Trichopodus poptae, a new anabantoid fish from Borneo (Teleostei: Osphronemidae)

Abstract

Trichopodus poptae, new species, is distinguished from all other species in the genus by an almost non-discernible colour pattern, except for a single black blotch at the base of the caudal fin; fewer lateral scales (34-38, vs. 40-65); fewer total dorsal-fin rays than T. pectoralis (14-16, vs. 17-18); more total dorsal-fin spines than T. microlepis (6-7, vs. 3-4); fewer total anal-fin rays than T. microlepis and T. pectoralis (38-41, vs. 44-50). An artificial key to the genus is provided.

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Ichthyol. Explor. Freshwaters, Vol. 25, No. 1
69
Copyright © Verlag Dr. Friedrich Pfeil
Ichthyol. Explor. Freshwaters, Vol. 25, No. 1, pp. 69-77, 3 figs., 1 tab., August 2014
© 2014 by Verlag Dr. Friedrich Pfeil, München, Germany – ISSN 0936-9902
Trichopodus poptae, a new anabantoid fish from Borneo
(Teleostei: Osphronemidae)
Bi Wei Low*, Heok Hui Tan** and Ralf Britz***
Trichopodus poptae, new species, is distinguished from all other species in the genus by an almost non-discernible
colour pattern, except for a single black blotch at the base of the caudal fin; fewer lateral scales (34-38, vs. 40-65);
fewer total dorsal-fin rays than T. pectoralis (14-16, vs. 17-18); more total dorsal-fin spines than T. microlepis (6-7,
vs. 3-4); fewer total anal-fin rays than T. microlepis and T. pectoralis (38-41, vs. 44-50). An artificial key to the
genus is provided.
* Freshwater and Invasion Biology Laboratory, Department of Biological Sciences, National University of
Singapore, Kent Ridge, Singapore 119260, Republic of Singapore. E-mail: biweilow@nus.edu.sg
** Lee Kong Chian Natural History Museum, National University of Singapore, Kent Ridge, Singapore 117546,
Republic of Singapore. E-mail: dbsthh@nus.edu.sg (corresponding author)
*** Natural History Museum, Department of Zoology, Fish Research Group, Cromwell Road, London.
E-mail: r.britz@nhm.ac.uk
Introduction
The gouramies of the Southeast Asian genus
Trichopodus La Cepède are laterally compressed,
deep-bodied fishes with a long-based anal fin
(Regan, 1910). They share with their closest rela-
tives, the species of the genus Trichogaster, the
reduction of the pelvic-fin spine and an extreme
elongation of the posterior branch of the first
pelvic-fin soft ray into a long thread-like filament
(Britz, 1995). The pelvic girdle and associated
muscles of Trichopodus and Trichogaster are also
modified to allow the elongated fin filament to be
moved in different directions including forward
(Steinbach, 1950; Liem, 1963). This filament carries
a series of taste buds innervated by the facial nerve
and is used as an organ of touch and taste (Schar-
rer et al., 1947; Steinbach, 1950; Weber, 1963).
Trichopodus gouramies are bubble-nest build-
ers and obligate air-breathers that possess an
auxiliary breathing apparatus associated with the
gills, the labyrinth organ, which allows for the
intake of atmospheric air and allows their persis-
tence in oxygen-deprived environments (Tweed-
ie 1953; Burggren, 1979). Consequently, they in-
habit a wide range of habitats, but are typically
found in heavily vegetated areas of shallow,
sluggish or standing waters, or seasonally-flood-
ed habitats (Smith, 1945; Mohsin & Ambak, 1983).
Trichopodus species have also been exploited as
ornamental fish and food fish (Ng & Tan, 1997;
Tan & Ng, 2005b).
Trichopodus gouramies were previously known
as Trichogaster Bloch & Schneider, following the
erroneous interpretation of the type species by
Myers (1923a, 1923b). This was until Derijst (1997)

70 Copyright © Verlag Dr. Friedrich Pfeil
Low, Tan & Britz: Trichopodus poptae
pointed out that the type species of Trichogaster
should have been what was known as Colisa
fasciata, and that the type species of the genus
Trichopodus La Cepède is what was known as
Trichogaster trichopterus (Britz, 2004; Tan & Kot-
telat, 2009). As it turns out, Trichopodus, previ-
ously mistaken as a synonym for Osphronemus
and Trichogaster, is valid. Following this revised
classification, there are currently four recognized
Trichopodus species, viz., T. pectoralis Regan, 1910,
T. trichopterus (Pallas, 1770), T. leerii (Bleeker, 1852)
and T. microlepis (Günther, 1861). Herein, we
describe a new species of Trichopodus from the
Barito River drainage in southern Borneo.
