The labyrinth fishes (Teleostei : Anabantoidei, Channoidei) of Sumatra, Indonesia

Abstract

Three families (Anabantidae, Helostomatidae, Osphronemidae) and 32 species of anabantoids, and nine species of channids (Channidae) are recorded from Sumatra and adjacent islands. Three new species are described. Betta cracens, new species, differs from other members of the B. pugnax group in having the most slender body (body depth 21.2-24.2% SL), more anal fin rays (27-29) and lateral scales (32-33). Betta fasca is redescribed and a lectotype is designated. An allied species, Betta raja is described, differing from B. fusca in having a longer head (HL 33.9-37.4% SL, vs. 33.2-33.8), more lateral scales (30-32, vs. 29) and a relatively longer pelvic fin. Betta rubra is redescribed from type and recent material. Luciocephalus aura, new species, is described from Jambi province; where it is syntopic with L. pulcher, but differs in having different body and fin patterns, most prominently the presence of numerous iridescent green spots on central stripe (vs. absence of green spots). The taxonomy of Luciocephalus pulcher is also clarified and a neotype is designated. The taxonomy of Channa marulioides and C. melanoptera is discussed.

Full text

115
THE RAFFLES BULLETIN OF ZOOLOGY 2005
THE RAFFLES BULLETIN OF ZOOLOGY 2005 Supplement No. 13: 115-138
THE LABYRINTH FISHES (TELEOSTEI: ANABANTOIDEI, CHANNOIDEI)
OF SUMATRA, INDONESIA
Tan Heok Hui
Department of Biological Sciences, National University of Singapore,
Kent Ridge, Singapore 119260, Republic of Singapore
Email: dbsthh@nus.edu.sg
Peter K. L. Ng
Department of Biological Sciences, National University of Singapore,
Kent Ridge, Singapore 119260, Republic of Singapore
Email: dbsngkl@nus.edu.sg
ABSTRACT. – Three families (Anabantidae, Helostomatidae, Osphronemidae) and 32 species of anabantoids,
and nine species of channids (Channidae) are recorded from Sumatra and adjacent islands. Three new species
are described. Betta cracens, new species, differs from other members of the B. pugnax group in having the
most slender body (body depth 21.2-24.2% SL), more anal fin rays (27-29) and lateral scales (32-33). Betta
fusca is redescribed and a lectotype is designated. An allied species, Betta raja is described, differing from
B. fusca in having a longer head (HL 33.9-37.4% SL, vs. 33.2-33.8), more lateral scales (30-32, vs. 29) and
a relatively longer pelvic fin. Betta rubra is redescribed from type and recent material. Luciocephalus aura,
new species, is described from Jambi province; where it is syntopic with L. pulcher, but differs in having
different body and fin patterns, most prominently the presence of numerous iridescent green spots on central
stripe (vs. absence of green spots). The taxonomy of Luciocephalus pulcher is also clarified and a neotype
is designated. The taxonomy of Channa marulioides and C. melanoptera is discussed.
KEY WORDS. – Taxonomy, Anabantoidei, Channoidei, new species, Sumatra.
INTRODUCTION
The ichthyofauna of Sumatra is relatively poorly known when
compared to Borneo or Malaysia. The definition of Sumatra
in this context includes all the surrounding islands (Riau
Archipelago, the islands Nias, Banka and Biliton), but
excludes the political boundary that includes the Anambas
and Natunas islands in the Riau Archipelago (sensu Whitten
et al., 1987). For the inland fishes of the Anambas and Natuna
islands, please refer to Tan & Lim (2004). With regards to
the air breathing labyrinth fish fauna (Anabantoidei and
Channoidei), recent collections in central Sumatra, Banka and
Pulau Bintan (Riau Archipelago) from 1993 to 1997 have
resulted in the discovery of eight new species of Betta and
a new species of Parosphromenus (Osphronemidae) from this
area, viz., Betta burdigala Kottelat & Ng, 1994; B.
chloropharynx Kottelat & Ng, 1994; B. miniopinna Tan &
Tan, 1994; B. schalleri Kottelat & Ng, 1994; B. spilotogena
Ng & Kottelat, 1994; B. simorum Tan & Ng, 1996; B. renata
Tan, 1998; B. falx Tan & Kottelat, 1998b; and
Parosphromenus bintan Kottelat & Ng, 1998.
