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Apistogramma kullanderi, new species, is described from the upper rio Curuá (Iriri-Xingu drainage) on Serra do Cachimbo, Pará, Brazil, and diagnosed by its maximum size of 79.7 mm SL (vs. 65.3 mm SL among wild-caught congeners); mature females having the unique combination of intense dark pigmentation continuous along base of dorsal fin and on ventral surfaces from gular region to anal-fin base; and mature males having a coarse, irregular pattern of dark spots and vermiculations on cheek and opercular series, and sides with 10-12 dark stripes, each stripe occupying proximal limits of adjacent scale rows and separated by paler region central to each scale. Apistogramma kullanderi is tentatively allocated to the A. regani lineage, although some characteristics (e.g., large body size) are more consistent with members the A. steindachneri lineage. Apistogramma kullanderi is endemic to an upland watershed isolated by large waterfalls and depauperate of cichlid diversity. Under those conditions, we speculate that ecological opportunities, reduced competition and sexual selection contributed to the evolution of large body size in A. kullanderi.
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Ichthyol. Explor. Freshwaters, Vol. 25, No. 3
Ichthyol. Explor. Freshwaters, Vol. 25, No. 3, pp. 243-258, 8 figs., 1 tab., December 2014
© 2014 by Verlag Dr. Friedrich Pfeil, München, Germany – ISSN 0936-9902
A titan among dwarfs:
Apistogramma kullanderi, new species
(Teleostei: Cichlidae)
Henrique R. Varella* and Mark H. Sabaj Pérez**
Apistogramma kullanderi, new species, is described from the upper rio Curuá (Iriri-Xingu drainage) on Serra do
Cachimbo, Pará, Brazil, and diagnosed by its maximum size of 79.7 mm SL (vs. 65.3 mm SL among wild-caught
congeners); mature females having the unique combination of intense dark pigmentation continuous along base
of dorsal fin and on ventral surfaces from gular region to anal-fin base; and mature males having a coarse, ir-
regular pattern of dark spots and vermiculations on cheek and opercular series, and sides with 10-12 dark stripes,
each stripe occupying proximal limits of adjacent scale rows and separated by paler region central to each scale.
Apistogramma kullanderi is tentatively allocated to the A. regani lineage, although some characteristics (e. g., large
body size) are more consistent with members the A. steindachneri lineage. Apistogramma kullanderi is endemic to
an upland watershed isolated by large waterfalls and depauperate of cichlid diversity. Under those conditions,
we speculate that ecological opportunities, reduced competition and sexual selection contributed to the evolution
of large body size in A. kullanderi.
* 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:
** The Academy of Natural Sciences of Philadelphia, 1900 Benjamin Franklin Parkway, Philadelphia, PA 19103,
U.S.A. E-mail:
Apistogramma Regan 1913 is composed of 84
valid species including the one described herein
(but not Apistogrammoides pucallpaensis Meinken,
1965), distributed in cis-Andean river systems
throughout much of tropical and subtropical
South America (Kullander, 2003; Römer, 2006;
Eschmeyer, 2014). Only Crenicichla, with 86 valid
species, is richer among cichlid genera (Kullander,
2003; Eschmeyer, 2014; HRV, unpubl. data).
The modern era of Apistogramma taxonomy
began with Kullanders (1980) monographic re-
vision, which reviewed the history of the genus
and established many of the standards used to
accurately compare and describe its morphologi-
cal diversity. Kullander (1980) recognized 36 valid
species in Apistogramma, 12 of which he newly
described. Based in part on coloration, squama-
tion, fin characteristics and body shape, Kullander
(1980) distinguished nine species groups, two of
them monotypic (A. cacatuoides and A. steindach-
neri). New species descriptions and regional revi-
sions (e. g., Mesa & Lasso, 2011) aside, substantial
contributions to our knowledge of the genus
include popular literature (e. g., Linke & Staeck,
1995; Mayland & Bork, 1997; Stawikowski & Wer-
ner, 1998), and a compendium on Apistogramma
Varella & Sabaj Pérez: Apistogramma kullanderi
and other dwarf cichlids by Römer (2006).
As developed by Kullander (1980, 1986, 1987)
and Kullander & Nijssen (1989), characteristics
of Apistogramma diagnostic in combination are:
epibranchial lobe present and with marginal
rakers; single supraneural bone; long, rod-like
interarcual cartilage; reduced number of ossified
external rakers on first ceratobranchial (commonly
one, occasionally absent or as many as five); gill
rakers laterally on lower pharyngeal tooth plate
(fifth ceratobranchial); two infraorbital bones
between lachrymal and sphenotic; and separate
skin openings for the posterior anguloarticular
canal foramen and anterior-most preopercular
foramen (except anguloarticular canal absent in
some species). Kullander (1987) considered the
last characteristic to be unique among cichlids to
Apistogramma and monotypic Apistogrammoides,
the latter distinguished most importantly by hav-
ing 6-9 (vs. usually 3) anal-fin spines (Kullander,
Kullander (1998) provided phylogenetic evi-
dence for placement of Apistogramma in the tribe
Geophagini (i. e., lobed geophagines), sister to
Gymnogeophagus, and considered monotypic
genera Apistogrammoides and Taeniacara to be most
similar to, if not congeneric with Apistogramma.
In a total evidence approach combining morpho-
logical and molecular data, López-Fernández et
al. (2005) supported a close relationship between
Apistogramma (inclusive of Apistogrammoides) and
Taeniacara, with those two genera consistently
Fig. 1. Apistogramma kullanderi, MZUSP 115057, holotype, 79.7 mm SL, adult, brooding male; Brazil: Pará: tribu-
tary of rio Curuá, rio Xingu basin; a, preserved; and b, alive, just after capture (photograph by José Birindelli).
Ichthyol. Explor. Freshwaters, Vol. 25, No. 3
sister to Satanoperca Günther. Kullander (1987)
had previously noted a striking similarity between
Apistogramma, Apistogrammoides and Satanoperca
with respect to the rod-like shape and articula-
tion of the interarcual cartilage. In a broader
molecular study, López-Fernández et al. (2010)
likewise supported the monophyly of a group
(apistogrammines) composed of Satanoperca sister
to Taeniacara + Apistogramma (inclusive of Apisto-
grammoides), and expanded Geophagini to include
Kullander’s Acarichthyini and Crenicaratini, as
well as genera Crenicichla and Teleocichla.
A recent expedition to Serra do Cachimbo,
Brazil, funded by the All Catfish Species Inventory
(NSF DEB-0315963), discovered a striking new
species of Apistrogramma. It was briefly noted and
figured under the informal name Apistogramma
gigas by Birindelli et al. (2009) and as Apisto-
gramma sp. by Sabaj Pérez (2009), both popular
articles. The objective of this paper is to formally
describe that species and comment on its place-
ment in Apistogramma.
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 typically occurs adjacent to a fin
base and is formed by scales approximately half
Fig. 2. Apistogramma kullanderi, MZUSP 97597, paratype, 58.7 mm SL, adult, brooding female; Brazil: Pará: tribu-
tary of rio Curuá, rio Xingu basin; a, preserved; and b, alive, just after capture (photograph by Mark Sabaj Pérez).
the size of a normal flank scale. The correspond-
ence between coloration and breeding condi-
tions of males and females was determined by
examining, under stereomicroscope, the gonads
of 20 specimens from 37.6 to 74.3 mm SL in an
ontogenetic series. Vertebral counts include the
last half-centrum and were taken from X-rays.
Osteological notes were based on specimens
prepared according to protocol of Datovo &
Bockmann (2010), in which bone and cartilage
are stained and muscle is not digested; those
specimens are indicated in the material examined
as ms (muscle). For illustration, sensory pores
were traced from photographs using Adobe Il-
lustrator CS6. Institutional abbreviations are as
follows: ANSP, Academy of Natural Sciences,
Philadelphia; MZUSP, Museu de Zoologia da
Universidade de São Paulo, São Paulo; NRM,
Naturhistoriska Riksmuseet, Stockholm.
Apistogramma kullanderi, new species
(Figs. 1-3, 6)
Apistogramma sp. Sabaj Pérez, 2009: 43-44 (pho-
tographs of live specimens and habitat).
Apistogramma sp. gigas Birindelli et al., 2009:
13-14 (photographs of live specimens and
Holotype. MZUSP 115057, 79.7 mm SL, adult
male; Brazil: Pará: Altamira: unnamed tributary
of rio Curuá (rio Iriri drainage, rio Xingu basin),
at route BR163 bridge; 8°49'12" S 54°58'11" W;
A. Netto-Ferreira, J. Birindelli, L. Sousa & P.
Hollanda-Carvalho, 22 Jan 2009.
