
faint pigmentation on its spine.
Adult male and females retaining most of
juvenile colour pattern, with some differences
attributed mainly to sexual dichromatism.
Adult male ground colour brown, darker
than females. Juvenile pattern of two abdominal
stripes replaced in adult male by irregular, faint
pigmentation on abdomen, occupying most of
area of four abdominal horizontal scale rows
below E0 series, from pectoral-fin base to bar 7.
Dorsal and anal fins of male more conspicuously
ornamented; dorsal fin darker with three series of
light spots on soft portion (besides basal blotches
also present in juveniles), soft portion of anal fin
with lighter and darker stripes interspaced by
dusky background.
Ground colour of adult females brown dor-
sally, pale yellowish on ventral half. Juvenile pat-
tern of abdominal stripes replaced by an irregular
pattern of dark pigmentation on abdomen; differ-
ent from males, this pigmentation is conspicuous
and expressed approximately in same position of
vertical bars, resulting in an overall barred pat-
tern. Midlateral band appearing more fragmented
than in males and juveniles, forming a series of
six blotches on midlateral area. Dark blackish
Fig. 2. Apistogramma flavipedunculata: a, MZUSP 115292, holotype, 45.5 mm SL, adult male; b, MZUSP 97594,
paratype, 30.5 mm SL, unsexed subadult; Brazil, Pará, rio Xingu basin, upper rio Curuá drainage, rio Escorpião.
a
b
Varella & Britzke: Apistogramma eleutheria and A. flavipedunculata
Ichthyol. Explor. Freshwaters, Vol. 27, No. 1
87
Copyright © Verlag Dr. Friedrich Pfeil
pigmentation of pelvic fin in adult females more
conspicuous than in adult males, occupying spine
and first soft ray. Suborbital stripe continuing onto
median area of gular region, occupying ventral
portion of opercular region and branchiostegal
membranes. Anterior to gular pigmented area,
another concentration of dark pigmentation
ventrally on dentary, separated from gular pig-
mentation by a small pale area (Fig. 5a).
Distribution and habitat. Apistogramma eleutheria
is known only from the rio Treze de Maio above
its major waterfalls (see photo in Sousa et al.,
2010: fig. 5a), rio Xingu basin, Brazil. Although
some maps and the Índice de Nomes Geográ-
ficos (IBGE, 2011) give the name rio Três de
Maio, Treze de Maio is the correct name based
on communications with locals and road signs
along BR163.
Etymology. The specific epithet eleutheria
(λευθερα) is a Greek noun that means freedom. It
makes reference to the name of the river, because
Treze de Maio is the date (13 May 1888) on which
Princess Isabel signed the Áurea law, marking
the official end of legal slavery in Brazil. A noun
in opposition.
Apistogramma flavipedunculata, new species
(Fig. 2)
Apistogramma sp. upper Curuá: Varella & Sabaj
Pérez, 2014: 255.
Holotype. MZUSP 115292, 45.5 mm SL, adult
male; Brazil: Pará: Altamira: rio Xingu basin: rio
Iriri drainage: rio Escorpião, tributary of the up-
per rio Curuá, at route BR163 bridge; 8°53'54" S
54°59'20" W; A. Netto-Ferreira, J. Birindelli, L.
Sousa, M. Sabaj-Pérez & N. Lujan, 29 Oct 2007.
56°48'W 56°00'W 55°12'W 54°24'W
7°48'S
8°36'S
9°24'S
Apistogramma eleutheria
Apistogramma flavipedunculata
Apistogramma kullanderi
Apistogramma sp. ‘Peixoto’
Apistogramma sp. ‘middle Curuá’
Apistogramma sp. ‘Jamanxim’
REBIO-Nascentes da
Serra do Cachimbo
Road BR 163
Isoline ca. 400 m
Fig. 3. Serra do Cachimbo, showing known distribution of Apistogramma eleutheria, A. flavipedunculata, A. kullan-
deri and other putative new species occurring in the rio Curuá and adjacent rivers. 1, waterfalls of the rio Treze
de Maio; 2, series of waterfalls of the rio Curuá.
88 Copyright © Verlag Dr. Friedrich Pfeil
Paratypes. MZUSP 97594, 5 immature specimens
identified as males, 26.3-39.6 mm SL, 1 adult male,
45.3 mm SL; collected with holotype.
Diagnosis. Adult males of A. flavipedunculata
are distinguished from males of all congeners by
having the posterior part of the flank and caudal
peduncle dominated by a light yellow pigmenta-
tion, resulting in the dark midlateral band frag-
mented posteriorly and the dark bars (present in
juveniles) masked or absent in the males.
