Gymnogeophagus taroba (Teleostei: Cichlidae), a new species from the río Iguazú basin, Misiones, Argentina

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Abstract
The Gymnogeophagus setequedas group is based on our results composed of three endemic species of which one, G. taroba sp. n., is described as a new species. The three species are diagnosable from each other and from other species of Gymnogeophagus by stable differences in several morphological characters among which the best are found in coloration patterns. Body and head shapes and meristic characters show lesser differentiation but several are also clearly diagnostic in the G. setequedas group. The G. setequedas group is strongly structured allopatrically. The prime candidates for this fragmentation and speciation are the origins of the waterfalls on the individual tributaries. The largest of the waterfalls, the famous Cataratas del Iguazú, with a height of 72 m, separate G. taroba sp. n. from its closest relatives G. che and G. setequedas. The original 28 m high Urugua-í falls separate G. che from G. setequedas. Gymnogeophagus setequedas is separated from G. che and G. taroba by large rapids (about 65m in total elevation above the río Paraná) and a former fall on the Acaray river and by the 45 m high Monday falls located a few km from the mouth of the Monday into the río Paraná just opposite the mouth of the Iguazú. Resumen. En base a nuestros resultados, el grupo Gymnogeophagus setequedas esta compuesto por tres especies endémicas, de las cuales, G. taroba sp. n., se describe como una nueva especie. Las tres especies son diagnosticables entre sí y de otras especies de Gymnogeophagus por diferencias estables en varios caracteres morfológicos, entre los cuales los mejores son los patrones de coloración. La forma del cuerpo, la cabeza y los caracteres merísticos muestran una menor diferenciación, pero algunos también son claramente diagnósticos en el grupo G. setequedas. El grupo G. setequedas está fuertemente estructurado alopátricamente. La causa principal para esta fragmentación y especiación es la formación de cascadas en los afluentes. La más grande de las cascadas, la famosa Catarata del Iguazú, con una altura de 72 m, separa G. taroba sp. n. de sus parientes más cercanos G. che y G. setequedas. El salto original del Urugua-í de 28 m de altura, separa G. che de G. setequedas. Gymnogeophagus setequedas está separada de G. che y G. taroba por grandes rápidos (unos 65 m de elevación total sobre el río Paraná) y un primer salto en el río Acaray y por otro salto de 45 m de altura en el río Monday localizado a unos pocos kilómetros de la desembocadura del Monday en el río Paraná justo enfrente de la desembocadura del Iguazú. palabras clave. Cíclidos, endemismo, morfología, Cuenca del Plata, Cuenca del río Paraná HISTORIA NATURAL Tercera Serie Volumen 7 (2) 2017/5-22
HISTORIA NATURAL Tercera Serie Volumen 7 (2) 2017/5-22 5
HISTORIA NATURAL
Tercera Serie Volumen 7 (2) 2017/5-22
ISSN 0326-1778 (Impresa)
ISSN 1853-6581 (En Línea)
Jorge Cascioa1,2, Adriana Almirón1, Lubomir Piálek3 and Oldřich Říčan3
1Universidad Nacional de La Plata, Facultad de Ciencias Naturales y Museo,
División Zoología Vertebrados, Paseo del Bosque (1900), La Plata, Buenos Aires, Argentina.
jrcas@fcnym.unlp.edu.ar; aalmiron@fcnym.unlp.edu.ar
2Comisión de Investigaciones Cientícas de la Provincia de Buenos Aires, Argentina.
3University of South Bohemia, Faculty of Science, Department of Zoology,
Branišovská 31 (CZ-370 05), České Budějovice, Czech Republic.
lubomir.pialek@prf.jcu.cz; oldrich.rican@prf.jcu.cz
Gymnogeophagus taroba (TELEOSTEI: CICHLIDAE),
A NEW SPECIES FROM THE RÍO IGUAZÚ
BASIN, MISIONES, ARGENTINA
Gymnogeophagus taroba (Teleostei: Cichlidae),
una nueva especie de la cuenca del río Iguazú, Misiones, Argentina
HISTORIA NATURAL Tercera Serie Volumen 7 (2) 2017/5-22
6
CasCiotta J., almirón a, Piálek l. and Říčan o.
Abstract. The Gymnogeophagus setequedas group is based on our results composed of three
endemic species of which one, G. taroba sp. n., is described as a new species. The three species
are diagnosable from each other and from other species of Gymnogeophagus by stable dierences
in several morphological characters among which the best are found in coloration paerns. Body
and head shapes and meristic characters show lesser dierentiation but several are also clearly
diagnostic in the G. setequedas group. The G. setequedas group is strongly structured allopatrically.
The prime candidates for this fragmentation and speciation are the origins of the waterfalls on the
individual tributaries. The largest of the waterfalls, the famous Cataratas del Iguazú, with a height
of 72 m, separate G. taroba sp. n. from its closest relatives G. che and G. setequedas. The original 28 m
high Urugua-í falls separate G. che from G. setequedas. Gymnogeophagus setequedas is separated from
G. che and G. taroba by large rapids (about 65m in total elevation above the río Paraná) and a former
fall on the Acaray river and by the 45 m high Monday falls located a few km from the mouth of the
Monday into the río Paraná just opposite the mouth of the Iguazú.
Key words. Cichlid shes, endemism, morphology, La Plata River basin, Paraná river basin
Resumen. En base a nuestros resultados, el grupo Gymnogeophagus setequedas esta compuesto por
tres especies endémicas, de las cuales, G. taroba sp. n., se describe como una nueva especie. Las tres
especies son diagnosticables entre sí y de otras especies de Gymnogeophagus por diferencias estables
en varios caracteres morfológicos, entre los cuales los mejores son los patrones de coloración. La
forma del cuerpo, la cabeza y los caracteres merísticos muestran una menor diferenciación, pero
algunos también son claramente diagnósticos en el grupo G. setequedas. El grupo G. setequedas
está fuertemente estructurado alopátricamente. La causa principal para esta fragmentación y
especiación es la formación de cascadas en los auentes. La más grande de las cascadas, la famosa
Catarata del Iguazú, con una altura de 72 m, separa G. taroba sp. n. de sus parientes más cercanos
G. che y G. setequedas. El salto original del Urugua–í de 28 m de altura, separa G. che de G. setequedas.
Gymnogeophagus setequedas está separada de G. che y G. taroba por grandes rápidos (unos 65 m de
elevación total sobre el río Paraná) y un primer salto en el río Acaray y por otro salto de 45 m de
altura en el río Monday localizado a unos pocos kilómetros de la desembocadura del Monday en el
río Paraná justo enfrente de la desembocadura del Iguazú.