Paratypes. 88 specimens, 14.5-76.0 mm SL. All
from Brazil: Pará: Altamira: rio Xingu basin:
upper rio Curuá (Iriri drainage). ANSP 194896,
17: 4 males, 58.4-74.3 mm SL, 2 females, 55.1-
56.1 mm SL, 9 unsexed or juveniles, 22.1-69.4 mm
SL; 2 ms, 20.7-22.3 mm SL; MZUSP 97584, 16: 2
males, 65.2-76.0 mm SL, 4 females, 50.1-57.7 mm
SL, 10 unsexed or juveniles, 21.2-47.0 mm SL;
rio Escorpião, tributary of upper rio Curuá, at
BR163 bridge; 8°53'54" S 54°59'20" W; J. Birindelli
et al., 29 Oct 2007. – MZUSP 96874, 5: 4 males,
51.8-58.4 mm SL, 1 female 50.5 mm SL; main
channel of rio Curuá immediately above ca. 10 m
falls (sampled after dry down during construc-
tion of PCH Salto Buriti); 8°46'09" S 54°57'02" W;
J. Birindelli et al., 21 Oct 2007. – MZUSP 97597,
14: 6 females, 48.0-58.7 mm SL, 8 unsexed or
juveniles, 22.1-48.5 mm SL; NRM 66406, 11: 5
females, 32.3-53.9 mm SL, 6 unsexed or juveniles,
23.6-42.6 mm SL; same locality as holotype; J.
Birindelli et al., 29 Oct 2007. – MZUSP 101380,
9: 4 males, 37.8-70.5 mm SL, 4 females, 42.3-
47.5 mm SL, 1 juvenile, 20.5 mm SL; NRM 66407,
Table 1. Standard length in millimeters and proportional measurements of the holotype and 45 paratypes of
Apistogramma kullanderi. Caudal-fin length was obtained from 42 specimens only. Holotype measurements in-
cluded in sample range. SD, standard deviation.
holotype range mean SD
Standard length (mm) 79.7 20.5-79.7 47.1
Percents of standard length
Body depth 37.9 34.2-41.3 37.9 1.4
Head length 36.7 35.1-39.8 37.7 1.3
Caudal peduncle depth 17.1 15.1-17.7 16.7 0.6
Caudal peduncle length 12.7 9.9-14.9 11.3 1.0
Last dorsal-fin spine length 12.9 11.6-16.0 13.9 1.2
Pectoral-fin length 27.3 26.1-32.1 29.2 1.4
Pelvic-fin length 39.8 25.0-39.8 29.4 2.6
Caudal–fin length* 32.3 30.2-36.9 32.8 1.9
Head depth 28.9 25.2-32.0 28.6 1.3
Head width 15.0 14.5-17.1 15.7 0.7
Orbital diameter 9.0 9.0-13.9 11.1 1.4
Interorbital width 10.4 8.2-10.9 9.7 0.7
Snout length 14.0 8.3-14.3 11.5 1.7
Preorbital depth 5.3 2.3-6.4 4.8 1.2
Upper jaw length 11.5 9.0-13.0 11.0 0.8
Lower jaw length 14.6 13.5-16.8 15.3 0.8
Varella & Sabaj Pérez: Apistogramma kullanderi
Ichthyol. Explor. Freshwaters, Vol. 25, No. 3
4: 3 males, 39.9-70.7 mm SL, 1 juvenile, 31.8 mm
SL; collected with holotype. – MZUSP 101388,
11, 14.5-35.2 mm SL, unsexed or juveniles; same
locality as ANSP 194896; A. Netto-Ferreira et al.,
22 Jan 2009. – MZUSP 115058, 1 mus, 68.5 mm
SL, male; from one of the collecting sites listed
above for 2009 expedition to rio Curuá (kept in
aquarium prior to preservation).
Diagnosis. Apistogramma kullanderi is diagnosed
by different aspects of coloration found separately
in mature females and males. Mature females
are diagnosed from all congeners by the unique
combination of intense dark pigmentation con-
tinuous along basal portion of dorsal fin and on
ventral surfaces from gular region to anal-fin base
(Figs. 2, 3c-d). Among nominal species of Apisto-
gramma, that pattern is most closely approximated
by mature females of A. baenschi Römer et al.,
2004, distinguished from A. kullanderi by having
dark pigmentation in dorsal fin discontinuous,
alternating with pale regions (see Römer, 2006:
429, bottom figure). Mature males of A. kullanderi
have a coarse, irregular pattern of dark spots and
vermiculations on cheek and opercular series,
and sides with 10-12 dark stripes, each stripe
occupying junction between adjacent scale rows
and separated by paler region central to each
scale (Figs. 1, 3a-b). Among nominal species of
Apistogramma, a similar pattern is found in mature
males of A. rubrolineata Hein et al., 2002 and A. tu-
curui Staeck, 2003. Mature males of A. kullanderi
are distinguished from those species by having
enlarged jaws with large, fleshy lips (vs. jaws and
lips not enlarged), five (vs. four) dentary pores,
and head profile acutely angular with distinct
concavity in interorbital region (vs. profile more
broadly rounded, lacking concavity in interorbital
region). Apistogramma kullanderi is also diagnosed
from all congeners by its maximum size, 79.7 mm
SL for a wild-caught adult male (vs. 65.3 mm
SL among wild-caught congeners). Additional
characteristics useful for identifying A. kullanderi
include: deep body (depth 34.2-41.3 % SL), caudal
fin broad and evenly rounded, first gill arch with
1-4 rakers, most scales in lateral line with distinct
tube (vs. simple perforation), and immature
individuals with dark midlateral band, lacking
midlateral blotch (Figs. 3e-g).
Description. Largest male 79.7 mm SL (Fig. 1),
largest female 58.7 mm SL (Fig. 2). Body deep
(34.2-41.3 % SL, mean 37.9 %), greatest depth
between verticals through second and fifth dorsal-
fin spines. Dorsal and ventral head profiles convex
in juveniles, approximately straight and with
interorbital concavity in adults. Snout relatively
long (8.3-14.3 % SL, mean 11.5 %); posterior bor-
der of maxilla reaching vertical through anterior
margin of orbit in most specimens, falling short
of orbital margin in largest specimens (> 60 mm
SL). Orbit dorsolateral, diameter negatively al-
lometric, decreasing with increase of standard
length; dorsal margin of orbit tangent with con-
tour of head in specimens smaller than 26 mm
SL, becoming more remote from head contour
as standard length increases. Posterior margins
of preopercle and supracleithrum entire; post-
temporal serrations lacking.
Ctenoid scales on flank; cycloid scales on
antero-ventral portion of cheek, preopercle, in-
teropercle, subopercle, opercle (except for some
ctenoid scales on dorsal border of opercular bone),
area anterior of dorsal fin, interpelvic area and
eventually around urogenital papilla. Scales in
E1 row 22 (n = 14) or 23 (30). Cheek fully scaled,
4-5 horizontal scale rows (Fig. 4). Predorsal scales
8-10. Prepelvic area scaled; 3-4 scales anterior to
tips of cleithra, 7-9 scales from tips of cleithra to
pelvic fin. Transverse row with 1 1
2 scales above
and 7 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 13-16/6-9 with frequencies
as follows: 13/6 (1), 13/7 (1), 14/8 (2), 14/9 (2),
15/5 (1), 15/7 (3), 15/8 (12), 15/9 (6), 16/7 (1),
16/8 (8), 16/9* (3). Tube-bearing scales range from
11-16/3-9. Scales between upper lateral line and
first dorsal-fin spine 2 (2), 3* (31), or 4 (7). Scales
between anterior ramus of lateral line and last
dorsal-fin spine 1
2 (4), 1 (12), 1 1
2 (11) or 2* (15).
Fins naked except caudal fin, which is scaled
basally for about one-third its length.
Infraorbital series composed of three bony ele-
ments: lachrymal, infraorbital one and infraorbital
2 + 3 (co-ossification interpreted by presence of
median pore). Infraorbital canal (Fig. 4) 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
Fig. 3. Apistogramma kullanderi, all MZUSP 101380, paratypes; Brazil: Pará: tributary of rio Curuá, rio Xingu ba-
sin; a, 70.7 mm SL, adult male, intermediate stage of maturation; b, 70.5 mm SL, adult brooding male; c, 42.4 mm
SL, adult female, intermediate stage of maturation; d, 42.3 mm SL, brooding female; e, 37.8 mm SL, unsexed
subadult; f, 31.8 mm SL, unsexed juvenile; and g, 20.4 mm SL, unsexed juvenile.
Varella & Sabaj Pérez: Apistogramma kullanderi
Ichthyol. Explor. Freshwaters, Vol. 25, No. 3
end, posterior one shared with anterior opening
of frontal canal. Frontal with four pores plus
coronalis pore.