Apistogramma flavipedunculata can be distin-
guished from all congeners except A. luelingi,
A. eleutheria and A. pleurotaenia by having exclu-
sively four anal-fin spines instead of three (four
spines occurring very rarely in few species; see
discussion). It differs from A. luelingi by the ab-
sence of midlateral spot, the lack of accentuated
elongation of lappets on spines of dorsal fin and
lyrate caudal fin in adult males, the absence of
conspicuous abdominal stripes (vs. three dark
stripes evident on abdomen), and by the presence
of numerous dark narrow bands on the caudal
fin (vs. dark markings on caudal fin absent or
inconspicuous).
Among species of the A. regani group, A. acr-
ensis, A. caetei, A. ortmanni, A. paulmuelleri and
A. regani resemble A. flavipedunculata with respect
to body shape and presence of narrow vertical
bands on the caudal fin. Apistogramma flavipe-
dunculata is distinguished from A. acrensis and
A. caetei by having more dorsal-fin spines (16-
17 vs. 15), from A. ortmanni, A. paulmuelleri and
A. regani by having two or three series of faint
dark markings on abdominal area not forming
well-defined stripes (vs. presence of three or
four well-defined dark abdominal stripes) and
caudal-fin blotch rounded and restricted to the
centralmost rays of the caudal fin (vs. caudal-fin
blotch vertically elongated in A. ortmanni, A. paul-
muelleri and A. regani, occupying mostly or totally
the depth of caudal-fin base).
Description. Largest male 45.5 mm SL, no female
specimens confirmed. Body deep (35.9-38.5 %
SL, mean 37.3), greatest depth between verticals
through dorsal-fin spines IV and V. Dorsal and
ventral head profiles convex in juveniles and
adults, with interorbital concavity in largest
males. Snout relatively short (10.1-11.4 % SL,
mean 10.9); posterior border of maxilla reaching
vertical through anterior margin of orbit in most
specimens, falling short of orbital margin in larg-
est specimens. Orbit dorsolateral, mostly but not
entirely situated in anterior half of head. Posterior
margins of preopercle and supracleithrum entire;
posttemporal serrations lacking.
Almost all body scales ctenoid, except for
cycloid scales on cheek, preopercle, interopercle,
subopercle, anteromedian portion of predorsal
and interpelvic area and eventually around uro-
genital papilla. Scales in E1 row 22 (2) or 23* (5).
Cheek fully scaled with 3 horizontal scale rows.
Predorsal scales 9-10. Prepelvic area scaled; scales
anterior to tip of cleithra absent or 3-4, and 6-7
scales from tip of cleithra to pelvic fin. Transverse
row with 1 scale above and 6 1
/
2 or 7 1
/
2 scales below
anterior ramus of lateral line. Circumpeduncular
scale rows 16; 2 lateral line scales plus 7 dorsally
and 7 ventrally. Lateral-line scales range from
14-17/7-8. Tube-bearing scales on upper branch
of lateral line 12 (1), 13* (3), 14 (3), plus 2-4 pore-
bearing scales. Tube-bearing scales on lower
branch of lateral line 2 (1), 5 (1), 6* (2), 7 (3) plus
1-4 pore-bearing scales. Scales between upper
lateral line and first dorsal-fin spine 3 or 3 1
/
2. Only
1
/
2 scale between anterior ramus of lateral line and
last dorsal-fin spine. Fins naked except caudal fin,
scaled basally on approximately one-third of its
length.
Configuration of infraorbital bones and later-
alis pores of head as in A. eleutheria.
Dorsal fin XVII.5 (5), XVII.6* (2). Lappets
of anteriormost eight dorsal-fin spines slightly
prolonged in mature males. Posterior margin of
soft dorsal fin pointed with second and third rays
longest, with tips reaching approximately one-
third length of caudal fin in smaller specimens
and half-length of caudal fin in largest specimens.
Anal fin IV.6* (7). Posterior margin of soft anal fin
pointed; 3rd and/or 4th ray longest, extending
slightly beyond half-length of caudal fin. Caudal
fin rounded with 3 procurrent and 8 principal
rays each lobe. Pectoral fin rounded, nearly sym-
metrical, with 11 (1), 12* (6) rays; 6th ray longest,
tip reaching genital papilla or extending slightly
beyond vertical through genital papilla. Pelvic fin
pointed, first ray elongated; tip of first pelvic-fin
ray usually reaching base of 1st anal-fin spine.
Jaw teeth unicuspid, erect, cusp slightly retrorse;
outer row teeth similar in size or slightly larger
than inner row teeth; 20-24 teeth in upper right
jaw outer hemi-series, 21-24 in upper left, 28-31
in lower right jaw outer hemi-series, 29-30 in
Varella & Britzke: Apistogramma eleutheria and A. flavipedunculata
Ichthyol. Explor. Freshwaters, Vol. 27, No. 1
89
Copyright © Verlag Dr. Friedrich Pfeil
lower left. Outer row of teeth occupying 2
/
3 length
of premaxilla and entire dentary margins; inner
row of premaxilla and dentary extending to ap-
proximately middle of jaw margin; additional
irregular, symphyseal row of teeth present on
upper and lower jaws.