Palabras clave. Cíclidos, endemismo, morfología, Cuenca del Plata, Cuenca del río Paraná
HISTORIA NATURAL Tercera Serie Volumen 7 (2) 2017/5-22 HISTORIA NATURAL Tercera Serie Volumen 7 (2) 2017/5-22
A New SpecIeS Of THe Gymnogeophagus
7
INTRODUCTION
The genus Gymnogeophagus inhabits the
La Plata basin, Laguna Los Patos system, the
Tramandaí basin and one record of G. balza-
nii is from the Guaporé basin (Malabarba et
al., 2015; Loureiro et al., 2016). Gymnogeopha-
gus is currently diagnosed by the presence
of a forward directed spine on the top of
the rst dorsal pterygiophore and the loss
of bony supraneurals (Reis and Malabarba,
1988). The species of the genus Gymnogeopha-
gus are informally included in two species
group, the G. gymnogenys species group,
characterized by mouthbrooding reproduc-
tive strategy and the G. rhabdotus species
group with substrate brooding (Loureiro et
al., 2015). The later group includes G. rhab-
dotus (Hensel, 1870) from río Uruguay and
Laguna dos Patos basins; G. meridionalis Reis
and Malabarba, 1988 with a wide geographi-
cal distribution including río Paraná, Uru-
guay and Río de la Plata basins; G. setequedas
Reis, Malabarba and Pavanelli, 1992 from
the middle río Paraná basin; G. che Cascioa,
Gómez and Toresanni, 2000 restricted to the
arroyo Urugua-í, río Paraná basin; and G. ter-
rapurpura Loureiro, Zarucki, Malabarba and
González Bergonzi, 2016 from lower río Uru-
guay, Río de la Plata coastal drainages and
Atlantic Ocean coastal drainages in Uruguay.
Cascioa et al. (2016) reportered for the rst
time specimens of Gymnogeophagus from the
río Iguazú basin above the Iguazú falls. The
specimens were preliminarily identied as G.
a. setequedas due to their closest similarity to
G. setequedas. Several diagnostic dierences
were however noted and the aim of this pa-
per is thus to describe these specimens as a
new endemic species of Gymnogeophagus.
MATERIAL AND METHODS
The Unied Species Concept (de Queiroz,
2007) was employed in the present study.
Consistent morphological dierences be-
tween morphogroups were used as a proxy
for lineage independence in the absence of a
molecular phylogeny which is the focus of a
parallel study.
Genetic sequences from type and non-
type specimens are included in the present
study and follow the GenSeq nomenclature
of Chakrabarty et al. (2013). Genomic DNA
was extracted from ethanol-preserved n
tissue using the JETQUICK Tissue DNA
Spin Kit (Genomed, Germany) following
standard protocol. The primers and reac-
tion conditions of polymerase chain reac-
tion (PCR) amplication for all loci are as
in Říčan et al. (2008). Each PCR reaction
volume of 25 µl contained 12.5 µl of Combi
PPP Master Mix (Top-Bio, hp://www.top-
bio.cz), 1.5 µl of each primer (10 pmol/µl),
and 1 µl of extracted DNA. PCR reactions
were performed in a Bioer XP Thermal Cy-
cler and PCR products were puried using
the JETQUICK PCR Purication Spin Kit
(Genomed). Sequencing reactions were per-
formed following standard protocol with
the use of the same primers as in Říčan et al.
(2008), and the products were analysed in an
ABI 3730XL automated sequencer (Applied
Biosystems; both steps done by Macrogen
Inc., Korea). Contiguous sequences of the
gene segments were created by assembling
DNA strands (forward and reverse) using
GENEIOUS v. 11.0.2 (hp://geneious.com,
Kearse et al., 2012) and were also translated
into protein sequences to check for possible
stop codons or other ambiguities.
Measurements and counts were taken as
described by Reis and Malabarba (1988)
and Malabarba et al. (2015), except body
depth taken at dorsal n origin. Measure-
ments were taken as straight line distances
with digital caliper to 0.1 mm on left side of
specimen. Morphometric characters are ex-
pressed as percents of the standard length
(SL) except for subunits of the head which
are recorded as percents of the head length
HISTORIA NATURAL Tercera Serie Volumen 7 (2) 2017/5-22
8
CasCiotta J., almirón a, Piálek l. and Říčan o.
(HL). Scale row nomenclature follows Kul-
lander (1996). Specimens were cleared and
counterstained (c&s) following Taylor and
Van Dyke (1985). Vertebral counts include
the last half centrum. In the descriptions,
the number of specimens is indicated in
parentheses and values of the holotype are
indicated by an asterisk. All material came
from Argentina except where noted. Non-
type specimens are listed in geographical
distribution, and mapped in distribution
maps, but were not used in species de-
scriptions. Institutional abbreviations are
as listed in Ferraris (2007), except IBIGEO
(Instituto de Bio y Geociencias del NOA-
Ictiología, Salta, Argentina).
RESULTS
Taxonomy
Family Cichlidae Bonaparte, 1835
Genus Gymnogeophagus Miranda Ribeiro,
1918
Gymnogeophagus taroba, new species (Figures
1-7; Table 1)
Gymnogeophagus a. setequedas Cascioa et
al., 2016 (photo of live holotype, rst men-
tion of the species)
All type and non type material from Argen-
tina, Misiones Province, río Iguazú basin,
above falls.
Holotype. MLP 11258, 1 ex., 86.9 mm SL.,
Parque Nacional Iguazú (PNI), boca del
arroyo Ñandú (25°42’10.9”S, 54°25’31.3”W),
Cascioa et al., December 2015, (Figure 1).
Paratypes. MLP 11259, 12 ex. (1 c&s), 18.8-
80.9 mm SL, arroyo Lobo (25º42’57”S-
54º05’59”W), Cascioa et al., November
2001. MLP 11260, 19 ex., 48.8-84.7 mm SL,
PNI, río Iguazú in Pasarela a Garganta
del Diablo (25°41’47.5”S, 54°26’20.4”W),
Cascioa et al., October, 2015. MLP 11138,
6 exs., 59.1-90.7 mm SL., same data as
holotype. MLP 11261, 2 ex., 80.9-83.8
mm SL., PNI, arroyo Yacuí in route 101
(25°40’50.2”S, 54°10’05.8”W), Cascioa et
al., October, 2015. IBIGEO-I 449, 7 ex., 64.0-
93.5 mm SL., PNI, arroyo Yacuí in camping
(25°35’41.50”S, 54°08’55.51”W), Říčan et al.,
February 2012 (Figures 1-7).
Non type material. MLP 11262, 4 ex., 65.2-
81.2 mm SL., arroyo Deseado in camping
(25° 40’ 16”S-53° 55’ 59”W), Říčan et al., No-
vember 2007. MLP 11263, 10 ex., 29.6-52.5
mm SL., PNI, bañado de Ope (25°41’26.4”S,
54°27’16.1”W), Gómez, November 1985.