Dorsal fin XIII.7 (1), XV.6 (1), XV.7* (14),
XV.8 (9), XVI.6 (7), XVI.7 (14). Dorsal-fin spines
with lappets evident in mature males. In holotype
(largest adult male examined), lappets longest
on spines IV-IX, their length approximately
one-third to one-half the length of correspond-
ing spine (except lappet of spine VIII, which is
slightly more than half length of spine); lappets
on posteriormost dorsal-fin spines X-XVI much
shorter, but extending well beyond tips of their
respective spines. Smaller adult males with less
developed lappets, their length approximately
one-third length of corresponding spines IV-VIII
or IX. In females and males < 50 mm SL, lappets
relatively inconspicuous, truncate or rounded.
In four largest males examined, soft dorsal fin
pointed with 4th ray longest and 3rd and 5th rays
nearly as long; tip of 4th ray reaching one-half to
three-quarters length of caudal fin. Soft dorsal fin
also pointed in mature females, but tip extending
only slightly beyond caudal fin base, to approxi-
mately one-fifth the length of caudal fin.
Pectoral fin rounded with 12* (31) or 13 (15)
rays; 6th ray longest, tip almost reaching vertical
through genital papilla. Pelvic fin pointed, first
ray elongated in males; tip of first pelvic-fin ray
reaching base of 3rd soft ray of anal fin in holotype
(79.7 mm SL), base of 3rd anal-fin spine or slightly
beyond in other large males (70.4-76.0 mm SL),
and base of 1st anal-fin spine in smaller males.
In females and unsexed juveniles, pelvic fin
pointed, tip reaching genital papilla or base of
1st anal-fin spine. Anal fin III.7 (10), III.8* (34),
IV.7 (1), IV.8 (1). Soft anal fin pointed in males and
females; 4th ray longest and 5th ray nearly as long;
4th ray extending beyond middle of caudal fin in
males and only slightly beyond caudal-fin base
in females. Caudal fin rounded with 3 procurrent
and 8 principal rays per half.
Jaw teeth unicuspid, erect, cusp strongly ret-
rorse; outer row teeth similar in size or slightly
larger than inner row teeth; 26-28 teeth in upper
right jaw outer hemi-series, 25-28 in upper left,
27-33 in lower right jaw outer hemi-series, 27-32
in lower left. Outer row of teeth occupying almost
entire premaxilla and dentary margins; inner row
of premaxilla expanded as far as outer row in large
specimens but only to one-third of jaw margin in
smaller ones; inner row of dentary expanded to
one third of jaw margin; few teeth forming one
additional, short, middle row near symphysis of
upper and lower jaws in some specimens.
Gill rakers on first epibranchial lobe 5 (9),
6* (24), 7 (10), 8 (2); one gill raker in angle and 1 (6),
2 (30), 3* (9), 4 (1) gill raker(s) on ceratobranchial.
Gill rakers laterally on lower pharyngeal tooth-
plate 10 (3), 11* (2), 12 (17), 13 (16), 14 (7). No tooth
i1 i2
10 mm
1 mm
Fig. 4. Apistogramma kullanderi, MZUSP 115058, 68.5 mm
SL; pattern of sensory lateralis pores on head. AA, an-
guloarticular pore; AN, anterior nasal pore; C, corona-
lis pore; D1-D5, dentary pores; F2-F4, frontal pores;
i1-i3, post-lachrymal infraorbital pores; L, lachrymal
pores; N/F1, pore shared by the nasal and first frontal
lateralis canal; Po1-Po6, preopercular pores.
Fig. 5. Apistogramma kullanderi, MZUSP 97597, 55.6 mm
SL, lower pharyngeal tooth plate, in dorsal view (pos-
terior right horn broken during preparation).
plates on ceratobrachial four. No microbranchio-
spines on gill arches.
Lower pharyngeal tooth-plate dissected from
three specimens (MZUSP 115058, 68.5 mm SL;
MZUSP 97584, 65.3 mm SL; and MZUSP 97597,
55.6 mm SL, Fig. 5): slightly wider than long
(length 90-91 % of width), posterior contour
deeply emarginated in dorsal view, with long
horns; strongly depressed in lateral view but
with deep, thin keel anteriorly; 15-17/16 teeth
in left/right posterior row, 5-7/6-7 teeth in left/
right median row. Medioposterior teeth largest
and more robust, teeth gradually decreasing in
height and more slender rostrally and laterally;
posterior and median teeth bicuspid, lateral and
rostral teeth unicuspid.
Vertebrae 11 + 12 = 23 (1), 11 + 13 = 24 (1), in-
cluding last half-centrum. Hypurals 1-5 separate.
Color pattern in alcohol. Within species of Apisto-
gramma, pigmentation patterns vary considerably
among males, females and juveniles. Some of that
variation stems from different social and behavio-
ral contexts, such as aggressive vs. neutral males
and brooding vs. non-brooding females (Römer,
2006; Rodrigues et al., 2009). The following de-
scriptions are based on wild-caught individuals
preserved at site of capture; sex, maturity and
breeding condition are inferred from comparisons
to other species of Apistogramma, particularly
those described and figured by Römer (2006).
Juveniles and immature specimens (22-45 mm
SL; Figs. 3e-g) with tan ground color, markings
light to dark brown, dorsal half of head and
body slightly darker than ventral half. Brown
preorbital stripe from anterior margin of orbit
to postlabial skin fold, aligned with postorbital
stripe from posterior margin of orbit across dorsal
margin of opercle, and continuing posteriorly as
midlateral band to caudal peduncle. Dark su-
praorbital, interorbital, lachrymal and suborbital
(cheek) stripes generally present, particularly in
smallest specimens (< 25 mm SL). Supraorbital
stripe runs obliquely from dorsoposterior margin
of orbit onto nape; reduced to rounded blotch
near orbit in some specimens. Interorbital stripe
indistinct, broad and irregularly pigmented be-
tween dorsal margins of orbits, but usually with
paired branches extending anteriorly, medial to
nostrils and confluent across snout. Lachrymal
stripe faint, from antero-ventral margin of orbit
to posterior terminus of jaws. Suborbital stripe
usually evident, oblique from just beneath ven-
tral margin of orbit to articulation of sub- and
interopercle, often with brief pale hiatus across
posterior margin of preopercle. Pre-, post-, and
supraorbital stripes dark brown and most distinct;
interorbital, lachrymal and suborbital stripes light
brown, more faint, particularly in larger juveniles.
Body with medium to dark brown midlateral
band finishing just before caudal flexure and sepa-
rate from darker caudal blotch. Midlateral band
occupying entire depth of E1 scale row plus the
adjacent thirds of scales in flanking rows E0 and
E2; band darkest in smallest specimens (< 25 mm
SL), uniformly pigmented or with middle portion
scarcely lighter (i. e., dark pigment less crowded
on central base vs. along margins of E1 scales);
dorsal and ventral limits of band relatively entire,
not jagged. Caudal blotch ovate to vertical rectan-
gular, situated between V3-4 and D3-4 caudal-fin
rays; posterior margin entire or with pale central
About eight shallow, irregularly pigmented
dark brown blotches from nape, along dorsal-
fin base, to caudal peduncle. Dark brown dorsal
blotches transition into about six wide, light to
medium brown vertical bars ending on lower
flank; additional anterior-most bar sometimes
evident on nape to dorsal margin of opercle
where it joins postorbital stripe. Vertical bars 1-4
anterior to anal-fin origin; bars 5 and 6 extending
between soft portions of dorsal and anal fins;
bar 7 on caudal peduncle. Vertical bars distinct in
small juveniles (< 25 mm SL) and larger immature
specimens (perhaps females), scarcely evident
in other immature specimens (perhaps males).
In lattermost case, vertical bars obscured by a
faint horizontal pattern of narrow dusky stripes
formed by a greater concentration of pigment in
dorsal and ventral thirds of each scale; typically,
one dusky stripe evident above midlateral band,
and three such stripes below.
Spinous portion of dorsal fin dusky with
melanophores scattered on spines and mem-
branes; soft portion with dusky blotches loosely
to regularly aligned to form 4-5 vertical bands
separated by pale interspaces. Anal fin similarly
patterned with spinous portion dusky and soft
portion having 3-4 vertical bands. Pectoral fin
hyaline except for dark hairlines outlining each
ray and its branches. Pelvic fin with spine, first
and sometimes second ray, and intervening
membrane(s) dusky with scattered dark pigment;
remaining portion relatively hyaline. Up to four
dusky vertical bands scarcely evident in caudal
Varella & Sabaj Pérez: Apistogramma kullanderi
Ichthyol. Explor. Freshwaters, Vol. 25, No. 3
fin; widest band across fin base, darkest band
occupies distal margin.