The intra-relationships of the suborder Anabantoidei were
recently revised (Britz, 1994a, 1995, 2001; Britz et al., 1995)
and several families have been shown to be monophyletic,
and Belontiidae and Luciocephalidae are synonymised with
Osphronemidae (Britz, 1994a; Kottelat & Whitten, 1996).
The suborder Anabantoidei is represented in mainland
Sumatra by three families (Anabantidae, Helostomatidae,
Osphronemidae) and 32 species; and the suborder Channoidei
is represented by one family (Channidae) and nine species
(Table 1). In addition, two species have been introduced -
Betta splendens and Trichogaster pectoralis. Herein, two new
species of Betta from the B. pugnax group are described - B.
cracens and B. raja. The lesser known B. fusca and B. rubra
are redescribed and a lectotype is designated for B. fusca.
Luciocephalus pulcher is redescribed, a neotype is designated
and L. aura, new species, is described from the Jambi
province.
MATERIAL AND METHODS
Material examined are deposited in Natural History Museum,
London (ex British Museum of Natural History, BMNH); the

116
Tan & Ng: Anabantoidei and Channoidei of Sumatra
collection of Maurice Kottelat, Cornol (CMK); Museo Civico
di Storia Naturale “Giacoma Doria”, Genova (MSNG);
Research and Development Centre for Biology (ex Museum
Zoologicum Bogoriense), The Indonesian Institute of
Sciences (MZB); Naturhistorisches Museum Basel (NMBA);
Naturhistorisches Museum Bern (NMBE); Nationaal
Natuurhistorisch Museum, Leiden (ex Rijksmuseum van
Natuurlijke Historie, RMNH); Instituut voor Systematiek &
Populatiebiologie, Universiteit van Amsterdam (ZMA); and
the Zoological Reference Collection of the Raffles Museum
of Biodiversity Research, National University of Singapore
(ZRC). Material examined are listed by Province (from north
to south) and followed by the collection abbreviation and
catalogue number with the number of specimens in brackets.
Full locality data is quoted only for new species and
redescriptions of poorly known taxa.
All measurements are taken from the left side of body
(whenever possible) with a pair of dial calipers (0.05 mm).
Taxonomy of the various taxa follow that of Tan & Ng (2005)
(present volume), Kottelat et al., 1993, and Lee & Ng (1994).
The synonymy is listed only for literature citation of species
from Sumatra. Species sold for food are denoted by *, species
in the aquarium trade are denoted by †. A list of the species
of anabantoids and channoids of Sumatra and their localities
by province and island is provided in Table 1, based on
literature and collections. Material examined is listed by
locality, catalogue number and the number of specimens in
brackets; more details are provided for new taxa and
redescriptions. Abbreviations used are SL: standard length,
TL: total length, HL: head length.
TAXONOMY
SUB-ORDER ANABANTOIDEI
FAMILY ANABANTIDAE
Anabas testudineus (Bloch) *
Anabas macrocephalus - Günther, 1861: 376; Volz, 1905: 125; 1907:
125.
Anabas oligolepis - Weber & de Beaufort, 1912: 539.
Anabas scandens - Volz, 1905: 124 ; 1907: 124; Weber & de
Beaufort, 1915: 270.
Anabas testudineus - Bleeker, 1879: 7; Weber & de Beaufort, 1912:
539; 1922: 334; Fowler & Bean, 1927: 4; de Beaufort, 1939:
193; SH Tan & HH Tan, 1994: 357.