MLP 11264, 3 ex., 75.7-93.0 mm SL., Parque
Nacional Iguazú (PNI), boca del arroyo
Ñandú (25°42’10.9”S, 54°25’31.3”W), Říčan
et al, November, 2016. MLP 11265, 7 ex. (2
c&s) 49.3-85.3 mm SL., same data as holo-
type. MLP 11266, 3 ex., 38.6-89.0 mm SL, ar-
royo Lobo (25º42’57”S, 54º05’59”W), Říčan
et al., November 2007.
Diagnosis. Gymnogeophagus taroba is distin-
guished from all species of Gymnogeophagus
in the G. gymnogenys group by the posses-
sion of 23 to 25 E1 scales and the absence
of a well developed adipose hump in adult
males. Gymnogeophagus taroba is most eas-
ily distinguished from all species in the G.
rhabdotus group (G. terrapurpura, G. rhab-
dotus, G. meridionalis, G. setequedas and G.
che) by the pigmentation of the dorsal n.
Figure 1 - Gymnogeophagus taroba, male, holotype,
MLP 11258, 86.9 mm SL., Parque Nacional Iguazú,
boca del arroyo Ñandú (25°42’10.9”S, 54°25’31.3”W).
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A New SpecIeS Of THe Gymnogeophagus
9
The spiny section of the dorsal n lacks any
markings vs. various types of markings. The
soft section has long and thin whitish light
blue lines parallel to rays vs. wide stripes
in G. rhabdotus vs. a combination of round-
ish spots at the base and stripes of various
length and width distally in G. meridionalis,
G. terrapurpura, G. setequedas and G. che. The
anal n also has long and thin whitish light
blue lines parallel to rays vs. spoed in G.
terrapurpura vs. a combination of round-
ish spots at the base and stripes of various
length and width distally in G. rhabdotus,
G. meridionalis, G. setequedas and G. che. In
G. setequedas only the posterior half of the
anal n has markings, the anterior half is
immaculate. Within the G. setequedas group
the following additional characters sepa-
rate G. taroba: opalescent lines on posterior
part of body and on caudal peduncle nar-
row, start behind the midlateral blotch and
are not found on the dorsum above the up-
per lateral line vs. wide, start in front of the
midlateral blotch and are found on the dor-
sum in G. setequedas and G. che; midlateral
blotch usually conspicuous and dominant
(as in G. che) vs. inconspicuous in G. seteque-
das; opalescent markings on head large and
dominant, present on opercular series (as in
G. che) vs. small and inconspicuous, usually
absent from opercular series in G. setequedas;
body scales or their centers are grey (as in
G. setequedas) vs. scale centers on at least the
posterior part of body and caudal peduncle
distinctly reddish-brown in G. che; caudal
n with best developed white longitudinal
lines and most pronounced dierence be-
tween a yellow to orange central portion of
the n and red dorsal and ventral margins.
Besides the color paerns, G. taroba can be
distinguished from G. setequedas by a longer
snout (44.4-53.7 vs. 30.8-44.7 % of HL) and
from G. che by deeper body (40.2-46.1 vs
37.3-40.7 % of SL).
Description. Meristic and morphometric
data are summarized in Table 1. Body com-
pressed, moderately elongated. Predorsal
and dorsal proles convex; decreasing in
depth to caudal peduncle. Caudal peduncle
rectangular, deeper than long, ventral pro-
le slightly concave. Body contour slightly
convex between lower lip and last anal-n
ray, with straight segment between pelvic
and anal ns. Snout pointed, nearly straight
dorsally and ventrally in lateral aspect; short
and anteriorly rounded in dorsal aspect. Eyes
close to dorsal prole of head; eye near mid-
dle of head length. Interorbital area slightly
concave. Posterior tip of maxilla not reach-
ing vertical through anterior margin of eye.
Mouth terminal, jaws isognathous. Body
scales ctenoid. Opercle scales ctenoid; scales
on preopercle small and cycloid. Cheek
scales cycloid in 4 rows. Other unpaired ns
without scales. Scales in E1 row 23(4*), 24(15),
25(6). Scales in transverse row 4/8(11*), 4/9(3),
5/8(10), 5/9(1). Upper lateral line scales 15(3),
16(12*), 17(7), 18(2), 19(1). Lower lateral line
scales 6 (1), 7(3), 7+1(1), 7+2(1), 8(6), 8+1(2),
8+2(1), 9(9*), 9+1(1). Dorsal n XII,10(1);
XII,11(3); XIII,9(2); XIII,10(10*); XIII,11(6);
XIV,9(2); XIV,10(1). Anal n III,7 (6); III,8(13*);
III,9(5), III,10(1). Caudal-n squamation not
reaching middle of n. Dorsal-n origin an-
terior to vertical line through posterior bony
margin of opercle, posterior end of depressed
n surpassing caudal-n base. Pectoral-n
edge surpassing the anal-n origin. Anal-n
edge surpassing caudal-n base. Caudal n
truncate or slightly concave.
Gill rakers externally on rst gill arch 9-11
(3). Premaxillary and dentary outer row teeth
slightly movable, inner ones fully depress-
ible. Premaxillary ascending process longer
than dentigerous one. Premaxilla with 14-17
(3) unicuspid teeth on outer row, larger than
inner ones. Four teeth rows near symphysis.
Dentary with 17-18 (3) unicuspid teeth on
outer row, 4 rows near symphysis. Lower
pharyngeal tooth plate with unicuspid and
bicuspid anteriorly curved teeth, those of
posterior and medial one to three rows larg-
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CasCiotta J., almirón a, Piálek l. and Říčan o.
Figure 2 - Gymnogeophagus taroba, lower pharyngeal
tooth plate in occlusal view, MLP11265, 83.4 mm SL.
Figure 3- Gymnogeophagus taroba, coloration of live specimens immediatelly after capture. (A-B) IBIGEO-I 449,
paratype specimens, 87.8 mm, 79.8 mm. (C-D) MLP uncat., Říčan et al., February 2014, data as holotype.
er than remaining ones (Figure 2). Upper
pharyngeal tooth plate with unicuspid and
bicuspid teeth. Frashed zone bearing one
concavity with small unicuspid teeth. Two
to four patch of unicuspid teeth on fourth
ceratobranchial. Vertebrae 26-27, 13+13 (1),
13+14 (1), 14+13 (1) (3 c&s).
Color in life. Background color of dorsal
region of body light olivaceous with ve
indistinct diuse vertical bars (Figure 3).
Black quadrangular mid-lateral spot at
third vertical bar, surrounded by bright,
yellow-green area in some specimens.