Coloration in adult females largely based on
specimens presumed to exhibit pigmentation
pattern of active brooders (Figs. 2, 3d). Ground
color medium brown on dorsal head and flank,
lighter on ventral flank; dark markings on head
and body brownish-black, extensive, dominating
overall pattern. Orbital stripes as described for
juveniles, except lachrymal stripe lacking, other
stripes more intensely pigmented, and suborbital
stripe almost entirely incorporated into dark col-
oration masking lower head. Entire ventral surface
darkly pigmented, brownish-black from gular
region across breast and abdomen to posterior
base of anal fin; dark pigmentation extending
dorsally onto cheek and separately along distal
margin of preopercle and anteroventral portion
of opercle; dark ventral mask joining suborbital
stripe on cheek to form broadly triangular blotch
with apex truncated by orbit.
As in juveniles, brooding females have about
eight shallow, irregular dark brown blotches from
nape, along dorsal-fin base, to caudal peduncle,
and six vertical bars on flanks, posterior to oper-
cle. Unlike juveniles, bars much more intensely
pigmented and confluent with dark pigmentation
masking ventral surfaces. In large female (ANSP
194896, 58.4 mm SL), flank almost completely
brownish-black except for narrow pale windows
separating vertical bars above and below continu-
ous midlateral band. In smaller brooding female
(ANSP 194896, 56.1 mm SL), relatively pale
brown region separating dark dorsal blotches
from vertical bars (vs. dorsal blotches and vertical
bars confluent in larger females). Also in smaller
female, midlateral band lacking, broken into a
series of seven dark blotches aligned with verti-
cal bars. In larger females, continuous midlateral
band persistent, but no more distinct than vertical
bars. In smaller female, dark caudal blotch vertical
rectangular with distinct outline; larger females
with caudal blotch somewhat enlarged, expanded
vertically and posteriorly onto caudal fin, outline
Dorsal and anal fins darkly pigmented except
soft portions of fins becoming pale distally; bands
in soft portion of dorsal fin scarcely evident, bands
lacking from soft portion of anal fin. Pectoral fin
hyaline except for small melanophores scattered
on and along rays. Pelvic fin darkly pigmented,
especially along base and anteriorly, becoming
slightly less so towards distal medial portion.
Caudal fin uniformly dusky, without distinct
Coloration in adult males variable, presum-
ably dependent upon condition at time of cap-
ture (e. g., aggressive vs. neutral, dominant vs.
subdominant); three patterns observed, the first
one most closely resembling that of juveniles
(Figs. 3a-b). Ground color tan to medium brown;
orbital stripes distributed as in juveniles; pre-,
supra-, post- and suborbital stripes conspicuous.
Cheek with additional dark brown narrow stripe
below lachrymal one, extending horizontally from
terminus of jaw to suborbital stripe; small dark
brown spots scattered on remaining portion of
cheek and on opercular series. Dark brown mid-
lateral stripe scarcely evident as such, pigmenta-
tion non-uniform with central portion of E1 scales
relatively pale. Horizontal pattern more conspicu-
ous, consisting of approximately ten narrow dark
brown stripes formed by greater concentration of
dark pigment in dorsal and ventral thirds of each
scale (limits and course of each stripe thereby de-
pendent on scale distribution). Two such stripes
marking dorsal and ventral limits of midlateral
band. Four stripes above midlateral band; two
dorsalmost stripes short, oblique, finishing before
middle of base of spinous dorsal fin; 3rd stripe
involving dorsal third of scales in anterior ramus
of lateral line and finishing below anterior base
of soft dorsal fin; 4th involving ventral limits of
scales in dorsal lateral line and finishing on caudal
peduncle. Four stripes below midlateral band;
three longest extending from pectoral-fin base to
caudal peduncle; 4th ventral to pectoral-fin base,
ending above base of soft anal fin. Scarcely evident
in adult males is conspicuous juvenile pattern
of shallow dark blotches along dorsal-fin base
transitioning into seven wide vertical bars (one
on nape plus six on flank). Caudal spot present,
dark brown, vertical rectangular. Fin pigmenta-
tion similar to juveniles, except vertical banding
pattern in caudal fin inconspicuous; caudal fin
dusky except for rows of pale windows restricted
to membranes, windows most evident along
membranes between the seven central fin rays.
Second pigmentation pattern observed in
mature males (Fig. 1) similar to above with
following exceptions: dark spots on cheek and
opercular series larger, some irregularly joining
each other and/or suborbital stripe to form coarse
vermiculations (thereby obscuring suborbital
stripe, sometimes rendering it inconspicuous),
horizontal pattern more conspicuous due to
56°W 52° W
Fig. 6. Apistogramma kullanderi, MZUSP 97584, 76.0 mm SL; Brazil: Pará: tributary of rio Curuá, rio Xingu basin;
mature male, live specimen photographed just after capture; dark coloration perhaps induced by stress during
capture and/or subsequent photography (photograph by Mark Sabaj Pérez).
Fig. 7. Distributions of Apistogramma kullanderi and other cichlids in upper rio Curuá on Serra do Cachimbo,
Brazil. Apistogramma kullanderi collected at sites 1, 2 (type locality) and 3; Apistogramma upper Curuá collected
at site 2; Apistogramma Treze de Maio collected at site 4; Aequidens aff. rondoni collected at sites 1, 2, 3 and 4.
A, 60 m (downstream) and 40 m (upstream) waterfalls; B, site of 10 m falls and weir for PCH Salto Buriti; C, res-
ervoir; D, major waterfalls on rio Treze de Maio. Brown dashed line approximates escarpment of Serra do
Varella & Sabaj Pérez: Apistogramma kullanderi
Ichthyol. Explor. Freshwaters, Vol. 25, No. 3
greater contrast between darker stripes and paler
ground color, and with two additional stripes
evident (i. e., twelve overall), one from gill cover
to anal-fin origin and second shorter stripe from
gill cover to pelvic-fin origin.
Third pigmentation pattern most extreme
and perhaps induced by stress during capture
and/or subsequent photography (Fig. 6). Sides
entirely black except for pale central portions of
a few scales scattered on flank and those in scale
row above dorsal lateral line. Head also darkly
pigmented, but with some contrast between black
markings (orbital stripes, vermiculations on cheek
and opercular series) and dark brown ground
color. All fins black except pectoral.
Color in life. Descriptions based on adults
presumably in breeding coloration and photo-
graphed live in the field immediately after capture
(Figs. 1b, 2b, 6). Males (Figs. 1b, 6) with ground
color yellow green on head, nape, breast, and up-
permost flank (above pectoral-fin base and along
dorsal fin base), transitioning to shades of aqua-
blue ventrally and posteriorly to caudal peduncle;
bluish hues most evident in regions occupied by
dark vertical bars described for preserved speci-
mens. Dark markings on head and body black,
strongly contrasted with lighter ground color. In
holotype (Fig. 1b), magenta wash completely cov-
ers snout and dark pre- and interorbital stripes.
Magenta coloration also on portions of jaws and
skin between jaws and snout, irregularly on nape
and scales along dorsal fin base, on distal portions
of dorsal and anal fins (particularly on prolonged
soft portions of those fins), and throughout caudal
fin (similar magenta coloration occurs in caudal
fin of wild caught males of A. cacatuoides; see
figure in Mayland & Bork, 1997: 51). Pectoral
fin yellow green. Small dark blotches evident in
basal portions of soft dorsal and anal fins, and on
central basal portion of caudal fin.
A female (Fig. 2b), presumably in brooding
coloration, with ground color gold to yellow or-
ange, lightest in region below midlateral band and
between four posteriormost vertical bars; dark
markings black, four posteriormost vertical bars
prominent, more or less uniformly pigmented
from dorsal to ventral contour of body. Dorsal fin
black except for yellow orange in distal quarter to
half of spinous portion, and distal three-quarters
Fig. 8. Type locality of Apistogramma kullanderi, unnamed tributary of rio Curuá (Iriri drainage), at BR163, bridge,
29 October 2007. (Photographs by Mark Sabaj Pérez).
a b
of soft portion. Anal fin black anteriorly, yellow
orange in posterior half of soft portion. Pectoral
fin hyaline; pelvic fin black. Caudal fin uniformly
yellow orange, appearing slightly darker than
similarly colored portions of dorsal and anal fins.
Similar pattern involving yellow orange ground
color and intense black markings reported for
brooding females of A. baenschi (see figures in
Römer, 2006: 446, 447).
Distribution and habitat. Apistogramma kulland-
eri is known only from the upper rio Curuá and
two of its larger tributaries on Serra do Cachimbo
(Fig. 7), an isolated plateau of the Brazilian (Gua-
poré) Shield. Apistogramma kullanderi has not been
recorded below the uppermost of three waterfalls
marking the Curuás departure from the plateau.