Gill rakers on first epibranchial lobe 4 (1), 5 (1),
6* (5); one gill raker in angle and 1 (1), 2 (2), 3 (3),
4* (1) gill rakers on ceratobranchial. Gill rakers
laterally on lower pharyngeal toothplate 10 (1),
11 (2), 12* (4).
Vertebrae 11 + 13 = 24 (5), 12 + 12 = 24* (1),
12 + 13 = 25, including last half-centrum. Hypurals
1-5 separate.
Color pattern in alcohol. Type material com-
posed of large males and smaller specimens
tentatively determined as males (by examination
of shape of genital papilla); same color pattern for
all specimens excepting some aspects as otherwise
noted. Ground colour yellowish tan, slightly
darker on dorsum closer to dorsal-fin base than
ventrally; colour markings on head and flanks
dark brown and blackish on distal margin of fins.
Brown preorbital stripe from anterior margin
of orbit to postlabial skin fold, extending onto
tip of upper jaw. Postorbital stripe aligned with
preorbital stripe, running from posterior margin
of orbit to posterior edge of opercle. Dark supraor-
bital stripe evident from dorsoposterior margin
of orbit obliquely onto median area of nape in
largest males; reduced to a rounded blotch near
orbit in few specimens. Suborbital stripe running
obliquely from just beneath ventral margin of
orbit to anterior margin of subopercle, often with
a brief pale hiatus across posteroventral margin
of preopercle; suborbital stripe more evident in
two largest males. Interorbital stripes distinct
only in holotype, fragmented into two paired
dark vermiculations from anterodorsal margin
of orbit onto median area.
About seven or eight irregularly pigmented
dark brown blotches from nape, along dorsal-fin
base, to caudal peduncle; anteriormost blotch on
nape, followed by one blotch on dorsal-fin origin,
five blotches along dorsal fin and one on caudal
peduncle. Blotches restricted to dorsum, not
extending ventrally to form bars and more evi-
dent in smallest specimens (i. e., three specimens
26.3-35.8 mm SL); posteriormost blotch, on caudal
peduncle, partially or totally indistinct in larger
specimens. Midlateral dark band aligned with
postorbital stripe, ending approximately at verti-
cal through caudal peduncle blotch and separate
from darker caudal-fin blotch in five specimens
(26.3-39.6 mm SL); midlateral band fragmented
posteriorly and faded past vertical through bar
5 in two largest males. Band occupying entire
depth of E1 scale row plus adjacent half of scales
in row E0 at least anteriorly (i. e., band fragmented
posteriorly in largest specimens). Two or three
faded abdominal series of irregular dark mark-
ings, composed of pigmentation on distal margin
of some scales (present only in four specimens
and more visible in holotype); abdominal series
occupying two or three horizontal scale rows
below E0 row. Dominance of light pigmentation
posteriorly to vertical through first soft dorsal-fin
ray onto base of caudal fin, masking melanin-
based colour markings except caudal fin blotch,
as most remarkable difference between largest
adult males and smaller specimens.
Caudal fin with a rounded blotch tangent to
caudal flexure and situated between caudal-fin
rays D3-4 and V4; posterior to caudal blotch,
eight or nine well-defined narrow bands to dis-
tal margin of fin. Dorsal fin dusky, with strong
blackish pigmentation on lappets of dorsal-fin
spines occupying most of first dorsal-fin spine and
distal half of second and third dorsal-fin spines;
distal portion of dorsal fin slightly darker than
basal portion. Dorsal fin with four slightly darker
vertical bands on distal portion of soft rays. Anal
fin dusky with dark gray pigmentation on distal
portion of spines and soft rays; four dark vertical
bars on two posteriormost soft rays. Pectoral fin
hyaline. Pelvic fin hyaline except black blotch
covering its spine and one or two adjacent soft
rays, much more evident in largest males than in
smaller specimens.
Distribution. Apistogramma flavipedunculata is
known only from the rio Escorpião, tributary of
the upper rio Curuá, rio Xingu basin, Brazil. The
species is syntopic with A. kullanderi in this river.
Etymology. The specific epithet flavipedunculata,
an adjective, refers to the yellow pigmentation
that dominates the posterior part of the body in
adult males; from the Latin flavus, yellow, and
pedunculatus, with a peduncle.
90 Copyright © Verlag Dr. Friedrich Pfeil
Discussion
The number of nominal species of Apistogramma
reported from the rivers of the Brazilian Shield
is relatively low when compared to that of the
Western Amazon, Orinoco basin and Guiana
Shield. Only the rio Madeira basin, along the
western border of the Brazilian Shield, presents
a high known diversity with 12 valid species, ten
of them endemic. This may be correlated with
the early scientific expeditions to this area (e. g.