Mid-lateral blotch usually conspicuous
and dominant, it can be extended dorsal-
ly to the base of dorsal n. Five horizon-
tal bright light blue narrow bands from
behind the black mid-lateral blotch to
caudal-n base. Ventral portion of body
and head whitish; isthmus and branchio-
stegal region black in adult males and fe-
males during reproductive season. Head
with black bar traversing eye from nape
to ventral edge of preopercle. Bright light
blue elongated or rounded dots usually
present on cheeks, usually forming one
interrupted nearly straight line from tip
of maxilla to upper portion of preopercle
with additional dots ventrally. Most speci-
mens with several bright light blue dots on
the opercle and one on the upper part of
rst lateral line scale. Dorsal n (Figures
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A New SpecIeS Of THe Gymnogeophagus
11
ures 3, 7) background yellow, orange to
light green with upper and lower margin
light purple, centre of n covered with nu-
merous elongated hyaline spots, arranged
in horizontal series between rays. Pectoral
n hyaline, light yellow. Pelvic n green to
ochre, with rst rays black; several whitish
light blue stripes surrounded by dark red
between rays.
3-5) with background light ochre; spiny
portion inmaculate, soft portion with long
and thin opalescent and whitish light
blue lines parallel to rays. Dorsal-n bor-
der black in reproductive adults. Anal-n
(Figures 3, 6) background yellow to ochre
with mostly long whitish light blue nar-
row bands, parallel and diagonal to rays,
surrounded by dark red. Caudal-n (Fig-
Figure 4 - Coloration of the spinous portion of the dorsal n in the G. setequedas group: (A-C) Gymnogeophagus
taroba, paratypes; (D-E) G. setequedas; (F-H) G. che.
HISTORIA NATURAL Tercera Serie Volumen 7 (2) 2017/5-22
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CasCiotta J., almirón a, Piálek l. and Říčan o.
Figure 5- Coloration of the soft-rayed portion of the dorsal n in the G. setequedas group: (A-C) Gymnogeophagus
taroba, holotype (A) and paratypes; (D-F) G. setequedas; (G-I) G. che.
Figure 6 - Coloration of the anal n in the G. setequedas group: (A-C) Gymnogeophagus taroba, holotype (A)
and paratypes; (D) G. che; (E-F) G. setequedas.
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A New SpecIeS Of THe Gymnogeophagus
13
Figure 7- Coloration of the caudal n in the G. setequedas group: (A-C) Gymnogeophagus taroba, holotype (A)
and paratypes; (D-E) G. che; (F-G) G. setequedas.
Figure 8 - Gymnogeophagus che, coloration of live specimens immediatelly after capture, non-type specimens:
(A) MLP 11296, 83.0 mm, Casciotta et al., 2010; (B) MLP11297, 98.5 mm, Říčan et al., 2016; (C) MLP 11296,
68.1 mm, Casciotta et al., 2010; (D) MLP11297, 89.4 mm, Říčan et al., 2016.
HISTORIA NATURAL Tercera Serie Volumen 7 (2) 2017/5-22
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CasCiotta J., almirón a, Piálek l. and Říčan o.
Color in alcohol. Background color of dor-
sal region of body light brown to gray with
ve to six diuse and darker bars (Figure
1). Black quadrangular mid-lateral spot at
third vertical band, surrounded by nar-
row light areas in some specimens. Ventral
portion of body and head whitish to light
brown; isthmus and branchiostegal region
black in most adult males and females
during reproductive season. Head with
black bar traversing eye from nape to ven-
tral edge of preopercle. Dorsal n back-
ground light gray; elongated hyaline spots
in soft portion. Dorsal n border black in
reproductive adults. Anal-n background
light gray with elongated hyalines spots.
Caudal-n background light gray entirely
covered with numerous elongated hyaline
spots, arranged in horizontal series be-
tween rays. Pectoral n hyaline; pelvic n
dusky, with the rst three rays and mem-
branes black.
Sexual dimorphism. Genital papilla in
males slender and longer than those of fe-
male. Some males with laments in dorsal,
anal and pelvic-ns. Pelvic ns of males
usually longer than those of females (2.3-
2.8 vs. 2.7-3.1 in SL).
Geographic distribution. Gymnogeophagus
taroba is endemic to the lower río Iguazú
basin above the Iguazú falls (Figure 10).
Habitat. Gymnogeophagus taroba is found
predominantly in pools and lentic stretch-
es of smaller streams with predominantly
muddy or sandy booms (Figures 11-12).
The type locality, boca del arroyo Ñandú
has a stone substrate covered with mud
(Figure 11). Some specimens (e.g. MLP
11260, 19 ex.) have also been collected in
pools between rock-gardens in the Iguazú
main channel (Pasarela a Garganta del Dia-
blo, Figure 13; pasarela a salto San Martín).
These pools are ecologically very similar
to the small tributary streams with similar
stony substrates and muddy booms.
Figure 9- Gymnogeophagus setequedas, coloration of live specimens immediatelly after capture, non-type specimens:
(A-B) Paraguay, tributary of Rio Monday above Salto Monday, 14.11.2009, 25°41’44.5”S, 55°05’09.7”W; (C)
Paraguay, tributary of Rio Acaray, 31.8.2016, 25°23’31.6”S, 55°25’26.2”W; (D) Paraguay, tributary of Rio Acaray,
30.8.2016, 25°07’53.3”S, 55°57’51.4”W.
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A New SpecIeS Of THe Gymnogeophagus
15
Figure 10 - Distribution of the G. setequedas group in Middle Paraná river basin. Colored
localities show distribution of the G. setequedas group and material examined in the present
study. Stars show type localities. Red: G. taroba, green: G. setequedas and violet: G. che.
Inset photos show two of the major waterfalls in the area (Cataratas del Iguazú, Salto Monday)
and localition of the study area within South America.
Figure 11 - Type locality of G. taroba, mouth of the arroyo Ñandu into the Iguazú river, Parque
Nacional Iguazú, Iguazú river basin (25°42’10.9”S, 54°25’31.3”W), December 2014.
HISTORIA NATURAL Tercera Serie Volumen 7 (2) 2017/5-22
16
CasCiotta J., almirón a, Piálek l. and Říčan o.
Figure 12- Paratype locality of G. taroba, pool in the arroyo Yacuí, Parque Nacional Iguazú, Iguazú river
basin (25°35’41.50”S, 54°08’55.51”W), February 2012.
Figure 13- Paratype locality of G. taroba, pools below Pasarela a Garganta del Diablo, Parque Nacional
Iguazú, Iguazú river basin (25°41’47.5”S, 54°26’20.4”W), December 2014.
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Etymology. The specic epithet taroba
refers to Tarobá, a warrior, and refers to a
legend of the Kaingang people; a noun in
apposition. The Kaingang were the origi-
nal rst inhabitants of the present prov-
ince of Misiones in Argentina and the
southern Brazilian states of Paraná, Santa
Catarina and Rio Grande do Sul and also
the southeastern state of São Paulo. Their
language and culture is quite distinct from
the neighboring Guaraní. The Kaingang
language is classied as a member of the
Gê language family. The legend tells that
at the beginning of time, the río Iguazú
was inhabited by a huge and monstrous
serpent called Mboi, a guardian god. The
Kaingang sacriced a beautiful young
maiden every year to Mboi. When Tarobá
meets Naipí, chosen for the sacrice, he
rebells against the elders of the tribe who
refuse to release her. Tarobá and Naipí try
to escape in a canoe by the river. Mboi be-
comes furious and brakes the course of the
river forming the Iguazú falls, catching the
lovers. Naipí is transformed into one of the
rocks of the falls, perpetually punished by
the turbulent waters, and Tarobá is turned
into a palm tree on the bank of the water-
fall. Mboi lives submerged in the Garganta
del Diablo, from where he watches over the
lovers, preventing them from joining again.