Below the falls, the rio Curuá flows north into the
rio Iriri, a major tributary to the lower rio Xingu
(Amazonas Basin). Tributaries to the upper Curuá
are clear to mild blackwaters; specimens were
captured in shallow, sluggish backwaters shaded
by bank vegetation (Fig. 8).
Etymology. The species is named in honor of
Swedish ichthyologist Sven Oscar Kullander
whose four decades of careful and comprehen-
sive work have transformed the taxonomy of
Neotropical Cichlidae and continue to inspire all
those fascinated with its diversity.
Sexual dimorphism. Apistogramma kullanderi
is sexually dimorphic for fin development, a
condition common among congeners. In mature
males, the pelvic fin is distinctly elongated, its
tip reaching the base of the 3rd soft anal-fin ray in
the largest male examined (vs. reaching anal-fin
origin in mature females), and soft portions of
the dorsal and anal fins are prolonged, reach-
ing more or less midlength of caudal fin. Other
sexually dimorphic fin features generally found
in Apistogramma are membranes between anterior
dorsal-fin spines prolonged as lappets, and cau-
dal fin with prolonged rays affecting lanceolate
or lyreate shapes. In A. kullanderi, the caudal fin
remains shallowly rounded to truncate, and is
not sexually dimorphic. Less clear is the condi-
tion of sexual dimorphism in the anterior dorsal
fin. In the largest males examined, the anterior
membranes are somewhat prolonged relative to
the remaining portion of the spinous dorsal fin,
whereas in mature females such prolongation is
not distinguishable from smaller mature males.
In all mature males of A. kullanderi, the spinous
dorsal fin certainly lacks the extreme lappets
found in species such as A. bitaeniata Pellegrin,
1936, A. cacatuoides Hoedeman, 1951, and A. mac-
masteri Kullander, 1979.
Relationships. Apistogramma kullanderi is not
clearly assignable to any of the species groups
proposed by Kullander (1980, 1986) and Kullander
& Nijssen (1989). Exclusive to members of Kul-
landers A. macmasteri group is the presence of a
dark blotch on the chest in females, a condition
shared with A. kullanderi; however, males of the
A. macmasteri group typically have a taller dorsal
fin with longer lappets. Apistogramma kullanderi
shares the characteristics used to distinguish the
A. regani group (sensu Kullander, 1986: 168), such
as four infraorbital and five dentary pores, mid-
lateral band relatively narrow and not reaching
caudal fin, caudal spot distinct, lateral spot absent,
dorsal-fin lappets short, caudal fin rounded, pre-
pelvic region fully scaled, and anterior dorsal-fin
membranes black; however, the dark chest blotch
is absent from females of the A. regani group (vs.
present in A. kullanderi).
On the basis of a cluster analysis of 116
species-level taxa of Apistogramma (63 nominal
and 53 undescribed), Römer (2006) recognized
three major lineages: A. agassizii, A. regani and
A. steindachneri. Kullanders A. macmasteri and
A. regani species groups are both placed in the
A. regani lineage of Römer, and the composition
of the former species group corresponds well
with Römers A. macmasteri complex. Members of
Kullanders A. regani group are distributed among
multiple complexes within Römers expanded
A. regani lineage. As noted earlier, mature males of
A. kullanderi have 10-12 dark longitudinal stripes,
a condition approximated by A. rubrolineata and
A. tucurui, both members of Römers A. regani
lineage (A. commbrae and A. linkei complexes,
Two characteristics of A. kullanderi are at
odds with its placement in the A. regani lineage
of Römer (2006). As previously noted, mature
females of A. kullanderi share striking similari-
ties in coloration (i. e., ventral surfaces intensely
black) with females of A. baenschi, a member
Römers A. steindachneri lineage placed in either
the A. cacatuoides complex (sensu Römer, 2004) or
Varella & Sabaj Pérez: Apistogramma kullanderi
Ichthyol. Explor. Freshwaters, Vol. 25, No. 3
A. nijsseni complex (sensu Römer, 2006). Secondly,
the large size of mature male A. kullanderi (up to
79.7 mm SL) is most closely approached by that of
A. steindachneri (65.3-75.8 mm SL). Despite those
characteristics, and in the absence of a compre-
hensive cladistic analysis of Apistogramma, we
tentatively consider A. kullanderi to be a member
of the A. regani lineage of Römer (2006), and the
A. regani species group of Kullander (1986).
Biogeography. The rio Curuá departs Serra do
Cachimbo via two major waterfalls approximately
40 meters (upstream) and 60 meters (downstream)
in height and separated by about 50 meters of
river channel (Birindelli & Britski, 2009). Above
the 40 m waterfall is a smaller falls (ca. 10 m)
and site of the weir (i. e. dam) for PCH Salto
Buriti, a run-of-the-river hydroelectric complex
constructed in 2007. The weir impounds the
rio Curuá upstream of the 10 m falls, creating a
reservoir from which water is diverted through
a penstock to power turbines before returning to
the river just downstream of the 60 m waterfall.
The waterfalls, collectively referred to as Salto
do Curuá, apparently isolate the upper Curuá
from the remaining portion of the watershed,
which drains into the rio Iriri, a major tributary
to the lower rio Xingu. Five characiforms were
recently described as endemic to the upper Cu-
ruá: Leporinus guttatus Birindelli & Britski, 2009
(Anostomidae); Jupiaba kurua Birindelli et al.,
2009, Moenkhausia petymbuaba Lima & Birindelli,
2006 (Characidae), and Lebiasina marilynae Netto-
Ferreira, 2012 and L. melanoguttata Netto-Ferreira,
2012 (Lebiasinidae). Apistogramma kullanderi ap-
pears to represent a sixth endemic species.
In 2007 and 2009, fishes were sampled in the
rio Curúa basin at three sites above the 10 m
waterfalls, one site just upstream of the 40 m
falls, and sites downstream of the 60 m falls. One
2007 collection was made during an artificial dry
down of the stretch of the main channel immedi-
ately above the 10 m falls (all water temporarily
re-routed for dam construction). Apistogramma
kullanderi was only recorded from the three sites
above the 10 m waterfalls, one in the upper Curuá
mainstem and two others in separate tributaries
(rio Escorpião and another unnamed). Only two
other species of cichlid have been recorded from
the upper Curuá, and both occur syntopically
with A. kullanderi.
One is tentatively identified as Aequidens aff.
rondoni and occurs in both the upper Curuá (same
three sites as A. kullanderi) and upper rio Treze
de Maio, a tributary of the Curuá. Headwaters
of the rio Treze de Maio similarly arise on Serra
do Cachimbo and are isolated by large waterfalls
from the lower Treze de Maio, which joins the
Curuá downstream of Salto do Curuá. Aequidens
rondoni was described by Miranda-Ribeiro (1918)
from rio do Sangue above Salto Bello and is dis-
tributed in the upper Tapajós basin.
The second species also belongs to the genus
Apistogramma and appears to be an undescribed
member of the A. regani group (sensu Kullander,
1980). Apistogramma upper Curuá was found
only in one of the two tributaries occupied by
A. kullanderi (i. e., type locality). Based on seven
specimens (mature male 46.7 mm SL, immature
male and five unsexed, immature specimens,
26.8-35.0 mm SL), Apistogramma upper Curuá
is easily recognized by having four anal-fin
spines vs. three (n = 44) or rarely four (n = 2) in
A. kullanderi, and by differences in coloration.
The mature male of Apistogramma upper Curuá
lacks the coarse pattern of dark markings on head
and the dark pigmentation in dorsal and anal fins
described for males of A. kullanderi, and has a con-
spicuous pattern of dark bars on caudal fin that
are absent in mature A. kullanderi. Apistogramma
kullanderi also has a conspicuous pattern of stripes
on flanks, whereas the undescribed species has
only two or three longitudinal series of faint
markings in the abdominal region formed by a
greater concentration of dark pigment on central
posterior border of each scale.
A second undescribed species of the Apisto-
gramma regani group (sensu Kullander, 1980)
occurs in the upper rio Treze de Maio, and is
not sympatric with A. kullanderi. It matures at a
much smaller size (ca. 40 mm SL), similar to other
species of Apistogramma. In mature females of
Apistogramma upper Treze de Maio, the dorsal-
fin base presents dark blotches alternating with
pale interspaces (vs. dorsal-fin base entirely dark
in A. kullanderi), and the dark pigmentation on
ventral part of body is restricted to gular region
(vs. covering almost entire ventral surface of body
in A. kullanderi). Males and juveniles of the upper
Treze de Maio species are distinguished by hav-
ing three to five longitudinal series of abdominal
markings formed by greater concentration of dark
pigment on central posterior border of each scale,
a pattern similar to Apistogramma upper Curuá,
but absent from A. kullanderi.