Johann Natterer in 1817-1835). Other tributaries
to the right margin of the rio Amazonas were
systematically explored later, more intensively in
the last fifty years. Only a few species have been
described from these drainages, as A. taeniata
(Günther, 1862) described from the lower rio
Tapajós (distributed also in the lower portion rio
Xingu) and A. tucurui Staeck, 2003 from the area of
Turucui reservoir in the rio Tocantins. Other spe-
cies of Apistogramma reported from these drain-
ages are tentatively identified as species widely
distributed in the main channel or lower portion
of tributaries of the Amazon river, such as A. cf.
pertensis (Haseman, 1911) in the rio Tapajós (RB,
pers. obs.) and A. agassizi in the lower rio Xingu, or
are putatively undescribed (e. g. Koslowski, 2002).
Apistogramma eleutheria and A. flavipedunculata,
together with the recently described A. kullan-
deri, are the only formally named species of the
genus from the middle and upper portions of
the rio Xingu basin. Their known distribution is
restricted to the Serra do Cachimbo in the upper
portion of the rio Curuá (A. flavipedunculata and
A. kullanderi) and the rio Treze Maio (A. eleutheria).
Apistogramma eleutheria is isolated from the lower
portion rio Treze de Maio by a waterfall with same
name (Souza et al., 2010: 259, fig. 5c). Similarly,
A. kullanderi and A. flavipedunculata are isolated
from the middle portion of the Curuá drainage
by a series of three waterfalls (indicated in Fig. 3
by arrow 2), as previously described by Birindelli
et al. (2009: 16, fig. 6) and Varella & Sabaj Pérez
(2014: 252, fig. 7).
Besides those three species, an expedition to
the Serra do Cachimbo in 2014 brought fishes of
another new species from the rio Curuá and rio
Treze de Maio below the waterfalls, provisionally
denominated here as Apistogramma sp. middle
Curuá (Fig. 4). This undescribed Apistogramma
is known only from juveniles and small adults
that resemble other forms encountered in the
lower portions of the rio Iriri and rio Xingu, and
shares many similarities with several species of
the A. regani group. Although the lack of mature
males and females of this species precludes a
definitive identification, Apistogramma sp. middle
Curuá can be easily distinguished from A. eleuthe-
ria and A. flavipedunculata by the presence of a
black blotch just below the posterior edge of the
mouth (arrow in Fig. 4; vs. absent in A. eleutheria
and A. flavipedunculata) and three anal-fin spines
(vs. modally or always four spines in A. eleutheria
and A. flavipedunculata, respectively).
Rivers adjacent to the rio Curuá (tributaries
of the rio Teles Pires and rio Jamanxim) present
Fig. 4. Apistogramma sp. middle Curuá, ca. 25 mm SL; Brazil, Pará, rio Xingu basin, rio Curuá basin, rio Treze
de Maio below the waterfalls; alive just after capture. Photograph by William Ohara.
Varella & Britzke: Apistogramma eleutheria and A. flavipedunculata
Ichthyol. Explor. Freshwaters, Vol. 27, No. 1
91
Copyright © Verlag Dr. Friedrich Pfeil
putative undescribed species deserving of com-
ments. Apistogramma sp. Jamanxim is most simi-
lar to A. flavipedunculata with respect to general
body shape, dorsal and anal fins comparatively
high and with their posterior margin extending
half the length of the caudal fin, and elements of
the color pattern such as caudal fin with numer-
ous dark narrow bands and conspicuous black
marking on the spine and first ray of pelvic fin in
males. Apistogramma flavipedunculata differs from
it by having four instead of three anal-fin spines,
by the peculiar color pattern on the posterior
portion of the flank and caudal peduncle (i. e.,
dominated by yellow pigmentation that masks
the melanin-based markings), and by not having
dark-margined scales on dorsum (vs. distal mar-
gin of scales on dorsum blackish or dark brown).
Apistogramma sp. Peixoto is an undescribed spe-
cies from the rio Braço Norte, tributary of the rio
Peixoto de Azevedo (rio Teles Pires basin), similar
in several aspects to A. eleutheria and A. kullanderi.