However, on sunny days and as a bridge of
love, the rainbow overcomes the power of
Mboi by rejoining Naipí and Tarobá.
Genseq-2 CYTB. IBIGEO-I 449; GenBank
accession number MG681194.
Genseq-3 CYTB. MLP 11262; GenBank ac-
cession number MG681195.
Genseq-3 CYTB. MLP 11264; GenBank ac-
cession number MG681196.
Morphological variation in the G. seteque-
das group
The G. setequedas group can be divided
into three morphospecies that are diagnos-
able by stable dierences in several mor-
phological characters (Figures 1, 3-7). The
best diagnostic characters in the G. seteque-
das group are found in coloration paerns.
The spinous portion of the dorsal n (Fig-
ure 4) ranges from completely unpaerned
(G. taroba), through only lightly paerned
and limited to the posterior part with short
triangular white spots (G. setequedas), to
heavily paerned throughout with long,
wide and dominant white bands (G. che).
The soft-rayed portion of the dorsal n
(Figure 5) is paerned in all species but also
diagnostic ranging from very light paern-
ing made of long thin lines following the
rays (G. taroba), through more distinct and
wider lines distally and more pronounced
spots proximally on the n (G. che), to
much more dominant blotches dispersed
all the way to the distal margin of the n in
more than the anterior half of the soft por-
tion with a much more reduced number of
lines in the posterior part (G. setequedas).
The anal n pigmentation (Figure 6) is
also diagnostic. All species have a paerned
anal n with the least paerning found in
G. setequedas, where it is limited to only the
posterior portion of the n, while G. che and
G. taroba have the whole n paerned. In G.
che the anterior portion is covered by dots
and the posterior part by dots at the base
and thick lines distally (the posterior part
is thus paerned as in G. setequedas), while
in G. taroba the whole anal n is paerned
by long white lines (that follow the n rays)
distally margined by red coloration in the
largest specimens. The red coloration in the
anal n is also found in the other species
but never to such extent as in G. taroba and
never arranged in clearly dened lines.
The caudal n coloration (Figure 7) is
least diagnostic among the ns but G.
HISTORIA NATURAL Tercera Serie Volumen 7 (2) 2017/5-22
18
CasCiotta J., almirón a, Piálek l. and Říčan o.
taroba has the caudal n most heavily pat-
terned by white longitudinal lines and has
the most pronounced dierence between a
yellow to orange central portion of the n
and red dorsal and ventral margins (the
red coloration covers about four rays in
both the dorsal and ventral margins).
Noticeable dierences are also found
in the coloration of the body and the head
(Figures 1, 8-9). Gymnogeophagus setequedas
is dierent from the other species in that its
midlateral blotch is always very diuse and
by far not so much well developed. The blue
opalescent pigmentation on the head (face)
is variable in its development but is invari-
ably least developed again in G. setequedas,
made of usually small and inconspicuous
dots which are absent from the opercular se-
ries. The opalescent lines on posterior part
of body and on caudal peduncle are wide in
G. setequedas and G. che but are narrow in G.
taroba. These opalescent lines start behind
the midlateral blotch and are absent on the
dorsum above the upper lateral line in G.
taroba while in G. setequedas and G. che they
start in front of midlateral blotch and are
conspicuous on the dorsum above the upper
lateral line. Body scales or their centers are
grey in G. taroba and in G. setequedas while in
G. che scale centers on at least the posterior
part of body and caudal peduncle are dis-
tinctly reddish-brown.
Body and head shapes are quite variable
within the G. setequedas group with much of
the variation being intraspecic. However,
several of these shape characters are clearly
diagnostic and distinguish some of the spe-
cies. These characters include body depth (G.
taroba vs. G. che) and snout length (G. taroba
vs. G. setequedas) (Table 1 and Reis et al., 1992
and Cascioa et al., 2000). The lower pharyn-
geal jaws are on the contrary not diagnos-
tic showing most variation intraspecically
(see Description) and varying in degree of
robustness and numbers of enlarged teeth
(in one to three medial rows) (Figure 2).
Variation in meristic characters is also
mostly intraspecic but G. che (Table 1 and
Reis et al., 1992 and Cascioa et al., 2000) can
be distinguished from G. setequedas and G.
taroba by a modally higher count of scales
along lateral line (26 vs. 24), of lower lateral
line scales (10 vs. 8), and marginally of dor-
sal n spines modal count (14 vs. 13).
DISCUSSION
The G. setequedas group is based on our
results composed of three species that are
diagnosable by stable dierences in several
morphological characters among which the
best are found in coloration paerns, namely
in the coloration of the dorsal n, anal n,
and to some extent also in body and head
coloration. Such dierentiation in live col-
oration paerns is typical for this colourfull
genus (Reis & Malabarba, 1988). The detailed
dierentiation in the coloration of unpaired
ns has recently been noted also in several
other Gymnogeophagus species where it was
also used to support the diagnoses of several
newly described species (Malabarba et al.,
2015; Loureiro et al., 2016). Body and head
shapes and meristic characters show lesser
dierentiation with a much larger propor-
tion of the variation being intraspecic as
in most other Gymnogeophagus species (see
above refs.). However, several of the shape
and meristic characters are clearly diagnostic
in the G. setequedas group.
Species of the G. setequedas group appear
to have a wide ecological valence since they
are found in a wide variety of habitats and
are common species of cichlids. Gymno-
geophagus setequedas is a relatively common
species found throughout eastern Paraguay
in the Paraná tributaries except for the drain-
ages emptying into the Itaipú reservoir and
the reservoir itself both in Paraguay (OŘ,
pers. obs.) and in Brazil (Paiz et al., 2017)
where the species has declined and is now
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A New SpecIeS Of THe Gymnogeophagus
19
rare. Paiz et al. (2017) listed G. setequedas as a
threated species (in Brazil) with a rheophilic
ecology but that is a misunderstanding de-
rived from too much focus on the Itaipú area
in Brazil. In Paraguay Gymnogeophagus
setequedas is a relatively common species
outside the Itaipú area, is predominantly
found in lentic habitats (hence is denitely
not rheophilic as also its body morphol-
ogy reveals) and is even common in and
around Paraguayan reservoirs (e.g. Embalse
de Yguazu). The marked decline of popu-
lations of G. setequedas in the Itaipú area is
more likely the result of the unprecedented
pressure from introduced non-native sh
species and the changed ecology of the res-
ervoir and of the tributaries (Agostinho et al.,
2004, 2005, 2016; Baumgartner et al., 2012) in
the Itaipú area. Gymnogeophagus che is com-
mon throughout the Urugua-í basin in all
types of habitats including in the articial
Urugua-í reservoir. Gymnogeophagus taroba
is based on our knowledge most common
in small auents of the Iguazú river with
habitats and localities that are slow owing
to standing (i.e. pools in small tributaries
and mouths of streams) with soft types of
booms (mud and sand). Some of our speci-
mens (e.g. MLP 11260, 19 ex.) have also been
collected in pools between rock-gardens in
the Iguazú main channel (see Habitat abo-
ve). Paiz et al. (2017) listed G. taroba (as G.