Material examined. – Sumatra Utara - ZRC 42472 (4). Riau -
ZRC 39045 (6). Jambi - ZRC 41695 (2).
FAMILY HELOSTOMATIDAE
Helostoma temminckii Cuvier *
Helostoma temmincki - Bleeker, 1879: 15; Volz, 1905: 125; 1907:
125; Weber & de Beaufort, 1912: 540; 1922: 340; Hardenberg,
1933: 308.
Helostoma temminckii - Günther, 1861: 377; Regan, 1910: 773;
Fowler & Bean, 1927: 4.
Material examined. – Jambi - ZRC 42342 (1); ZRC 41736 (5).
FAMILY OSPHRONEMIDAE
Belontia hasselti (Cuvier & Valenciennes) *
Polyacanthus einthoveni - Volz, 1905: 125 ; 1907: 125.
Polyacanthus einthoveni - Günther, 1861: 378.
Polyacanthus hasselti - Günther, 1861: 378; Volz, 1905; 125 ; 1907:
125; Weber & de Beaufort, 1922: 338; Fowler & Bean, 1927:
4; de Beaufort, 1939: 193.
Polyacanthus hasseltii - Bleeker, 1879: 12; Regan, 1910: 772; Weber
& de Beaufort, 1912: 540.
Material examined. – Sumatra Utara - ZRC 42479 (1). Jambi -
ZRC 41737 (1); ZRC 46212 (16). Sumatra Selatan - ZRC 43247
(2). Banka - ZRC 31015-16 (2). Biliton - ZRC 46301 (1).
Betta Bleeker
For the genus Betta, the species are listed by species groups
according to Tan (1998), Tan & Kottelat (1998a) and Tan &
Ng (2005)(present volume).
Betta bellica species group
Betta bellica Sauvage
Betta bellica - Kottelat et al., 1993: 161; Tan & Ng, 1996: 144, fig.
4B; Tan & Ng, 2005: 58 (present volume).
Betta fasciata - Regan, 1910: 782, pl. 77, Fig. 4.
Material examined. – Sumatra Utara - BMNH 1889.12.26:30
(lectotype of B. fasciata); ZRC 42468 (10); ZRC 42478 (4). Riau
Archipelago - ZRC 44057 (14).
Remarks. – Betta bellica is found only in Sumatra Utara and
Riau Archipelago, whereas B. simorum is found in Riau,
Jambi and Sumatra Selatan. Betta bellica differs from B.
simorum in the following characters: lower anal fin ray count
(mode 31, vs. 32); lower dorsal fin ray count (mode 12, vs.
11); lower lateral scale count (mode 33, vs. 34); lower
postdorsal scale count (mode 8, vs. 9); shorter pelvic fin (23.6-
38.8% SL, vs. 31.3-48.3; reaching end of 8th anal ray, vs.
14th); shorter pelvic fin flange (ending halfway pelvic
filament, vs. two-thirds); non-overlap of adpressed pelvic fin
branched rays with anal fin origin (vs. overlap); and a less
sloping lateral head profile (vs. more sloping with a slight
concavity (see Tan & Ng, 1996).
Betta simorum Tan & Ng †
Betta bellica (non Sauvage, 1884) - Kottelat et al., 1993: 161 (part),
pl. 75.
Betta simorum Tan & Ng, 1996: 151, figs. 3C, 4C.