Brooding females of A. eleutheria, A. sp. Peixoto
and A. kullanderi share the possibly apomorphic
states of the gular region covered by a blackish
pigmentation, and the presence of dark markings
on the chest and belly (Fig. 5). In A. eleutheria, the
conspicuous ventral marking is restricted to the
gular area, but there are fainter markings on mid-
ventral area, on spine of pelvic fin and laterally
on chest close to opercle (Fig. 5a). Apistogramma
sp. Peixoto presents conspicuous black marking
on gular region, extending posteriorly onto the
median portion of chest, the spine and lateralmost
rays of pelvic fin; separate from that, another
strong black marking on midventral area extends
posteriory onto anal fin (Fig. 5b). Apistogramma
kullanderi has the sexually dimorphic colora-
tion in females over-expressed, resulting in the
ventral body almost totally covered by intense
black pigmentation (Fig. 5c). Thus, in A. eleuthe-
ria, A. sp. Peixoto and A. kullanderi, which are
allopatrically isolated in rivers of the Serra do
Cachimbo, females develop sexual dimorphism
based on more or less the same color pattern ele-
ments, differing from each other by the intensity
and distribution of the color expression. On the
other hand, males of A. kullanderi are much larger
than females and seem to show very distinctive
Fig. 5. Schematic drawing of ventral body in species of Apistogramma showing extent and condition of dark
markings present on brooding females. a, A. eleutheria, MZUSP 97095, 38.9 mm SL; b, Apistogramma sp. Peixoto,
MZUSP 116959, 38.7 mm SL; and c, A. kullanderi, MZUSP 97597, 58.7 mm SL.
a b c
92 Copyright © Verlag Dr. Friedrich Pfeil
dimorphic color pattern consisting of many dark
stripes on flanks and a coarse pattern of dark
vermiculations on the head. Males of A. eleutheria
are slightly darker than females, and males of
Apistogramma sp. Peixoto have few faint dark
markings on the head and some pale spots on
the flank, but neither species present the strong
sexually dimorphic coloration as in A. kullanderi.
Varella & Sabaj Pérez (2014) inferred that large
body in A. kullanderi may be due to its isolation in
the upper part of rio Curuá, which is depauperate
of cichlids and potential predators (i. e., phylo-
genetic constraints on body size reduced when
competition and predatory pressure are limited).
Ready et al. (2006) studied allopatric populations
from the Guamá and adjacent rivers in Pará,
previously identified as A. caetei Kullander, 1980,
but with slight differences in color pattern. They
found three closely related lineages with strong
prezygotic isolation and sexual selection through
female mate choice involved in the diversification
in Apistogramma. Following the same idea, the
evolution of large size and sexually dimorphic
coloration in A. kullanderi would be possibly
correlated: an unrestrained male-biased sexual
dimorphism in an isolated area with reduced en-
vironmental constraints resulting in a large body.
Occurring syntopically with Apistogramma sp.
Peixoto in the rio Braço Norte, there are other
cichlids (e. g. species of Satanoperca, Crenicichla
and Cichla) and other fish groups (e. g. species
of Serrasalmus and Pimelodus), which can be con-
sidered as potential competitors and predators.
They may represent environmental constraints
for the development of a very flashy dimorphic
coloration and large body like that found in A. kul-
landeri. However, A. eleutheria occurs above large
waterfalls in the upper rio Treze de Maio, which is
depauperate like the upper rio Curuá. Body size
and sexual dimorphism in A. eleutheria, however,
have apparently not evolved like in A. kullanderi.
Anal-fin spines. The number of anal-fin spines
is traditionally useful in the taxonomy of species
and genera of Cichlidae, as well as to reconstruc-
tions of their phylogenetic relationships, notably
within the Neotropical clade. Kullander (1983a,
1989, 1996) discussed in detail this character and
its implications for the delimitation of supraspe-
cific groups. In many published or unpublished
cladistics analysis (e. g. Cichocki, 1976; Kul-
lander, 1998; Landim, 2006; Chakrabarty, 2007;
Graça, 2008; Musilová et al., 2009), a condition
of three to four anal-fin spines was codified as
plesiomorphic among cichlids. Reduction to two
anal-fin spines is an autapomorphy of Biotoecus
dicentrarchus Kullander, 1989, and the condition
of five or more spines is synapomorphic for the
tribe Heroini (sensu Kullander, 1998).
The only geophagine with five or more anal-fin
spines is Apistogrammoides pucallpaensis Meinken,
1965 (6-9 spines), which has been considered
closely related to, if not congeneric with Apisto-
gramma (e. g. Kullander, 1998; López-Fernández et
al., 2005, 2010). Almost all species of Apistogramma
have three anal-fin spines. Exceptions with four
anal-fin spines include a few specimens of A. agas-
sizi and A. cacatuoides (1 specimen each; Kullander,
1980), A. cruzi (1 specimen; Kullander, 1986) and
A. kullanderi (2 of 44 specimens). Among species
typically with three anal-fin spines, A. commbrae is
the only one that frequently presents four anal-fin
spines (15 of 56 specimens examined here; also
about 15 % of the material examined by Kullander,
1982). Apistogramma flavipedunculata exclusively
and A. luelingi Kullander, 1976 almost exclusively
(40 of 42 specimens, according to Kullander, 1986)
have four-anal fin spines. Apistogramma eleutheria
presented 5 of 7 specimens with four spines,
perhaps an indication that it is the most common
count for the species. Finally, the holotype (only
specimen known) of A. pleurotaenia (Regan, 1909)
also possesses four spines.