Material and methods
Material examined is deposited in: BMNH,
Natural History Museum, London; MZB, Re-
search and Development Centre for Biology, The
Indonesian Institute of Sciences (LIPI, formerly
the Museum Zoologicum Bogoriense), Cibinong;
and ZRC, Lee Kong Chian Natural History Mu-
seum, National University of Singapore.
Method for meristic and measurements follow
that of Ng & Kottelat (1994) and Tan & Ng (2005a).
Lateral scales were counted as the number of
scales in a longitudinal series above the lateral
line, from behind the opercular margin to the
posterior end of the hypural complex. Measure-
ments were taken with digital calipers. SL is
standard length, TL is total length and HL is head
length.
Selected specimens were cleared and double
stained following the method of Taylor & Van
Dyke (1985).
Trichopodus poptae, new species
(Figs. 1-2)
Holotype. MZB 21466, 64.3 mm SL; Indonesia:
Kalimantan Tengah: Barito River drainage: Desa
Kerendan, catch by locals; D. Siebert et al., 23 Sep
1995.
Paratypes. BMNH 2001.1.15.7668-7692, 14; MZB
21467, 4, ZRC 54264, 5, 31.8-66.1 mm SL; same
data as holotype. – BMNH 2001.1.15.7649-7667,
18, 30.2-58.1 mm SL; same data as holotype. –
BMNH 2001.1.15.1998, 1, 60.1 mm SL; Indonesia:
Barito River drainage: Kalimantan Tengah: Sun-
gai Tengkon; D. Siebert et al., 24 Sep 1995. – BMNH
2001.1.15.1999, 1, 45.4 mm SL; Indonesia: Barito
River drainage: Kalimantan Tengah: Sungai Paku-
merah, a tributary of Sungai Kerendan; D. Siebert
et al., 28 Sep 1995.
Diagnosis. Trichopodus poptae is distinguished
from all congeners in having a black blotch on
the caudal peduncle with no other distinct mark-
ings on the body (vs. two black blotches, one in
the middle of the body and the other on the
caudal peduncle in T. trichopterus; a black stripe
extending from the mouth to the caudal peduncle
in T. pectoralis and T. leerii; no distinct body mark-
ings in T. microlepis), and fewer lateral scales
(34-38, vs. 49-63 in T. pectoralis, 40-52 in T. tricho-
pterus, 43-50 in T. leerii, 57-65 in T. microlepis).
Trichopodus poptae can be further distinguished
from its congeners by the following suite of char-
acters: more dorsal-fin spines than T. microlepis
(6-7, vs. 3-4); fewer total dorsal-fin rays than
T. pectoralis (14-16, vs. 17-18); fewer total anal-fin
rays (38-41, vs. 44-50 in T. pectoralis, 44-49 in
T. microlepis and 41-46 in T. trichopterus); fewer
predorsal scales (30-34, vs. 37-44 in T. pectoralis,
36-41 in T. leerii and 45-52 in T. microlepis);
fewer transverse scales at dorsal fin origin (8-9,
1, 8-10, vs. 11-13, 1, 15-17 in T. pectoralis and
14-17, 1, 19-22 in T. microlepis); fewer longitudi-
nal scale rows on caudal peduncle (5-6, 1, 4-5,
vs. 7-9, 1, 6-9 in T. pectoralis and 8-9, 1, 6-8 in
T. microlepis); greater caudal peduncle depth
(16.0-17.2, vs. 13.3-15.8 % SL in T. pectoralis, 14.2-
15.7 % SL in T. leerii and 12.2-14.8 % SL in T. mi-
crolepis); greater body depth at anus (43.4-47.4,
vs. 36.3-42.1 % SL in T. pectoralis and 39.5-42.8 %
SL in T. leerii); smaller postdorsal length (21.9-
24.5, vs. 25.1-27.9 % SL in T. leerii and 25.8-32.9 %
SL in T. microlepis); greater orbital diameter (28.8-
33.8, vs. 18.7-27.1 % HL in T. pectoralis and 23.2-
27.0 % HL in T. microlepis); fewer pterygiophores
between first and second haemal spine (4, vs. 5
in T. trichopterus, T. pectoralis and T. microlepis).
Description. General body form as in Figures
1-2; meristics and morphometrics listed in Ta-
ble 1. Body oval, laterally compressed and deep.