setequedas) only from two localities appar-
ently within the Iguazú river proper (one
above and one below the Iguazú falls). No
habitat information or precise description of
the localities was however given; nor does
the NUP catalogue (hp://www.splink.org.
br/form?lang=pt&collectioncode=NUP) for
these specimens. Based on our records the
localities of Paiz et al. (2017) are also most
likely pools or slow-owing sections of the
river.
The G. setequedas group is strongly struc-
tured allopatrically and made of three en-
demic species. The prime candidates for this
fragmentation and speciation are the origins
of the waterfalls on the individual tributar-
ies (Figure 10). The largest of the waterfalls
are the famous Cataratas del Iguazú, one
of the largest waterfalls in the world with a
2.7-kilometres-long ledge and a height of 72
to 78 m, located 25 km before the Iguazú en-
ters the río Paraná. The Cataratas del Iguazú
separate G. taroba from its closest relatives G.
che and G. setequedas. The 28 m high Urugua-
í falls located eight km from the conuence
with the Paraná river (in 1989 replaced by a
hydroelectric dam) just south of the Iguazú
separate G. che. Gymnogeophagus setequedas
is separated from G. che and G. taroba by a
former fall (of unknown height, that is now
also replaced by a hydroelectric dam) and
by large rapids (about 65m in total elevation
above the río Paraná) on the Acaray river
and by the 45 m high Monday falls (Saltos
del Monday) on the Monday river located
seven km from its mouth into the río Paraná
just opposite the mouth of the Iguazú (and
hence the Cataratas del Iguazú).
Comparative Material
Gymnogeophagus balzanii: Misiones
Province: MLP 8617, 1 ex., 106.9 mm,
arroyo Don Lorenzo, Departamento Ca-
pital. Formosa Province:
IML 452, 1 ex,
57.6 mm, río Paraguay. IML 304, 3 exs.,
50.3-61.5 mm. Salta Province: IML 440, 1
ex., 91.5
mm, San Martín. Chaco Provin-
ce: MLP 7925, 1 ex.,
96.4 mm, Resistencia
.
Corrientes Province: MLP
8616, 7 exs.,
48.9-134.2 mm, riacho Carrizal, Bella
Vis-
ta. MLP 8615, 6 exs., 81.7-169.0 mm, la-
guna cerca
de la escuela Nº 12, camino
Bella Vista-San Roque. MACN
7038, 1
ex., 54.8 mm, río Santa Lucía. MACN
7036, 2
exs., 57.4-63.6 mm, riacho Carrizal,
Bella Vista. MACN
6401, 1 ex., 91.4 mm,
Esteros del Iberá. Entre Ríos
Province
:
MLP 8631, 1 ex., 66.7 mm, Salto Gran-
HISTORIA NATURAL Tercera Serie Volumen 7 (2) 2017/5-22
20
CasCiotta J., almirón a, Piálek l. and Říčan o.
de.
Santa Fé
Province
: MLP 8380 1 ex., 9
mm, laguna
Vargas, Departamento San
Javier. MLP 7292, 2 exs.,
53.8-124.0 mm,
río Colastiné. MLP 8375, 1 ex., 62.0
mm,
río Santa Fé. MLP 7268, 1 ex., 83.1mm,
madrejón
Flores. MLP 6693, 1 ex., 74.7
mm, Parque Belgrano.
Brasil: MSNG
7683 (holotype), 79.5
mm, Villa María,
río Paraguay, Mato Grosso. Paraguay:
ZMB17109, 1 ex., 77.5 mm, Asunción.
ZMB 17108, 2 exs.
52.0-76.4 mm, Asunción.
Gymnogeophagus che: Misiones Province,
río Paraná basin: MLP 8747, (holotype),
91.9 mm, arroyo Urugua-í y ruta provinci-
al 19 frente a la pasarela en parque provin-
cial Islas Malvinas. MLP 8748, (paratypes),
5 exs., 67.1-96.3 mm, arroyo Urugua-í en
Isla Palacios. MLP 8749, (paratype) 1 ex.,
71.3 mm, arroyo Uruzú
y ruta provinci-
al 19, Parque provincial Islas Malvinas.
MACN 8220, (paratype) 3 exs., 89.0-101.0
mm, arroyo Urugua-í y ruta provincial
19. MLP 8750, (paratype)
1
ex. 60.9 mm,
arroyo Urugua-í, en el establecimiento
«Alto
Paraná». MLP 8751, (paratypes) 3
exs., 40.0-86.7
mm, arroyo Uruzú y ruta
provincial 19, Parque provincial Islas
Malvinas. MLP
8752, (paratypes) 2 exs.,
68.5-88.6 mm, arroyo Grapia, auente
del arroyo Urugua-í, 10 km al NE de la
ruta provincial 18. MLP 8758, (paraty-
pes) 17
exs., 15.6-21.0 mm arroyo Urugu-
a-í y ruta
12.
MLP11296
, 7 exs.,62.7-83.0
mm, Embalse Urugua-í at Policia la-
custre. MLP 11297, 2 exs., 84.9-98.5 mm,
arroyo Falso Urugua-í.
Gymnogeophagus
gymnogenys: Brasil: MLP11248 1 ex.,(c&s),
68.8 mm, Brasil, RS, São
Sebastião do Caí,
río Caí. Gymnogeophagus labiatus: Brasil:
MLP11246, 1 ex.,
98.6 mm SL., Brasil, RS,
Sto. Antonio da Patrulla, Vila N. S. Mon-
te Serrat, río Dos Sinos. Gymnogeophagus
meridionalis: MLP 8633, (paratypes) 12
exs.,
42.5-68.5 mm, Corrientes Province,
pond in road Bella Vista to San Roque, near
school Nº12. MLP 8404, 29 exs., 22.0-71.0
mm, Buenos Aires Province, río Salado ba-
sin, laguna Chascomús. MLP 4748, 3 exs.,
58.0-67.2 mm, Buenos Aires Province, Río
de La Plata basin, arroyo Pereyra.
Gymno-
geophagus sp. (since G. terrapurpura was
diagnosed by colour paern in specimens
alive (Loureiro et al., 2016) it is impossible
to decide if all these samples are G. terra-
purpura, G. meridionalis or both species).