Body size. Apistogramma kullanderi is the largest
known member of the genus with wild-caught
males reaching at least 79.7 mm SL (holotype).
Apistogramma steindachneri (Regan, 1908) is
the next largest congener with males reaching
65.3 mm SL in the wild, and at least 75.8 mm SL
under aquarium conditions (Kullander & Nijssen,
1989: 82). Records of Apistogramma erythrura Stae-
ck & Schindler, 2008 reaching 140 mm SL (e. g.,
Steele & López-Fernández, 2014: table S1) are in
error; that value pertains correctly to TL (60.7 mm)
as a percentage of SL (43.4 mm), reported as 139.8
by Staeck & Schindler (2008: 200, tab. 1).
With respect to body size among Neotropical
cichlids (subfamily Cichlinae), Apistogramma rules
the small extreme. In a recent study of body size
diversity in 498 species of Cichlinae by Steele &
López-Fernández (2014), Apistogramma included
both the smallest species (i. e., A. staecki, 21 mm
SL) and the most species below 36 mm SL (n = 22)
and 50 mm SL (n = 53). Only ten species from
other genera (e. g., Dicrossus, Teleocichla) shared
the small-bodied morphospace occupied by Apis-
Athough body size appeared to be phylo-
genetically constrained within Apistogramma,
it was divergent at a more inclusive level, the
Satanoperca (CAS) clade
of Geophagini (Steele & López-Fernández, 2014).
Body size frequency distributions of the CAS clade
were consistent with three possible optima around
35, 100 and 250 mm SL, with Apistogramma and
Satanoperca corresponding to the first two optima,
respectively. Those two genera are closely related
and form a well-supported clade with Taeniacara
(López-Fernández et al., 2010), yet occupy distinct
areas of body size space (Steele & López-Fernán-
dez, 2014). Body size divergence, under an eco-
logical opportunity model, may help co-existing,
closely related taxa to minimize competition for
limited resources such as food and habitat (Mahler
et al. 2010; Steele & López-Fernández, 2014).
Furthermore, ecological opportunity predicts that
speciation rates and morphological evolution,
such as changes in body size, will be elevated
in a taxon after its colonization of insular areas
unoccupied by competitors (Harmon et al., 2008).
Ecological opportunity may play a role in the
extreme body size of A. kullanderi, known only
from the rio Curuá above Salto do Curuá. Only
two other species of cichlid, Aequidens aff. rondoni
and Apistogramma upper Curuá are known to
co-exist with A. kullanderi in the upper Curuá.
The maximum sizes observed for the species of
Aequidens and Apistogramma upper Curuá are
126.7 and 47.3 mm SL, respectively. The insular
nature of the upper Curuá, coupled with its ap-
parently depauperate cichlid fauna, may provide
ecological opportunities for body size diversifi-
cation in an otherwise small-bodied taxon such
as Apistogramma. Concordantly, large body size
may have evolved in A. kullanderi to minimize
competition with the smaller Apistogramma upper
Curuá, and is presumably constrained to some
degree by the larger species of Aequidens.
In addition to its extremely large body size
relative to congeners, A. kullanderi exhibits male-
biased sexual size dimorphism wherein mature
males are larger than mature females. Within a
taxon, the evolution of larger body size in males
vs. females is generally attributed to sexual
selection mediated by male-male competition
and/or female choice (Woolbright, 1983). Larger
males tend to be more reproductively successful
when mating involves behaviors (e. g., court-
ship displays and nest defense) that are both
energetically costly and incur a greater risk of
predation (Emery, 1968; Knapp, 1995; Johnson &
Hixon, 2011). Within Apistogramma, male-biased
sexual size dimorphism is particularly strong in
polygynous species (Römer, 2006), wherein males
defend spawning territories. We speculate that in
the context of an insular watershed depauperate
of cichlids, ecological opportunities and reduced
competition relax phylogenetic constraints on
body size in Apistogramma, allowing sexual selec-
tion to drive male body size to a larger optimum.
Comparative material. Apistogramma sp. upper Cu-
ruá: MZUSP 95794, 7, 26.8-47.3 mm SL, 2 males and
five immature unsexed specimens, examined by photos;
Brazil: Pará: Altamira: rio Xingu basin: unnamed tribu-
tary of rio Curuá (Iriri drainage), at route BR163 bridge;
8°49'12" S 54°58'11" W.
Apistogramma sp. upper Treze de Maio: MZUSP
97095, 2, 25.0-39.1 mm SL, 1 female, 1 juvenile; MZUSP
101422, 5, 23.8-39.5 mm SL, 1 male, 1 female, 3 unsexed
or juveniles; Brazil: Pará: Altamira: rio Xingu basin: rio
Treze de Maio, a tributary of rio Curuá (Iriri drainage),
upstream of PCH Salto Treze de Maio dam; 8°45'06" S
55°02'05" W.
Aequidens aff. rondoni: all from Brazil: Pará: Al-
tamira: rio Xingu basin. MZUSP 96872, 4, main channel
of rio Curuá immediately above ca. 10 m falls (sampled
after dry down during construction of PCH Salto Bu-
riti); 8°46'09" S 54°57'02" W; J. Birindelli et al., 21 Oct
2007. – MZUSP 97094, 2, 34.4-60.2 mm SL; rio Treze de
Maio, tributary of rio Curuá (rio Iriri drainage), up-
Varella & Sabaj Pérez: Apistogramma kullanderi
Ichthyol. Explor. Freshwaters, Vol. 25, No. 3
stream of PCH Salto Treze de Maio dam; 8°45'06" S
55°02'05" W. – MZUSP 97580, 17 of 22, 58.5-126.0 mm
SL; Brazil: Pará: rio Escorpião, tributary of upper rio
Curuá, at BR163 bridge; 8°53'54" S 54°59'20" W. – MZUSP
97582, 48 of 54, 40.5-126.7 mm SL; unnamed tributay
of upper rio Curuá (rio Iriri drainage), at route BR163
bridge; 8°49'12" S 54°58'11" W.
Other comparative material. Apistogramma agassizi:
MZUSP 7916, 4, 12.4-30.6 mm SL. Apistogramma bitae-
niata: MZUSP 15321, 2, 22.2-33.3 mm SL; MZUSP 46716,
2, 27.2-28.8 mm SL. Apistogramma borelli: MZUSP 25269,
2, 25.4-26.2 mm SL. Apistogramma cacatuoides: MZUSP
46714, 53, 11.8-33.6 mm SL. Apistogramma diplotaenia:
MZUSP 28214, 23, paratypes, 15.2-29.2 mm SL. Apis-
togramma gossei: MZUSP 38982, 1, paratype, 25.5 mm
SL; MZUSP 38991, 1, paratype, 24.3 mm SL. Apistogram-
ma hippolytae: MZUSP 6657, holotype, 30.8 mm SL;
MZUSP 19396-19428, 33, paratypes, 21.8-34.4 mm
SL. Apistogramma inconspicua: MZUSP 38944, 1, para-
type, 23.3 mm SL. Apistogramma iniridae: MZUSP 14888,
1, paratype, 28.1 mm SL. Apistogramma linkei: MZUSP
28726, 8, paratypes, 17.4-25.9 mm SL. Apistogramma
macmasteri: ANSP 127328, 9, 20.7-38.8 mm SL; ANSP
127330, 1, 50.8 mm SL; ANSP 127389, 13, 20.4-45.1 mm
SL. Apistogramma paucisquamis: MZUSP 36952, holo-
type, 29.2 mm SL; MZUSP 42322, 4, paratypes, 17.5-
27.9 mm SL. Apistogramma pulchra: MZUSP 38989, 2,
paratypes, 20.0-26.7 mm SL. Apistogramma regani:
MZUSP 38992, 2, paratypes, 29.4-27.4 mm SL; MZUSP
38967, 6, paratypes, 24.0-27.3 mm SL. Apistogramma
wapisana: MZUSP 100673, holotype, 27.9 mm SL; MZUSP
100674, 1, paratype, 24.8 mm SL.
Thanks to José L. O. Birindelli, Pedro Hollanda-Carval-
ho, Nathan K. Lujan, André L. Netto-Ferreira and Le-
andro M. Sousa for help collecting the type series.
Added thanks to J. L. O. Birindelli for live photograph
of holotype. Thanks to Ricardo Britzke for his assistance
during the development of this study and comments
on the manuscript. Fieldwork was supported in part
by the All Catfish Species Inventory (NSF DEB-0315963,
senior personnel M. Sabaj Pérez); museum work was
supported in part by iXingu Project (NSF DEB-1257813
CoPI M. Sabaj Pérez); visit to ANSP by HRV was sup-
ported in part by ANSP Böhlke Award. H. Varella is
financially supported by FAPESP (grant 2011/14630-0).