Head laterally compressed with pointed snout
and slight convexity at supra-orbital area. Mouth
superior, very small and protractile. Lateral line
complete or interrupted, with 34-38 scales in a
longitudinal series above lateral line. Dorsal fin
inserted slightly behind middle of body with

Ichthyol. Explor. Freshwaters, Vol. 25, No. 1
71
Copyright © Verlag Dr. Friedrich Pfeil
Fig. 1. Trichopodus poptae, MZB 21466, holotype, 64.3 mm SL; Indonesia: Kalimantan Tengah: Barito River drain-
age (copyright BMNH).
Fig. 2. Trichopodus poptae, BMNH 2001.1.15.7669, paratype, 30.8 mm SL; Indonesia: Kalimantan Tengah: Barito
River drainage.
VI-VII, 8-9 rays; dorsal-fin base short, with 14-16
subdorsal scales. Caudal fin emarginate to forked
with 7-8 + 8 principal soft rays. Anal fin inserted
before dorsal-fin origin with XIII-XV, 25-27 rays;
anal-fin base about 3
/
4 of SL. Pelvic fins with i, 2-4
rays, with first ray filamentous and elongated.
Pectoral fin rounded.
Vertebral count: 9-10 + 21-22 = 30-31 (mode

72 Copyright © Verlag Dr. Friedrich Pfeil
= 30 or 31, n = 10). First dorsal-fin pterygiophore
inserted between neural spines 10 and 11 (7) or
11 and 12 (3). Anal-fin pterygiophore complex in
front of first haemal spine with 6 (8) or 7 (2) fin
spines; 4 (9) rarely 5 (1) anal-fin pterygiophores
between first and second haemal spine.
Colouration. See Figures 1-2 for overall appear-
ance. Dorsal surface of head and body dark brown,
fading to cream on lateral and ventral surfaces,
with 15-18 (mode 16 or 17, n = 22) faint light brown
stripes that follow longitudinal rows of scales on
body. Stripes on upper half of body are more or
less regular and horizontal, whereas on lower
half they are curved downwards posteriorly and
continued onto anal fin. All fins relatively hyaline,
more opaque at base. Caudal and posterior part
of anal fin with faint pattern of cream spots (more
visible on fish with white background). A distinct
black blotch at lower half of caudal peduncle,
roughly covering 2 1
/
2 scale rows deep and 3 scale
rows wide. Juvenile (around 30 mm SL; Fig. 2)
with similar colouration pattern as adult.
Live colouration not documented.
Distribution. Trichopodus poptae is currently
known only from the middle Barito River drain-
age in Southern Kalimantan, Borneo, in the
tributaries of Sungai Tengkon and Sungai Keren-
dan.
Table 1. Meristic and morphometric data of Trichopodus poptae (n = 10).
holotype type series
Standard length (in mm) 64.3 44.1-66.1
Meristics (mode)
Anal-fin rays XIII, 27 XIII-XV, 25-27 (XIII, 27)
Dorsal-fin rays VI, 8 VI-VII, 8-9 (VII or VII, 8)
Caudal-fin rays 8, 8 7-8, 8 (8, 8)
Pelvic-fin rays i, 3 i, 2-4 (i, 3)
Pectoral-fin rays 10 9-10 (9)
Subdorsal scales 16 14-16 (14)
Transverse scales at dorsal-fin origin 8, 1, 10 8-9, 1, 8-10 (8 or 9, 1, 8 or 9)
Lateral scales 38 34-38 (38; n = 9)
Lateral scale at dorsal-fin origin 15 14-15 (14; n = 9)
Lateral scale at anal-fin origin 33-4 (3; n = 9)
Predorsal scales 33 30-34 (33)
Postdorsal scales 24 21-25 (24)
Peduncular scales 6, 1, 5 5-6, 1, 4-5 (6, 1, 5; n = 9)
Morphometrics (mean)
In percentage of standard length
Total length 142.8 137.9-149.4 (143.7)
Body length 70.0 68.5-71.0 (70.0)
Predorsal length 59.7 56.4-62.1 (59.4)
Postdorsal length 22.7 21.9-24.5 (23.5)
Caudal peduncle depth 16.1 16.0-17.2 (16.6)
Preanal length 43.4 41.8-46.1 (44.6)
Head length 31.9 31.1-33.6 (32.4)
Body depth as anus 44.3 43.4-47.4 (45.6)
Pelvic-fin length 129.1 123.3-150.7 (137.8)
Anal-fin base length 75.9 73.2-79.4 (75.6)
Dorsal-fin base length 24.6 22.3-25.9 (24.2)
Prepelvic length 35.8 34.1-37.2 (35.5)
Head width 16.5 15.9-17.7 (16.6)
In percentage of head length
Orbit diameter 29 29-34 (30.9)
Postorbital length 44 44-46 (44.6)
Interorbital width 44 39-45 (41.3)
Snout length 31 28-32 (29.9)
Low, Tan & Britz: Trichopodus poptae

Ichthyol. Explor. Freshwaters, Vol. 25, No. 1
73
Copyright © Verlag Dr. Friedrich Pfeil
Field notes. All the specimens were obtained
from local fishers and villagers. Collection pots
with preservative were left behind by the main
expedition team in Desa Kerendan when the team
progressed further upstream. Upon returning
from survey work upstream several days later,
these collection pots were retrieved from the local
fishers and were filled with T. poptae (D. Siebert,
pers. comm.).