117
THE RAFFLES BULLETIN OF ZOOLOGY 2005
Table 1. Distribution of Anabantoids (Anabantidae, Helostomatidae, Osphronemidae) and Channoids (Channidae) of Sumatra (based on literature and collections)
Aceh Sumatra Utara Nias Sumatra Barat Riau Riau Archipel. Jambi Sumatra Selatan Banka Biliton Bengkulu Lampung
Anabantidae
Anabas testudineus +++ ++++ + ++ +
Helostomatidae
Helostoma temminckii ++ ++ + + ++
Osphronemidae
Belontia hasselti ++ + + + ++
Betta bellica ++
B. burdigala +
B. chloropharynx +
B. coccina ++ +
B. cracens +
B. edithae +++
B. falx ++ + +
B. fusca ++
B. hipposideros ++
B. imbellis +
B. miniopinna +
B. pugnax ++++
B. raja +++
B. renata ++ +
B. rubra ++
B. schalleri +
B. simorum ++
B. spilotogena +
Luciocephalus aura +
L. pulcher +++++++
Osphronemus goramy +++++ +
Parosphromenus bintan ++
P. deissneri ++
P. sumatranus +
Sphaerichthys osphromenoides ++ + + ++
S. cf. selatenensis +
Trichogaster leerii ++ + + + +
T. trichopterus ++++++++++
Trichopsis vittata ++++
Channidae
Channa bankanensis ++ + + ++
C. cyanospilos +++ +
C. gachua +++++++++
C. lucius ++ ++++ + ++
C. marulioides +++++++
C. melasoma ++ ++ +++
C. micropeltes ++ + + + ++
C. pleurophthalma ++
C. striata +++++++++
Total no. of species 12 17 3 7 20 19* 26* 20 18* 16 6 5
*reflects extensive collection done

118
Tan & Ng: Anabantoidei and Channoidei of Sumatra
Material examined. – Holotype - Jambi - MZB (holotype), 63.8
mm SL.
Paratypes - Jambi - ZRC 38549 (4 paratypes); CMK 11182 (5
paratypes); ZRC 39118 (19 paratypes).
Others - Riau - ZRC 39249 (2), 50.3-52.1 mm SL. Sumatra Selatan
- ZMA 121.825 (3). Kalimantan Barat - ZRC 46185 (8).
Remarks. – Betta simorum is restricted to the southern half
of Sumatra and was recently recorded from the Kapuas basin
in Kalimantan Barat (pers. observ.). It is closely allied to B.
bellica (see Tan & Ng, 1996).
Betta coccina species group
Betta burdigala Kottelat & Ng
Betta burdigala Kottelat & Ng, 1994: 70, fig. 4.
Material examined. – Holotype - Banka - ZRC 35162 (holotype),
23.8 mm SL.
Paratypes - ZRC 35163-35164 (2 paratypes), CMK 9597 (2
paratypes).
Remarks. – Betta burdigala is restricted to the island of
Banka. It differs from B. coccina in the following characters:
lack of lateral iridescent blotch (vs. presence); white pelvic
fin tip (vs. black); more dorsal fin rays (14-15, vs. 9-12); more
subdorsal scales (11-111/2, vs. 7-8); fewer anal fin rays (24-
26, vs. 27-29); fewer predorsal scales (15-16, vs. 18-21);
smaller total length (131.0-132.3% SL, vs. 132.7-138.5);
greater postorbital length (16.7-17.4% SL, vs. 14.3-15.5);
greater dorsal fin base length (26.8-28.6% SL, vs. 15.2-20.4).
Betta burdigala differs from B. miniopinna in the following
characters: red body colour (vs. black); presence of iridescent
green spots on median fins (vs. tabsence); more dorsal fin
rays (14-15, vs. 10-11); more subdorsal scales (11-111/2, vs.
8); fewer anal fin rays (24-26, vs. 27-28); fewer predorsal
scales (15-16, vs. 18); greater dorsal fin base length (26.8-
28.6% SL, vs. 16.0-24.6); greater postdorsal length (16.7-
17.4% SL, vs. 12.3-13.3); smaller predorsal length (52.8-
54.9% SL, vs. 56.2-65.4); and a greater preanal length (42.5-
45.5% SL, vs. 37.9-42.6).
Betta coccina Vierke †
Betta coccina Vierke, 1979: 288 (type locality: Jambi); Kottelat et
al., 1993: 161; Tan & Ng, 2005: 80 (present volume).