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Received 7 February 2014
Revised 26 September 2014
Accepted 5 November 2014
Varella & Sabaj Pérez: Apistogramma kullanderi
... Covain et al. (2016) The Rio Curuá departs Serra do Cachimbo through two major waterfalls approximately 40 m (upstream) and 60 m (downstream) (Fig. 5) in height, separated from each other by about 50 m of river channel. Above the 40 m waterfall, there is a sequence of three smaller waterfalls of approximately 10 m each (Varella & Sabaj Pérez 2014). In the same way, the Rio Treze de Maio leaves the Serra do Cachimbo through a large waterfall with the same name, and then runs for about 20 km, discharging into the Rio Curuá near the village of Cachoeira da Serra downstream to the two major waterfalls of Rio Curuá. ...
... plumbea Sousa et al., 2010;Jupiaba kurua Birindelli et al., 2009;Lebiasina marilynae Netto-Ferreira, 2012;L. melonoguttata Netto-Ferreira, 2012; L. minuta Netto-Ferreira, 2012; Apistogramma kullanderi Varella & Sabaj Pérez, 2014;A. eleutheria Varella & Britzke, 2016;A. ...
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Three new species of Harttia from the Rio Xingu basin in the region of Serra do Cachimbo, Pará State, Brazil, are described. Harttia rondoni n. sp., H. panara n. sp. and H. villasboas n. sp. were collected in the Rio Curuá, a tributary of Rio Iriri, Rio Xingu drainage. Harttia rondoni was additionally recorded from the Rio Fresco basin, a tributary of the middle Rio Xingu. The three new species represent the first records for the genus in this drainage. Harttia rondoni differs from H. panara and H. villasboas by having the abdomen partially covered by plates. Harttia panara differs from H. rondoni and H. villasboas by having a larger body depth and interorbital distance. Harttia villasboas differs from H. rondoni and H. panara by having the anterior profile of the head elliptical in dorsal view and a more elongated snout. Additionally, the 26 species of Harttia described are organized in three species group based on the pattern of abdominal covering.
... This high number of endemic species can be attributed by the high number of headwaters of the Xingu and Tapajós rivers [41]. There, H. rondoni and H. villasboas have a probable geographic barrier due to a series of waterfalls 40-60 m height, over a 50 m stretch, where H. rondoni and H. villasboas occur below and above this set of waterfalls, respectively [41,44]. Despite some karyotype similarities that they share, such as the higher number of m and sm chromosomes, a vicariance event may have facilitated the fixation of another series of chromosomal rearrangements in both groups, including the origin of two distinct, but related sex chromosome systems (a proto or-neo-XY system in H. rondoni and a multiple X1X1X2X2/X1X2Y system in H. villasboas). ...
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The armored Harttia catfishes present great species diversity and remarkable cytogenetic variation, including different sex chromosome systems. Here we analyzed three new species, H. duriventris, H. villasboas and H. rondoni, using both conventional and molecular cytogenetic techniques (Giemsa-staining and C-banding), including the mapping of repetitive DNAs using fluorescence in situ hybridization (FISH) and comparative genomic hybridization (CGH) experiments. Both H. duriventris and H. villasboas have 2n = ♀56/♂55 chromosomes, and an X1X1X2X2 /X1X2Y sex chromosome system, while a proto or neo-XY system is proposed for H. rondoni (2n = 54♀♂). Single motifs of 5S and 18S rDNA occur in all three species, with the latter being also mapped in the sex chromosomes. The results confirm the general evolutionary trend that has been noticed for the genus: an extensive variation on their chromosome number, single sites of rDNA sequences and the occurrence of multiple sex chromosomes. Comparative genomic analyses with another congeneric species, H. punctata, reveal that the X1X2Y sex chromosomes of these species share the genomic contents, indicating a probable common origin. The remarkable karyotypic variation, including sex chromosomes systems, makes Harttia a suitable model for evolutionary studies focusing on karyotype differentiation and sex chromosome evolution among lower vertebrates.
... Although distributed over vast areas, the species are generally restricted to small watersheds or endemic to a single river (e.g., [18][19][20]). Apistogramma species represent the smallest cichlids known, ranging from 2 to 8 cm in size [21,22], with a marked sexual dimorphism, where males are larger and exhibit more exuberant color patterns [18]. ...
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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.
... Microschemobrycon cryptogrammus is only found in the upper Rio Curuá, a left-bank tributary of Rio Xingu formed mostly by rivers draining the Serra do Cachimbo, and this region seems to house several species with restricted distribution (e.g. Birindelli et al., 2009;Netto-Ferreira, 2012;Varella & Sabaj-Pérez, 2014;Garcia-Ayala et al., 2017;Oyakawa et al., 2018). On the other hand, several species inhabiting right-bank tributaries of the Rio Xingu exhibit a wider geographic distribution not only in the Rio Xingu, but also in other drainages from the Brazilian Shield (see the "Central Brazilian Shield" described by Lima & Ribeiro, 2011: 158-160 ...
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A new species of Microschemobrycon with a restricted distribution was recently discovered in the Rio Curuá, Rio Xingu basin, Pará, Brazil. Microschemobrycon cryptogrammus new species can be promptly distinguished from all congeners by the presence of a rounded, dark, conspicuous caudal-peduncle blotch and by a subjacent dark midlateral stripe visible in life. Additionally, the new species can be distinguished from all its congeners, except M. elongatus, by the presence of a longitudinal dark stripe along the lower jaw. The new species can be distinguished from M. elongatus by presenting lateral line with 36–38 pored scales, anterior and posterior nostrils coalescend, dark chromatophores concentrated along the ventral margin of the caudal peduncle and by the presence in life of a dark spot at the base of the dorsal-fin origin.
... 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). ...
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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.
... However, due to its diminutive nature, bony hooks on other than pelvic and anal fins may have been overlooked in several taxa and in fact to be more widespread in Characidae than presently known. The upper rio Curuá presents a high level of fish endemism (Birindelli et al., 2009;Varella & Sabaj Pérez, 2014). Brachychalcinus reisi seems to be another example of a taxon restricted to this system, although a single known specimen from rio Tapajós system (LBP 23315) presents the main diagnostic characters of B. reisi (i.e., a series of dark wavy longitudinal stripes and 7 longitudinal scale rows between dorsal-fin origin and lateral line). ...
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Brachychalcinus reisi, a new species of characid fish, is described from the rio Curuá, tributary of rio Iriri, rio Xingu basin, Serra do Cachimbo, Pará State, Brazil. The new species can be distinguished from its congeners by the presence of a series of longitudinal black wavy stripes on the entire body and by a lower number of longitudinal scale rows between dorsal-fin origin and lateral line (7–8 vs. 8–12). Additionally, the new species differs from B. copei, B. parnaibae, and B. retrospina by the lower number of branched dorsal-fin rays (9 vs.10). This is the first description of a new species of the subfamily since the revisionary study of Stethaprioninae, published almost 30 years ago.
... melanophore and carotenoid-based pigments). Among Neotropical fishes, sexual dichromatism is commonly reported in representatives of the families Cichlidae (Lucena & Kullander, 1992;Kullander & Lucena, 2006;Kullander et al., 2009;Varella & Sabaj Pérez, 2014;Malabarba et al., 2015;Varella, 2016) and Cynolebiidae (Costa, 1998(Costa, , 2006(Costa, , 2007Costa et al., 2012). Sexually dichromatic males of these taxa develop a very bright colour pattern, often making them very popular in the aquarium trade. ...
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Hyphessobrycon myrmex sp. nov., is described from the Rio Formiga, upper Rio Juruena, upper Rio Tapajós basin, Mato Grosso, Brazil. The new species can be distinguished from congeners by having the lower half of the body deeply pigmented with dark chromatophores, chromatophores concentrated above the anal fin and forming a broad, diffuse, dark midlateral stripe and by having a dense concentration of dark chromatophores along unbranched dorsal-fin rays and distal portions of the two or three subsequent branched rays. In life, H. myrmex exhibits a conspicuous sexual dichromatism, with adult males red to orange and females and immatures pale yellow. A list containing 108 sexually dichromatic taxa in six families of Characiformes is provided and the distribution of this poorly known type of dimorphism across the Characiformes is discussed.