Trichopodus poptae had been collected only in
the area of Desa Kerendan, a small village in the
upper Sungai Lahei watershed within the Barito
River drainage. A few specimens were collected
from small streams, with gravel bottoms, which
villagers said flowed from relatively undisturbed
watersheds. Most specimens were collected from
streams and ponds in the near vicinity of Desa
Kerendan by village children.
Fishes caught with T. poptae in streams were:
Cyprinidae – Barbodes binotatus, B. everetti, Cyclo-
cheilichthys apogon, Labiobarbus fasciatus, Malayo-
chela maassi, Osteochilus sp., Rasbora dusonensis,
R. elegans; Cobitidae – Acanthopsoides sp., Pangio
shelfordii; Botiidae – Chromobotia macracantha,
Syncrossus hymenophysa; Balitoridae – Homalo-
pteroides sp.; Nemacheilidae – Nemacheilus lactoge-
neus, N. spiniferus; Vaillantellidae – Vaillantella
euepiptera.; Bagridae – Mystus castaneus, M. nigri-
ceps, M. singaringan; Siluridae – Ompok leiacan-
thus, Silurichthys gibbiceps; Clariidae – Clarias
leiacanthus; Zenarchopteridae – Hemirhamphodon
sp.; Pristolepididae – Pristolepis fasciata; Osphro-
nemidae – Betta obscura, B. pallifina; Channidae –
Channa lucius, C. marulioides, C. striata (D. Siebert,
pers. comm.; taxonomy updated).
Etymology. Named for Canna Maria Louise
Popta (1860-1929), curator of fishes for the Rijks-
museum van Natuurlijke Historie from 1891 to
1928 (Brown, 1994); in honour for her pioneer
taxonomic work on the freshwater fish fauna of
Borneo, based on the trans-Borneo expeditions
conducted between 1893 to 1900.
Artificial key to the genus Trichopodus
1. – Body with single longitudinal black stripe
extending from mouth to caudal peduncle.
....................................................................... 2
– Body with one or two black blotches along
mid-body, or no distinct markings.
....................................................................... 3
2. – Body with grey to black oblique bars; 33-39
branched anal-fin rays; 27-31 transverse
scales; dorsal-fin base length 25.9-31.5 %
SL. Distribution: Indochina.
..................................................... T. pectoralis
– Body with a mosaic of small, light silvery
spots with dark outlines; 25-30 branched
anal-fin rays; 19-25 transverse scales; dor-
sal-fin base length 19.4-23.8 % SL. Distribu-
tion: Peninsular Malaysia, Sumatra, south-
ern Borneo.
............................................................. T. leerii
3. – Body silvery or grey with no distinct mark-
ings; 3-4 dorsal-fin spines; 57-65 lateral
scales; 34-40 transverse scales; dorsal-fin
base length 14.5-17.3 % SL. Distribution:
Indochina.
.................................................... T. microlepis
– Body with one or two black blotches on
mid-body; 5-9 dorsal-fin spines; 34-52
lateral scales; 17-25 transverse scales; dor-
sal-fin base length 21.9-28.7 % SL.
....................................................................... 4
4. – Body with two black blotches, one in mid-
dle of body and other on caudal peduncle,
and faintly distinct grey oblique bars;
9-12/31-38 anal-fin rays; 40-52 lateral
scales; head length 27.3-30.6 % SL. Distribu-
tion: Indochina, Peninsular Malaysia, Su-
matra, Java, Borneo.
................................................. T. trichopterus
– Body with a single black blotch on caudal
peduncle; 13-15/25-27 anal-fin rays; 34-38
lateral scales; head length 31.3-33.6 % SL.