Kullander (1988) and Varella & Moreira
(2013) described two species of Teleocichla having
exclusively four anal-fin spines (T. centrarchus
and T. wajapi, respectively). These are the only
species with four instead of three anal-fin spines
among species of the Crenicichla-Teleocichla clade,
although four spines are rarely recorded for a few
species typically having three (Varella & Moreira,
2013). For this reason, the authors considered this
condition probably apomorphic and indicative of
a closer-relationship between T. centrarchus and
T. wajapi, which would demand the division of
the character state if in a cladistic analysis. Also
Kullander (1996) suggested considering modal
anal-fin counts rather than ranges in phylogenetic
analyses, if there are adequate data available
(i. e. large amount of specimens of the terminal
taxa). In the particular case of a comprehensive
phylogeny on Apistogramma, to follow this sug-
gestion might avoid missing important variation
present between the included species.
Varella & Britzke: Apistogramma eleutheria and A. flavipedunculata
Ichthyol. Explor. Freshwaters, Vol. 27, No. 1
93
Copyright © Verlag Dr. Friedrich Pfeil
Comparative material. Apistogramma sp. middle Cu-
ruá: Brazil: Pará: Altamira: rio Xingu drainage: MZUSP
97617, 2 juveniles, 9.6-10.2 mm SL; flooding at brigde
of road BR-163, close to district of Castelo dos Sonhos;
8°15'17" S 55°06'40" W. – MZUSP 116007, 4, 19.4-25.0 mm
SL; tributary of rio Treze de Maio below the waterfalls,
rio Curuá drainage; 8°39'08" S 55°02'08" W. – MZUSP
116025, 5, 17.0-25.2 mm SL; tributary of rio Curuá, close
to district of Cachoeira da Serra; 8°36'18.51" S 55°05'
28.67" W. – MZUSP 116620, 3, 17.6-24.4 mm SL; tributary
of rio Curuá, close to district of Cachoeira da Serra;
8°36'41.80" S 55°07'25.00" W.
Apistogramma sp. Jamanxim: Brazil: Pará, Novo
Progresso: rio Tapajós drainage: MZUSP 97261, 1,
22.7 mm SL; rio Jamanxim, approximately 30 km from
Castelo dos Sonhos; 8°15'17" S 55°06'40" W. – MZUSP
97561, 18, 12.4-37.9 mm SL; rio Jamanxim, flooded
shoreline at BR163 bridge, south of Vila Mil; 7°51'47.00" S
55°10'47.00" W.
Apistogramma sp. Peixoto: Brazil: Pará: Novo
Progresso: rio Teles Pires drainage (rio Tapajós basin):
MZUSP 96770, 1, 21.1 mm SL; right margin tributary of
rio Peixoto de Azevedo, at BR163 bridge, close to Força
Aérea Brasileira (FAB) base: 10°17'14" S 54°50'57" W. –
MZUSP 96828, 43, 13.7-48.3 mm SL; MZUSP 116052,
10, 28.0-61.7 mm SL; tributary of rio Braço Norte (rio
Peixoto de Azevedo drainage), at BR163 bridge, close
to FAB base; 9°25'55" S 54°52'11" W. – MZUSP 96846,
141, 13.0-46.7 mm SL; MZUSP 101428, 3, 28.1-30.0 mm
SL; MZUSP 116053, 20: 26.6-53.3 mm SL; tributary of
rio Braço Norte (rio Peixoto de Azevedo drainage), at
BR163 bridge, close to FAB base; 9°28'20" S 54°52'11" W.
– MZUSP 101434, 1, 40.5 mm SL; tributary of rio Braço
Norte, rio Peixoto de Azevedo, at BR163 bridge, close
to FAB base; 9°19'17" S 54°50'26" W. – MZUSP 116595,
7, 13.9-38.7 mm SL; tributary of rio Braço Norte (rio
Peixoto de Azevedo drainage), on road BR163 close to
border between States of Pará and Mato Grosso,
09°28'20" S 54°51'32" W.
A. acrensis: MCP 29419, 2, 30-32 mm SL; Brazil:
Acre: rio Purus basin, rio Branco.
A. agassizii: ANSP 197892 (examined from photo-
graph); Brazil: Pará: lower rio Xingu close to Porto do
Moz.
A. commbrae: MCP 22849, 1, 25 mm SL; Brazil: Rio
Grande do Sul: Uruguaiana: rio Uruguai basin. – MZUSP
59972, 22 (with 3 anal-fin spines), 13.5-23,3 mm SL, 13
(with 4 anal-fin spines), 15.7-21.6 mm SL; Brazil: Mato
Grosso do Sul: Corumbá: rio Paraguai basin: rio Ver-
melho. – MZUSP 100185, 2 (with 4 anal-fin spines),
20.5-22.4 mm SL, 12 (with three anal-fin spines), 14.1-
21.3 mm SL: Brazil: Mato Grosso: Corumbá: rio Paraguai
basin: rio Miranda.