Misiones Province: MLP 6853, 1 ex., 37.6
mm, arroyo Piray Mini.
Corrientes Provin-
ce: MLP 8634,
1
ex., 40.6
mm, riacho Carri-
zal, Bella Vista. Entre
Ríos Province: MLP
3346, 5 exs., 22.3-51.2 mm, laguna Mazaru-
ca. MLP 3309, 2 exs., 57.7-73.9 mm, Concor-
dia. MLP 6082, 2 exs., 68.4-78.8 mm, Entre
Ríos,
no other data
. MLP 7684, 2 exs., 28.0-
33.0 mm, río Nogoyá, cerca de Tres Bocas,
Departamento de Gualeguaychú. Santa Fé
Province: MLP 8382, 2 exs.
55,6-63,2 mm,
laguna Setubal.
Buenos Aires Province:
MLP 7684, 21 exs., 26.3-39.0 mm, Delta del
Paraná.. MLP 8636, 6 exs., 30.5-
45.8 mm,
cantera de Los Talas, Berisso. MLP 5633, 2
exs., 43.1-67.9 mm, Magdalena. MLP 4748,
3 exs., 57.9-68.7 mm, arroyo Pereyra. MLP
6306, 7 exs., 51.0-91.1
mm, cantera de Los
Talas, Berisso. MLP 8637, 2 exs.,
19.5-26.2
mm, arroyo Villoldo, Punta Indio. MLP 8638,
2 exs., 18.5-24.7 mm, arroyo Castelli, Punta
Indio.
Gymnogeophagus rhabdotus: Bra-
sil: MLP11247,
1
ex., 61,9 mm,
RS, Brasil,
Passo de Taquara, Gravataí, col: Reis et
al., 5/7/82. Uruguay: MLP11249, 2 exs.,
(c&s) 58.6-66.7 mm, río Uruguay, basin,
arroyo Catalán Grande.
Gymnogeopha-
gus setequedas: Paraguay: MHNG 2518.19
(holotype), 1 ex., 80.3 mm, reservorio río
Iguazú
en Juan E. O‘Leary, Provincia el
Alto Paraná. MHNG 2269.49 (paratypes)
6 exs. (1 ex. c&s) 12.1-72.4
mm, reservorio
río Iguazú en Juan E. O‘Leary; Provincia
el Alto Paraná. MHNG 2027 92-95 (para-
types) 4
exs., 44,7-64.7 mm, Alto Paraná:
Itabo-Guazu. MHNG
2027.68-69 (para-
HISTORIA NATURAL Tercera Serie Volumen 7 (2) 2017/5-22 HISTORIA NATURAL Tercera Serie Volumen 7 (2) 2017/5-22
A New SpecIeS Of THe Gymnogeophagus
21
types) 2 exs., 57,4-67.8 mm, Provincia
de
Alto Paraná, río Acaray, near del Ciudad
del Este.
MCP 14636 (paratype), 1 ex., 68.8
mm, Alto
Paraná, Juan E. O‘Leary, área de
inundación del río Iguazú. MCP 1437 (pa-
ratype) 1 ex., 96.8 mm, Alto Paraná, Juan
E. O‘Leary, arroyo Iguazú, sistema
do río
Paraná. Gymnogeophagus cf. setequedas:
MLP11250, 2 exs., 93.1-99.8 mm, Entre
Ríos Province, río Paraná basin, Parque
Nacional Pre-Delta, Tapera de Chano.
Gymnogeophagus terrapurpura: MLP
11227, 1 ex., 62.0 mm, Entre Ríos Provin-
ce,
Municipio Villa Elisa, balneario munici-
pal, arroyo Perucho Verna.
ACKNOWLEDGEMENTS
We thank Soledad Gouric for the picture
of the LPJ and the authorities of Ministerio
de Ecología y Recursos Naturales Renovab-
les de la Provincia de Misiones and Admini-
stración de Parques Nacionales for the awar-
Holotype Range Mean SD
Standard length (mm)) 86.9 59.1-90.7
Body depth 43.4 40.2-46.1 43.3 1.55
Head length 35.9 32.6-37.4 34.8 1.15
Dorsal-n base 58.6 53.3-59.0 56.1 1.54
Pectoral-n length 38.9 33.4-40.5 37.1 1.59
Caudal peduncle length 13.3 13.3-16.6 15.1 0.84
Caudal peduncle depth 16.1 13.3-16.1 14.8 0.73
Eye diameter* 29.8 26.4-31.4 28.6 1.37
Interorbital width* 32.7 26.6-34.1 30.5 2.22
Pre-orbital length* 27.9 23.2-30.7 27.2 1.99
Snout length* 49.4 44.4-53.7 49.7 2.07
Upper jaw length* 34.6 30.2-37.7 34.0 2.30
ded shing permits. Financial support was
provided by Comisión de Investigaciones
Cientícas de la provincia de Buenos Aires
(CIC), Facultad de Ciencias Naturales y Mu-
seo (UNLP) and Administración de Parques
Nacionales.
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marked with *, which are percentages of head length.
HISTORIA NATURAL Tercera Serie Volumen 7 (2) 2017/5-22
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Recibido: 06/11/2017 - Aceptado: 18/12/2017 - Publicado: 29/12/2017
  • Article
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    A new mouth breeder species of Gymnogeophagus is described from a tributary of the río Uruguay. It is distinguished from most species of the genus by the presence of hypertrophied lips, and from G. labiatus and G. pseudolabiatus by the color pattern. The presence of successive allopatric species of the Gymnogeophagus gymnogenys clade inhabiting the tributaries of the río Uruguay is discussed.
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    Gymnogeophagus jaryi, new species, is described from Southern tributaries of the Middle Paraná basin in Misiones. It can be distinguished from all other members of the genus, except from G. australis and G. caaguazuensis, by the presence of a hyaline to grey anterior portion of the dorsal fin. Gymnogeophagus jaryi differs from G. caaguazuensis by a longer caudal peduncle, caudal fin not lyrate, central portion of scales on dorsal portion of trunk light iridescent blue and by white spots in soft portion of dorsal fin in adult males, and from G. australis by the light iridescent blue coloration of central portion of scales on the dorsal portion of trunk and tail, and by the lack of scales on the soft portion of the dorsal fin. Additionally, it can be diagnosed by the following unique combination of characters: 10–11 dorsal-fin branched rays, 27–30 E1 scales, absence of lips thickening, and, in males, by the possession of a hump in adults, caudal fin not lyrate, presence of large white spots forming transversal stripes distally and in anterior area of the dorsal fin’s soft portion, central area of scales on the dorsal portion of the trunk light iridescent blue, lack of scales on the base of the dorsal fin’s soft portion, absence of a conspicuous and oblique dark band from the eye to the anterior border of the head, anterior portion of dorsal fin hyaline to grey, scales of the midlateral spot each bearing a semicircular light blue blotch, head hump starting at the horizontal through the eyes, concave anterior profile in lateral view, base of unpaired fins yellow, and whitish hyaline spots on caudal fin. The new species, based on mtDNA phylogeny, is the sister species of G. caaguazuensis from the Paraguay basin and is closely related to G. australis.