Distribution: southern Borneo.
.......................................................... T. poptae
Discussion
The genus Trichopodus belongs to the family Os-
phronemidae, in which it forms the sister group
to Trichogaster based on the results of morpho-
logical and molecular analyses (Britz, 1995; Rüber
et al., 2006). The two genera were classified by
Britz (2001) in the subfamily Luciocephalinae
along with the genera Parasphaerichthys, Ctenops,
Sphaerichthys, and Luciocephalus, the monophyly
of which is supported by the shared presence of
a posterior process of the basioccipital on which
Baudelot’s ligament inserts and the loss of the

74 Copyright © Verlag Dr. Friedrich Pfeil
first branchiostegal ray. The molecular analysis
of Rüber et al. (2006), however, did not recover
Trichopodus and Trichogaster as members of the
Luciocephalinae, but as the sister group to the
Macropodinae.
Notwithstanding the different hypotheses
about their phylogenetic position, Trichopodus and
Trichogaster are sister taxa and share a number of
skeletal and muscular modifications of the pelvic
girdle and fin that enable them to move the
highly elongated first pelvic soft ray in any direc-
tion including at least 180 degrees backwards to
their tail and forwards to beyond the tip of their
head (Steinbach, 1950). These filaments are used
as exploratory organs of touch and taste (Scharrer
et al., 1947; Steinbach, 1950; Weber, 1963). While
elongated first pelvic-fin rays occur in a number
of anabantoids (Britz, 1995: fig. 16), their move-
ment is very restricted, especially anteriorly, and
there are no modifications to the pelvic girdle.
Trichopodus and Trichogaster also share addi-
tional uniquely derived dorsal gill arch characters
that identify them as sister taxa among anaban-
toids that we take the opportunity to discuss here
(see Fig. 3). Primitively among anabantoids,
epibranchial 2 runs parallel to epibranchials 3 and
4 and its tip articulates with a cartilage capped
dorsally directed process in the middle of pharyn-
gobranchial 2. This condition can be found in
representatives of all anabantoid genera with the
exception of Trichopodus and Trichogaster. In these
two genera, pharyngobranchial 2 is elongated,
reaches the midline and has shifted anteriorly in
its distal portion, so that its tip no longer articu-
lates with pharyngobranchial 2 but ends freely.
This anterior shift is associated with the loss of
the cartilage capped process on pharyngo-
branchial 2, which only connects to the remaining
upper gill arch elements via its cartilage capped
process that articulates with a similar process on
pharyngobranchial 3. This highly unusual ar-
rangement of epibranchial 2 was illustrated by
Rosen & Patterson (1990) for Trichogaster lalia, but
not described or discussed. The only other ana-
bantoid, in which we also found the dorsally
directed process on pharyngobranchial 2 missing
is Luciocephalus, but here epibranchial 2 is still
aligned in parallel with the other epibranchials
and not shifted anteriorly. We interpret the lack
of the pharyngobranchial process as indepen-
dently derived in Luciocephalus and probably
related to its highly predatory behaviour and its
greatly flattened dorsal gill arches.
The dorsal gill arch skeleton also shows in-
teresting differences between Trichopodus and
Trichogaster. In Trichopodus the distal end of epi-
branchial 2 is flattened and greatly expanded,
while that of Trichogaster is round in diameter.
Trichogaster on the other hand has a band of tiny
serially arranged toothplates in front of pharyn-
gobranchial 2, not illustrated and probably over-
looked by Rosen & Patterson (1990: fig. 29).
Trichopodus and Trichogaster can also be easily
distinguished from each other, by the short dor-
sal fin with a reduced number of fin spines and
rays in the former and the long dorsal fin with
numerous spines and rays in the latter. A series
of ray-less pterygiophores in front of the dorsal
fin in Trichopodus are remnants of the formerly
longer dorsal fin. Further differences between the
two genera are the complete lack of pelvic-fin
rays other than the filamentous first ray on
Trichogaster, while Trichopodus still has an addi-
tional 3-4 small to tiny rays depending on the
species.
The number of pterygiophores between hae-
mal spines 1 and 2 differs among Trichopodus
species and separates them into two groups:
T. pectoralis, T. trichopterus, and T. microlepis usu-
ally have 5 pterygiophores, while T. leeri and
T. poptae usually show 4. This lower number in
T. leeri and T. poptae correlates with fewer total
anal-fin rays in these two species (37-44, vs. 41-
50 in the other three species). Trichopodus poptae
is most easily distinguished from its congeners
by its colour pattern, consisting of a black blotch
on the caudal peduncle and no other markings
on the body except some pale longitudinal lines.