A. hongsloi: MZUSP 96446, 15, 13.0-46.3 mm SL,
Venezuela, Bolivar, Caicara del Orinoco. – LBP 18704,
18, 21.2-47.1 mm SL, Colombia Meta, Canõ Iraca, Río
Guaviare basin.
A. inconspicua: ROM 69320, 2, 17.6-24.3 mm SL,
Bolivia, Santa Cruz, Río Negro, Vers Perseverancia.
A. luelingi: ROM 82894, 2, 21.6-28.5 mm SL, Peru,
Madre de Dios, Tambopata, Tambopata River.
A. ortegai: MUSM 48898, holotype, 38.4 mm SL;
Peru, Loreto, Distrito Pebas: small stream tributary of
Ampiyacu River.
A. ortmanni: ROM 91316, 8, 35.1-55.7 mm SL, Guy-
ana, Imbaima creek, tributary of Kuribong River.
A. rubrolineata: ROM 63879, 8, 20.5-34.5 mm SL,
Peru, Madre de Dios, Tambopata, Planchon creek.
A. taeniata: ANSP 195006 (examined from photo-
graph); Brazil, Pará, lower rio Xingu close to Porto de
Moz.
Acknowledgements
Thanks to André Netto-Ferreira, José Luís Birindelli,
Leandro Sousa, Mark Sabaj Pérez, Nathan Lujan and
Pedro Hollanda-Carvalho for collecting the type spec-
imens. Fieldwork was supported in part by the All
Catfish Species Inventory (NSF DEB-0315963). Informa-
tion on Apistogramma from the lower Xingu and sug-
gestions for improving the manuscript were provided
by Mark Sabaj Pérez (supported by the iXingu Project,
NSF DEB-1257813). Flávio Bockmann and Ricardo
Castro allowed us to take radiographs in the Laborató-
rio de Ictiologia de Ribeirão Preto (LIRP), using the
machine obtained by the project Melhoria e Capacita-
ção das Coleções Científicas do Departamento de Bio-
logia da Faculdade de Filosofia, Ciências e Letras de
Ribeirão Preto-USP (FAPESP 09/54931-0). Visit of the
authors to MCP collection was supported by the project
Qualificação taxonômica da base de dados da coleção
de peixes do Museu de Ciências e Tecnologia – PUCRS
para uso on-line (MCT/CNPq: 45/2012: 504177/2012-
5). H. Varella is financially supported by FAPESP (grant
2011/14630-0) and R. Britzke by FAPESP (grant
2011/00269-4) and PROPe (004/2014).
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Received 18 June 2015
Revised 17 February 2016
Accepted 13 March 2016
96 Copyright © Verlag Dr. Friedrich Pfeil
Varella & Britzke: Apistogramma eleutheria and A. flavipedunculata
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INSTRUCTIONS TO CONTRIBUTORS
Ichthyological Exploration of Freshwaters
An international journal for field-orientated ichthyology
Articles appearing in this journal are indexed in:
AQUATIC SCIENCES and FISHERIES ABSTRACTS
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C O N T E N T S
Chen, Yan-Qiao, Chang-Lian Peng and E Zhang: Sinocyclocheilus guanyangensis, a new
species of cavefish from the Li-Jiang basin of Guangxi, China (Teleostei: Cyprinidae) . 1
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Covain and Juan I. Montoya-Burgos: Hypostomus formosae, a new catfish species from
the Paraguay River Basin with redescription of H. boulengeri (Siluriformes: Loricari-
idae) .............................................................................................................................................. 9
Britski, Heraldo A. and José L. O. Birindelli: Redescription of Leporinus altipinnis, a senior
synonym of L. falcipinnis, and comments on L. holostictus (Characiformes: Anostomi-
dae) ................................................................................................................................................ 25
Britz, Ralf: Pillaiabrachia siniae, a new species of earthworm eel from northern Myanmar
(Teleostei: Synbranchiformes: Chaudhuriidae) ..................................................................... 41
Decru, Eva, Emmanuel Vreven, Oumar Sadio and Jos Snoeks: Brycinus epuluensis, a new
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species of loach from a cave in eastern Yunnan, China (Teleostei: Cobitidae)................. 61
de Oliveira, Renildo R., Lucia Rapp Py-Daniel, Claudio H. Zawadzki and Jansen Zuanon:
Two new Amazonian species of Ancistrus with vestigial adipose fin, with an appraisal
on adipose fin loss in neotropical armoured catfishes (Teleostei: Loricariidae) .............. 67
Varella, Henrique R. and Ricardo Britzke: Apistogramma eleutheria and A. flavipedunculata,
two new species of dwarf cichlids from the rio Curuá on Serra do Cachimbo, Brazil
(Teleostei: Cichlidae) .................................................................................................................. 81
Ichthyological Exploration of Freshwaters
An international journal for field-orientated ichthyology
Volume 27 Number 1 April 2016
Cover photograph
Pillaiabrachia siniae (photograph by Ralf Britz)
Ralf Britz
(this volume pp. 41-47)
... Apistogramma fulfills these conditions in many areas of its distribution, in which a high level of endemism and species diversity occur. This condition stands out in the Western Amazon, particularly in Peru (21 endemic species), Orinoco River (19 endemic species), and Negro River basins (13 endemic species) [42]. Its species have a very extensive geographic distribution in several South American watersheds, the reflex of a complex biogeographic history [4], showing a temporal sequence of cladogenetic events associated with dramatic chromosomal changes. ...