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    We provide cytogenetic data for the threatened species Gymnogeophagus setequedas, and the first record of that species collected in the Iguaçu River, within the Iguaçu National Park’s area of environmental preservation, which is an unexpected occurrence for that species. We verified a diploid number of 2n = 48 chromosomes (4sm + 24st + 20a) and the presence of heterochromatin in centromeric and pericentromeric regions, which are conserved characters in the Geophagini. The multiple nucleolar organizer regions observed in G. setequedas are considered to be apomorphic characters in the Geophagini, whereas the simple 5S rDNA cistrons located interstitially on the long arm of subtelocentric chromosomes represent a plesiomorphic character. Because G. setequedas is a threatened species that occurs in lotic waters, we recommend the maintenance of undammed environments within its known area of distribution.
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    Describimos una nueva especie de Gymnogeophagus con incubación de sustrato, en base a caracteres cromáticos. La nueva especie puede ser diferenciada de las restantes especies del género por la exclusiva pigmentación de la aleta dorsal, la que consiste de bandas oblicuas de color celeste sobre fondo rojo en la porción espinosa y una combinación de manchas redondas, elípticas, o alargadas azul brillantes sobre fondo rojo en la porción blanda. Además puede ser distinguida de las otras especies de Gymnogeophagus por la siguiente combinación de caracteres: banda horizontal azul iridiscente situada por encima de la línea lateral superior discontinua en la región humeral, manchas azules brillantes redondeadas sobre fondo rojo anaranjado en la aleta anal, bandas horizontales azul brillante en el cuerpo conspicuas. La nueva especie habita una amplia gama de hábitats de agua dulce en el sector inferior de la cuenca del río Uruguay, los drenajes costeros del Río de la Plata y drenajes costeros del Océano Atlántico en Uruguay.
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    Gymnogeophagus is a Neotropical cichlid genus distributed in the río Paraguay, rio Paraná and rio Uruguay drainages and also in the coastal rivers of Uruguay and southern Brazil. Its monophyly is supported by two derived features: the absence of supraneurals and the presence of a forward spine in the first dorsal-fin pterygiophore. Herein, five new species of Gymnogeophagus are described from middle to upper tributaries of the rio Uruguay drainage and from the rio Negro. All these new species belong to a clade, which includes G. gymnogenys, easily recognized by sharing two synapomorphies: the absence of an oblique bar between the dorsal border of the eye and the nape, and the possession of a black bar originating in the dorsal contour near the dorsal-fin origin and directed downward and backward on the dorsum. A key to all species of Gymnogeophagus is provided.
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    Gymnogeophagus che, a new species of the family Cichlidae frori the Paranii river basin (Percifonnes, Labroidei). Gyn~~togeophagu~ che, new species, is described from the Paranh river basin in northeastern Argentina. This specics can be distinguished fiom its congeners by the following combination of characters: Absence of small rounded spots on dorsal, anal, and caudal fms, few hyaline stripes on caudal fin, low body depth and head profile, and ascending arm of the premaxilla bearing a wide sqmphyseal articular facet.
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    A checklist of Recent and fossil catfishes (Order Siluriformes) is presented, summarizing taxonomic literature published through 2005. From 4624 nominal species group names and 810 genus group names, 3093 species are recognized as valid, and are distributed among 478 genera and 36 families. Distributional summaries are provided for each species, and nomenclatural synonymies, including relevant information on all name-bearing types, are included for all taxa. One new name is proposed herein: Clariallabes teugelsi, as a replacement for Clarias (Allabenchelys) dumerili longibarbis David & Poll, 1937, which is preoccupied by Clarias longibarbis Worthington, 1933, but has been treated as a valid species of Clariallabes by Teugels. Acrochordonichthys melanogaster Bleeker, 1854, is designated as type species of Acrochordonichthys Bleeker, 1857, inasmuch as no earlier valid designation has been found. A new genus Pseudobagarius, is proposed for the "pseudobagarius group" of species formerly placed in Akysis. The status of 228 species group names remains unresolved and 31 names based on otoliths ascribed to catfishes are listed but not placed into the checklist. The current emphasis given to catfish taxonomy at present is likely to result in a dramatic increase in the total number of valid taxa as well as major changes in the membership of some of the higher level taxa recognized here.
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    An improved and expanded nomenclature for genetic sequences is introduced that corresponds with a ranking of the reliability of the taxonomic identification of the source specimens. This nomenclature is an advancement of the "Genetypes" naming system, which some have been reluctant to adopt because of the use of the "type" suffix in the terminology. In the new nomenclature, genetic sequences are labeled "genseq," followed by a reliability ranking (e.g., 1 if the sequence is from a primary type), followed by the name of the genes from which the sequences were derived (e.g., genseq-1 16S, COI). The numbered suffix provides an indication of the likely reliability of taxonomic identification of the voucher. Included in this ranking system, in descending order of taxonomic reliability, are the following: sequences from primary types - "genseq-1," secondary types - "genseq-2," collection-vouchered topotypes - "genseq-3," collection-vouchered non-types - "genseq-4," and non-types that lack specimen vouchers but have photo vouchers - "genseq-5." To demonstrate use of the new nomenclature, we review recently published new-species descriptions in the ichthyological literature that include DNA data and apply the GenSeq nomenclature to sequences referenced in those publications. We encourage authors to adopt the GenSeq nomenclature (note capital "G" and "S" when referring to the nomenclatural program) to provide a searchable tag (e.g., "genseq"; note lowercase "g" and "s" when referring to sequences) for genetic sequences from types and other vouchered specimens. Use of the new nomenclature and ranking system will improve integration of molecular phylogenetics and biological taxonomy and enhance the ability of researchers to assess the reliability of sequence data. We further encourage authors to update sequence information on databases such as GenBank whenever nomenclatural changes are made.
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    The Neotropical cichlid genus Gymnogeophagus Ribeiro, 1918 is revised. The following species are considered valid and are redescribed: G. rhabdotus (Hensel, 1870), G. gymnogenys (Hensel, 1870), G. labiatus (Hensel, 1870), G. balzanii (Perugia, 1891) and G. australis (Eigenmann, 1907). In addition, two new species are described: G. lacustris, sp. n., from the coastal region of southern Brazil and G. meridionalis, sp. n., from the lower Rio Paraná and Rio Uruguay systems. Lectotypes are designated for Geophagus bucephalus Hensel, 1870 (= G. labiatus) and Geophagus scymnophilus Hensel, 1870 (= G. labiatus) and the phylogenetic relationships among the species are analyzed. An osteological description based mainly on G. meridionalis is presented. A map of species distribution and a key to the species are provided.