The cream spots on its fins are somewhat remi-
niscent of those of T. leeri, the so-called pearl
gourami. In the molecular phylogeny of Rüber et
Fig. 3. Dorsal gill arches 2 to 4 in dorsal (left side) and
frontal view (right side) of: a, Sandelia bainsii (Anabanti-
dae), BMNH 2014.4.2.1, 98 mm SL; b, Belontia hasselti
(Osphronemidae, Belontiinae), BMNH 2014.4.2.2, 74 mm
SL; c, Trichopodus microlepis (Osphronemidae, Lucio-
cephalinae), BMNH 2014.4.2.3, 65 mm SL, epibranchial
2 removed in frontal view; d, Trichogaster lalia (Os-
phronemidae, Luciocephalinae), BMNH 2014.4.2.4,
38 mm SL, epibranchial 2 removed in frontal view, band
of tiny ossicles in front of pharyngobranchial 2 marked
with arrowheads. Note lack of articular process in mid-
dle of pharyngobranchial 2 and anterior shift of tip of
epibranchial 2 in c and d. Eb 2, 3, 4, epibranchial 2, 3, 4;
Pb 2, 3, pharyngobranchial 2, 3; UP 4, upper pharyn-
geal toothplate 4. Scale bars 1 mm.
/
Low, Tan & Britz: Trichopodus poptae

Ichthyol. Explor. Freshwaters, Vol. 25, No. 1
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Copyright © Verlag Dr. Friedrich Pfeil
Eb4
Pb3
Pb2
Eb2
UP4
Eb3
Eb4
Pb3
Pb2
Eb2
UP4
Eb3
Eb4 Pb3 Pb2
UP4
Eb3
Eb4 Pb3
Pb2
Eb2
UP4
Eb3
a
d
Eb4
Pb3
Pb2
UP4
Eb3
Eb4
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76 Copyright © Verlag Dr. Friedrich Pfeil
al. (2006) T. leeri was recovered as the sister group
to the other three Trichopodus species known at
that time. It will be interesting to see where
T. poptae fits phylogenetically among the different
Trichopodus species, but its similarities to T. leeri,
which are most likely plesiomorphies, point to a
position at the base of Trichopodus, either as the
sister group of all other species or as the sister
group to T. leeri.
Comparison material. Trichopodus leerii: ZRC 38614,
8, 47.0-61.9 mm SL; Indonesia: Sumatra: Jambi: Danau
Rasau. – ZRC 43248, 1, 66.8 mm SL; Indonesia: South
Sumatra: Sungei Sentang.
Trichopodus microlepis: ZRC 53428, 6, 74.4-98.7 mm
SL; Cambodia: Phnom Penh: Pochengtong Market. –
BMNH uncat, E3 223, 1, 71.1 mm SL; E3 224, 1, 69.0 mm
SL; E3 225, 1, 65.8 mm SL; not labelled, 1, 66.3 mm SL;
Cambodia: local markets.
Trichopodus pectoralis: ZRC 1621, 6, 93.6-168.0 mm
SL; Malaysia: Penang: Bayan Lepas Fishponds. – BMNH
uncat, E3 233, 1, 106.9 mm SL; E3 234, 1, 105.2 mm SL;
E3 235, 1, 100.4 mm SL; E3 236, 1, 101.4 mm SL; Cam-
bodia: local markets.
Trichopodus trichopterus: ZRC 43871, 3, 72.8-75.2 mm
SL; Indonesia: Java: Banjar market, about 500 m from
train station. – ZRC 53311, 7, 42.2-73.5 mm SL; Singa-
pore: Sungei Buloh Wetland Reserve, Outdoor Class-
room pond.
The following references were used to supplement
comparison materials in the diagnosis of T. poptae
(relevant species in parentheses): Regan, 1910
(T. pectoralis, T. trichopterus, T. leerii and T. micro-
lepis); Smith, 1945 (T. pectoralis, T. trichopterus,
T. leerii and T. microlepis); Mohsin & Ambak, 1983
(T. pectoralis and T. trichopterus); Rainboth, 1996
(T. pectoralis, T. trichopterus, T. leerii and T. micro-
lepis); Kottelat, 2001 (T. microlepis).