Article
Full-text available
Neotropical cichlid fishes are one of the most diversified and evolutionarily successful species assemblages. Extremely similar forms and intraspecific polychromatism present challenges for the taxonomy of some of these groups. Several species complexes have a largely unknown origin and unresolved evolutionary processes. Dwarf cichlids of the genus Apistogramma, comprising more than a hundred species, exhibit intricate taxonomic and biogeographic patterns, with both allopatric and sympatric distributions. However, karyotype evolution and the role of chromosomal changes in Apistogramma are still unknown. In the present study, nine South American Apistogramma species were analyzed using conventional cytogenetic methods and the mapping of repetitive DNA sequences [18S rDNA, 5S rDNA, and (TTAGGG)n] by fluorescence in situ hybridization (FISH). Our results showed that Apistogramma has unique cytogenetic characteristics in relation to closely related groups, such as a reduced 2n and a large number of bi-armed chromosomes. Interspecific patterns revealed a scenario of remarkable karyotypic changes, including a reduction of 2n, the occurrence of B-chromosomes and evolutionary dynamic of rDNA tandem repeats. In addition to the well-known pre-zygotic reproductive isolation, the karyotype reorganization in the genus suggests that chromosomal changes could act as postzygotic barriers in areas where Apistogramma congeners overlap.
... Crenicichla and Apistogramma jockey for the most species-rich genus of Neotropical Cichlidae, the former with about 91 valid species (Kullander and Varella, 2015;herein) and the latter with about 88 (Britzke et al. 2014;Varella and Sabaj Pérez, 2014;Varella and Britzke, 2016;Römer et al. 2017). To help classify diversity within Crenicichla, at least eight species groups have been proposed, and most of them have been recovered as monophyletic (Kullander, 1990;Ploeg, 1991;Lucena and Kullander, 1992;Kullander et al., 2009;Piálek et al., 2011). ...
... Crenicichla and Apistogramma jockey for the most species-rich genus of Neotropical Cichlidae, the former with about 91 valid species (Kullander and Varella, 2015;herein) and the latter with about 88 (Britzke et al. 2014;Varella and Sabaj Pérez, 2014;Varella and Britzke, 2016;Römer et al. 2017). To help classify diversity within Crenicichla, at least eight species groups have been proposed, and most of them have been recovered as monophyletic (Kullander, 1990;Ploeg, 1991;Lucena and Kullander, 1992;Kullander et al., 2009;Piálek et al., 2011). ...
Article
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Crenicichla dandara, new species, is endemic to the rio Xingu above the Belo Monte narrows, and its major left bank tributary the rio Iriri. The new species is distinguished from all congeners by the following combination of characters: body uniformly black (live fishes) or dark brown (preserved specimens) in adults and 74-86 scales in the E1 row. Crenicichla dandara is considered herein to be the tenth species of the C. lugubris group, but its allies within the group are not clear. Among members of this group, C. dandara is most similar in general body shape, snout characteristics and squamation to C. lugubris and C. adspersa, but lacks a caudal-fin blotch as does C. johanna and C. monicae, a condition rare within the Crenicichla-Teleocichla clade. The overall black coloration in Crenicichla dandara is shared with the syntopic, but not closely related, Teleocichla preta. Both species inhabit clear water rapids where they hide among the shadows of large rocks, suggesting that the dark coloration is for camouflage and ambush predation. Based on a preliminary assessment of its conservation status following the criteria and categories of IUCN classification, Crenicichla dandara is considered as a species of least concern (LC). © 2018 by the Academy of Natural Sciences of Drexel University.
... The South American cichlid genus Crenicichla Heckel, 1840, with 90 species (Kullander, 2003;Kullander & Varella, 2015), ranks as the most species-rich cichlid genus next to Apistogramma Regan, with 92 species (Kullander, 2003;Varella & Britzke, 2016;Römer et al. 2017). The division of Crenicichla into less inclusive, diagnosable groups has been a common procedure to make taxonomic revisions of the genus practicable. ...
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