... In recent decades numerous new species of the genus Apistogramma Regan, 1913 have been discovered in Brazil, Venezuela, and especially Peru. Some of these forms have been scientifically studied and the vast majority of them described for the first time only in recent years ( BRitzke et al., 2015;kullandeR, 1980, 1986kullandeR & FeRReiRa, 2005;Mesa & lasso 2011a-b;Ready & kullandeR, 2008;RöMeR, 1994RöMeR, , 1997RöMeR & HaHn, 2008, 2013RöMeR et al., 2003RöMeR et al., , 2004aRöMeR et al.,-b, 2006bRöMeR et al.,-d, 2011RöMeR et al., , 2012RöMeR et al., , 2013scHindleR & staeck, 2013;staeck, 1991staeck, , staeck & scHindleR, 2008VaRella & BRitzke, 2016;VaRella & saBaj PéRez, 2014), bringing the total number of valid species to significantly over 90 and making Apistogramma the largest genus among the Neotropical cichlids. However, there are still several forms awaiting formal description. ...
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Apistogramma megastoma sp. n. is described based on a total of 18 specimens from small forest streams in the Departamento Loreto, Peru, tributaries of the Río Jutai in the border area between Peru and Brazil south and west of the city of Leticia (Colombia) (near 04°12′ S / 70°06′ W). Apistogramma megastoma sp. n., which may be confused only with A. barlowi, is distinguished from all other Apistogramma species by the combination of: noticeably disproportionately large head; exceptionally massive jaws, lyrate densely vertically banded caudal fin, extended dorsal-fin membranes in males; in aggressive females sooty head pattern and up to 8 series of small black dashes on flanks; small round caudal spot; and, exceptional for Apistogramma, maternal mouth-brooding behaviour. Distinguished from A. barlowi, by differences in colour pattern and by higher number of scale rows on cheeks. Apistogramma megastoma sp. n. inhabits in small fastflowing streams.
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Supplement to Cichlid Atlas Volume 1: This volume includes a revised and updated key for the identification of live dwarf cichlids of the genus Apistogramma, information about evolution and biogeography of that genus, results of field observational work in Amazonia, as well as scientific descriptions of 4 new species and a new genus of dwarf cichlids. Detailed information is given about ecology, maintenance, and breeding. Together with the chapter printed in Volume 1, the bibliography included in Cichlid Atlas 2 covers nearly all relevant citations about Neotropical cichlids. The volume is illustrated with over 400 drawings, 125 maps, and appoximately 1280 colour photographs. 1st English language edition published December 15th 2006.
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Apistogramma erythrura sp.n. is described from the drainage of the lower rio Mamoré in the province of Beni in Bolivia. It is most similar to Apistogramma trifasciata, but differs from this species by the complete absence of any abdominal markings below the lateral band. It can be distinguished from all the other described Apistogramma species by the following combination of characters: only two postlacrymal infraorbital lateralis canal pores, five dentary lateralis canal pores, a broad lateral band with even borders that widens posteriorly to a height of at least two and a half scales and extends into the caudal fin Without a caudal spot. Adult males usually with a partly or even completely bright red colouration of their rounded caudal fin, extremely produced anterior dorsal-fin lappets and prolonged pelvic-fin filaments reaching the posterior end of the anal-fin base.|
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First comprehensive treatise of known South American dwarf cichlids until 1998. Contains 1400 photographs, 150 drawings, key for identification, results of research on sex determination, distribution, ecology, behaviour, maintenance and breeding in captivity, descriptions of several species of the genera Apistogramma, Apistogrammoides, Biotoecus, Crenicara, Crenicichla, Dicrossus, Laetacara, Mazarunia, Mikrogeophagus, Nannacra, Taeniacara, Teleocichla, and an extended bibliography. 1st English language edition.
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Body size is an important correlate of life history, ecology and distribution of species. Despite this, very little is known about body size evolution in fishes, particularly freshwater fishes of the Neotropics where species and body size diversity are relatively high. Phylogenetic history and body size data were used to explore body size frequency distributions in Neotropical cichlids, a broadly distributed and ecologically diverse group of fishes that is highly representative of body size diversity in Neotropical freshwater fishes. We test for divergence, phylogenetic autocorrelation and among-clade partitioning of body size space. Neotropical cichlids show low phylogenetic autocorrelation and divergence within and among taxonomic levels. Three distinct regions of body size space were identified from body size frequency distributions at various taxonomic levels corresponding to subclades of the most diverse tribe, Geophagini. These regions suggest that lineages may be evolving towards particular size optima that may be tied to specific ecological roles. The diversification of Geophagini appears to constrain the evolution of body size among other Neotropical cichlid lineages; non-Geophagini clades show lower species-richness in body size regions shared with Geophagini. Neotropical cichlid genera show less divergence and extreme body size than expected within and among tribes. Body size divergence among species may instead be present or linked to ecology at the community assembly scale.
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Lebiasina marilynae n. sp., L. melanoguttata n. sp. and L. minuta n. sp. são descritas das cabeceiras do rio Curuá, na Serra do Cachimbo, Pará, Brasil, e representam os únicos membros de Lebiasininae no escudo brasileiro, até o momento. Uma relação próxima entre estas espécies é proposta com base em: 1) presença de um par de forames por onde passa o ramo palatino do nervo facial, uma modificação única em Lebiasinidae e aparentemente em Characiformes, 2) a expansão do osso extra-escapular, 3) a ausência da faixa secundária, e 4) os lobos da nadadeira caudal com comprimento similar. Lebiasina marilynae difere ainda de seus congêneres pela faixa primária estendendo-se da ponta do focinho à porção distal do pedúnculo caudal, a presença de duas séries de manchas escuras paralelas à faixa primária, e a superfície dorsal do mesetmoide arredondada. Lebiasina melanoguttata e Lebiasina minuta também diferem de seus congêneres pela ausência da faixa primária e da mancha caudal, e pela presença de três séries longitudinais de manchas escuras na base das escamas das séries 3-5. Lebiasina melanoguttata difere de Lebiasina minuta pela ausência de uma mancha escura na base dos raios médios da nadadeira dorsal, presença de dentes cônicos no segundo infra-faringobranquial, a coloração avermelhada dos olhos e nadadeiras e pelas manchas escuras não coalescendo (vs. mancha escura presente na nadadeira dorsal; o segundo infrafaringobranquial sendo edêntulo; olhos escuros em tom oliváceo e coloração amarelada do corpo e nadadeiras; e as manchas escuras nas escamas das séries longitudinais 3 e 4 coalescendo no ponto onde escamas longitudinais adjacentes se sobrepõem). A ocorrência de espécies de Lebiasininae no escudo brasileiro é discutida e o padrão de distribuição das espécies descritas é comparado com o de outras espécies endêmicas da Serra do Cachimbo, uma área de alta biodiversidade isolada do restante da bacia amazônica.
López-Fernández, H., Honeycutt, R. L., Stiassny, M. L. J. & Winemiller, K. O. (2005). Morphology, molecules, and character congruence in the phylogeny of South American geophagine cichlids (Perciformes, Labroidei). —Zoologica Scripta, 34, 627–651. Phylogenetic relationships among the Neotropical cichlid subfamily Geophaginae were examined using 136 morphological characters and a molecular dataset consisting of six mitochondrial and nuclear genes. Topologies produced by morphological and combined data under parsimony were contrasted, congruence among different partitions was analysed, and potential effects of character incongruence and patterns of geophagine evolution on phylogenetic resolution are discussed. Interaction of morphological and molecular characters in combined analysis produced better resolved and supported topologies than when either was analysed separately. Combined analyses recovered a strongly supported Geophaginae that was closely related to Cichlasomatinae. Within Geophaginae, two sister clades included all geophagine genera. Acarichthyini (Acarichthys+Guianacara) was sister to the ‘B clade’, which contained the ‘Geophagus clade’ (‘Geophagus’steindachneri+Geophagus sensu stricto, and both sister to Gymnogeophagus) as sister to the ‘Mikrogeophagus clade’ (Mikrogeophagus+‘Geophagus’brasiliensis), and in turn, the Geophagus and Mikrogeophagus clades were sister to the crenicarine clade (Crenicara+Dicrossus) and Biotodoma. The second geophagine clade included the ‘Satanoperca clade’ (Satanoperca+Apistogramma and Taeniacara) as sister to the ‘Crenicichla clade’ (Crenicichla+Biotoecus). Several lineages were supported by unique morphological synapomorphies: the Geophaginae + Cichlasomatinae (5 synapomorphies), Geophaginae (1), Crenicichla clade (3), crenicarine clade (1), the sister relationship of Apistogramma and Taeniacara (4) and of Geophagus sensu stricto and‘Geophagus’steindachneri (1), and the cichlasomine tribe Heroini (1). Incorporation of Crenicichla in Geophaginae reconciles formerly contradictory hypotheses based on morphological and molecular data, and makes the subfamily the most diverse and ecologically versatile clade of cichlids outside the African great lakes. Results of this study support the hypothesis that morphological differentiation of geophagine lineages occurred rapidly as part of an adaptive radiation.