Acknowledgements
Many thanks to the following: Darrell Siebert, Oliver
Crimmen (BMNH); Renny Hadiaty (MZB), Kelvin Lim
(ZRC), for access to material. We are grateful to Phil
Crabb, Photo Unit BMNH, for photographing the
holotype of T. poptae and Jonathan Jackson (BMNH) for
locating the digital photograph. Funding from the Raf-
fles Museum of Biodiversity Research and research
grant R-154-000-318-112 from the National University
of Singapore is kindly acknowledged.
Literature cited
Bleeker, P. 1852. Diagnostische beschrijvingen van
nieuwe of weinig bekende vischsoorten van Suma-
tra. Natuurkundig Tijdschrift voor Nederlandsch
Indië, 3: 569-608.
Britz, R. 1995. Zur phylogenetischen Systematik der
Anabantoidei (Teleostei, Percomorpha) unter be-
sonderer Berücksichtigung der Stellung des Genus
Luciocephalus. Morphologische und ethologische
Untersuchungen. PhD dissertation, Eberhard-Karls-
Universität Tübingen, 125 pp.
— 2001. The genus Betta – monophyly and intrarela-
tionships, with remarks on the subfamilies Ma-
cropodinae and Luciocephalinae. Ichthyological
Exploration of Freshwaters, 12: 305-318.
— 2004. Why Colisa has become Trichogaster and
Trichogaster is now Trichopodus. Labyrinth, 136: 6-7.
Brown, P. S. 1994. Early women ichthyologists. Envi-
ronmental Biology of Fishes, 41: 9-30.
Burggren, W. W. 1979. Bimodal gas exchange during
variation in environmental oxygen and carbon di-
oxide in the air breathing fish Trichogaster tricho-
pterus. Journal of Experimental Biology, 82: 197-213.
Derijst, E. 1997. Nota over de geldigheid van de genus-
namen: Trichogaster Bloch & Schneider, 1801; Tri-
chopodus Lacepède, 1801; Polyacanthus Cuvier, 1829
en Colisa Cuvier, 1831 (Perciformes: Belontiidae);
met commentaar over de publicatiedata van de
werken van Bloch & Schneider en van Lacepède,
beide verschenen in 1801 en over het auteurschap
van de genusnaam Colisa. Aquariumwereld, 50 (9):
217-226.
Günther, A. 1861. Catalogue of the acanthopterygian
fishes in the collection of the British Museum, vol-
ume 3. London: Trustees of the British Museum,
xxxv + 586 pp.
Kottelat, M. 2001. Fishes of Laos. WHT Publications,
Colombo, 198 pp.
Liem, K. F. 1963. The comparative osteology and phy-
logeny of the Anabantoidei. Illinois Biological
Monographs, 30: 1-149.
Mohsin, A. K. M. & M. A. Ambak. 1983. Freshwater
fishes of Peninsular Malaysia. Penerbit Universiti
Pertanian Malaysia, Serdang, 284 pp.
Myers, G. S. 1923a. Notes on the nomenclature of certain
anabantids and a new generic name proposed.
Copeia, 118: 62-63.
— 1923b. Further notes on anabantoids. Copeia, 124:
111-112.
Ng, P. K. L. & M. Kottelat. 1994. Revision of the Betta
waseri group (Teleostei: Belontiidae). Raffles Bul-
letin of Zoology, 42: 593-611.
Ng, P. K. L. & H. H. Tan. 1997. Freshwater fishes of
Southeast Asia: potential for the aquarium trade
and conservation issues. Aquarium Science Con-
servation, 1: 79-90.
Low, Tan & Britz: Trichopodus poptae

Ichthyol. Explor. Freshwaters, Vol. 25, No. 1
77
Copyright © Verlag Dr. Friedrich Pfeil
Pallas, P. S. 1770. Spicilegia Zoologica quibus novae
imprimis et obscurae animalium species iconibus,
descriptionibus atque commentariis illustrantur.
Lange, Berolini [Berlin], 1 (fasc. 8): 1-56, pls. 1-5.
[Also 1 (fasc. 7 [1769]): 1-42, pls. 1-6.].
Rainboth, W. J. 1996. FAO species identification field
guide for fishery purposes. Fishes of the Cambo-
dian Mekong. FAO, Rome, 265 pp., 27 pls.
Regan, C. T. 1910. The Asiatic fishes of the family Ana-
bantidae. Proceedings of the Zoological Society of
London, 1909: 767-787, pls. 77-79.
Rosen, D. E. & C. Patterson. 1990. On Müller’s and
Cuvier’s concepts of pharyngognath and labyrinth
fishes and the classification of percomorph fishes,
with an atlas of percomorph dorsal gill arches.
American Museum Novitates, 2983: 1-57.
Rüber, L., R. Britz & R. Zardoya. 2006. Molecular phy-
logenetics of labyrinth fishes, evolution of their
breeding behaviour and remarks on their historical
biogeography (Perciformes; Anabantoidei). System-
atic Biology, 55: 374-397.
Scharrer, E., S. W. Smith & S. L. Palay. 1947. Chemical
sense and taste in the fishes, Prionotus and Tricho-
gaster. Journal of Comparative Neurology, 86:
183-198.
Smith, H. M. 1945. The fresh-water fishes of Siam, or
Thailand. Bulletin of the United States National
Museum, 188: 1-622, 9 pls.
Steinbach, G. 1950. Funktionelle und vergleichend
anatomische Untersuchungen über die Bauchflos-
senvon Trichogaster trichopterus Pallas. Deutsche
Zoologische Zeitschrift, 1: 24-60.
Tan, H. H. & P. K. L. Ng. 2005a. The fighting fishes
(Teleostei: Osphronemidae: genus Betta) of Singa-
pore, Malaysia and Brunei. Raffles Bulletin of Zool-
ogy, Supplement 13: 43-99.
Tan, H. H. & P. K. L. Ng. 2005b. The labyrinth fishes
(Teleostei: Anabantoidei, Channoidei) of Sumatra,
Indonesia. Raffles Bulletin of Zoology, Supplement
13: 115-138.
Tan, H. H. & M. Kottelat. 2009. The fishes of the Batang
Hari drainage, Sumatra, with description of six new
species. Ichthyological Exploration of Freshwaters,
20: 13-69.
Taylor, W.R. & G. C. Van Dyke. 1985. Revised proce-
dures for staining and clearing small fishes and
other vertebrates for bone and cartilage study.
Cybium, 9: 107-119.
Tweedie, M. W. F. 1953. Malayan aquarium fishes.
Malayan Nature Journal, 8: 47-51, pls. 9-10.
Weber, H. 1963. Die Sinnesfunktionen der freien Bauch-
flossenstrahlen der Labyrinthfische (Anabantidae)
und ihr Zusammenwirken mit den Augen. Zeit-
schrift für Vergleichende Physiologie, 47: 77-110.
Received 21 November 2013
Revised 1 April 2014
Accepted 2 April 2014

78 Copyright © Verlag Dr. Friedrich Pfeil
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INSTRUCTIONS TO CONTRIBUTORS
Ichthyological Exploration of Freshwaters
An international journal for field-orientated ichthyology

Articles appearing in this journal are indexed in:
AQUATIC SCIENCES and FISHERIES ABSTRACTS
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C O N T E N T S
Freyhof, Jörg and Müfit Özulug: Acanthobrama thisbeae, a new species of bream from south-
ern Anatolia, Turkey (Teleostei: Cyprinidae) ......................................................................... 1
Freyhof, Jörg, Hamid Reza Esmaeili, Golnaz Sayyadzadeh and Matthias Geiger: Review of
the crested loaches of the genus Paracobitis from Iran and Iraq with the description of
four new species (Teleostei: Nemacheilidae) ......................................................................... 11
Esguícero, André L. H. and Ricardo M. C. Castro: Knodus figueiredoi, a new characid from
the Rio das Garças, upper Rio Araguaia basin, Brazil, with comments on the taxo-
nomic limits of the genera Knodus and Bryconamericus (Teleostei: Characidae) ............... 39
Lamboj, Anton: Two new species of Parananochromis from Cameroon, Central Africa (Tele-
ostei: Cichlidae) ........................................................................................................................... 49
Kim, Daemin, Hyung-Bae Jeon and Ho Young Suk: Tanakia latimarginata, a new species of
bitterling from the Nakdong River, South Korea (Teleostei: Cyprinidae) ........................ 59
Low, Bi Wei, Heok Hui Tan and Ralf Britz: Trichopodus poptae, a new anabantoid fish from
Borneo (Teleostei: Osphronemidae) ........................................................................................ 69
Costa, Wilson J. E. M.: Six new species of seasonal killifishes of the genus Cynolebias from
the São Francisco river basin, Brazilian Caatinga, with notes on C. porosus (Cyprinodonti -
formes: Rivulidae) ...................................................................................................................... 79
Ichthyological Exploration of Freshwaters
An international journal for field-orientated ichthyology
Volume 25 • Number 1 • August 2014
Cover photograph
Parananochromis orsorum, female (photograph by A. Lamboj)
Anton Lamboj
(this volume pp. 49-57)