Material examined. – Riau Archipelago - CMK 7290 (10); ZRC
21051-52 (2); ZRC 44056 (36). Jambi - ZRC 38257 (2); ZRC 38611
(4); ZRC 39672 (7).
Remarks. – Betta coccina belongs to the B. coccina group
and is also found in Peninsular Malaysia. It is a stenotopic
blackwater species. It differs from B. burdigala in the
following characters: presence of lateral iridescent blotch in
males (vs. absence); black pelvic fin tip (vs. white); fewer
dorsal fin rays (9-12, vs. 14-15); fewer subdorsal scales (7-
8, vs. 11-111/2); more anal fin rays (27-29, vs. 24-26); more
predorsal scales (18-21, vs. 15-16); greater total length
(132.7-138.5% SL, vs. 131.0-132.3); smaller postorbital
length (14.3-15.5% SL, vs. 16.7-17.4); smaller dorsal fin base
length (15.2-20.4% SL, vs. 26.8-28.6). Betta coccina differs
from B. miniopinna in the following characters: presence of
lateral iridescent blotch in males (vs. absence); red body
colour (vs. black); black pelvic fin tip (vs. white); presence
of iridescent green spots on median fins (vs. absence); greater
postorbital length (14.3-15.5% SL, vs. 12.3-13.3); smaller
pelvic fin length (7.8-8.8% SL, vs. 8.7-12.0).
Betta miniopinna Tan & Tan
Betta miniopinna HH Tan & SH Tan, 1994: 42, fig. 1; SH Tan &
HH Tan, 1994: 357.
Material examined. – Holotype - Riau Archipelago - ZRC 32504
(holotype), 24.1 mm SL.
Paratypes - ZRC 32505 (1 paratype); MZB (1 paratype); ZRC 32503
(1 paratype).
Others - ZRC 37510 (10).
Remarks. – Betta miniopinna is restricted to Pulau Bintan of
the Riau Archipelago. Betta miniopinna differs from B.
burdigala in the following characters: black body colour (vs.
red); absence of iridescent green spots on median fins (vs.
presence); fewer dorsal fin rays (10-11, vs. 14-15); fewer
subdorsal scales (8, vs. 11-111/2); more anal fin rays (27-28,
vs. 24-26); more predorsal scales (18, vs. 15-16); smaller
dorsal fin base length (16.0-24.6% SL, vs. 26.8-28.6); smaller
postdorsal length (12.3-13.3% SL, vs. 16.7-17.4); greater
predorsal length (56.2-65.4% SL, vs. 52.8-54.9); smaller
preanal length (37.9-42.6% SL, vs. 42.5-45.5). Betta
miniopinna differs from B. coccina in the following
characters: absence of lateral iridescent blotch (vs. presence);
black body colour (vs. red); white pelvic fin tip (vs. black);
absence of iridescent green spots on median fins (vs.
presence); and a smaller postorbital length (12.3-13.3% SL,
14.3-15.5% SL); greater pelvic fin length (8.7-12.0% SL, vs.
7.8-8.8).
Betta edithae species group
Betta edithae Vierke
Betta edithae - Witte & Schmidt, 1992: 324, 326; Kottelat et al.,
1993: 162; SH Tan & HH Tan, 1994: 357.
Material examined. – Riau Archipelago - ZRC 37551 (4); ZRC
34721-42 (22); ZRC 34877 (7); ZRC 42512 (13); ZRC 42513 (26);
ZRC 42514 (2); ZRC 42515 (13). Sumatra Selatan - ZRC 46193
(24). Banka - ZRC 31031-44 (14); ZRC 30899-65 (17); ZRC 35159-
61 (3). Biliton - ZRC 46182 (3).
Remarks. – Betta edithae belongs to its own species group.
It has a widespread distribution and is found in Kalimantan

119
THE RAFFLES BULLETIN OF ZOOLOGY 2005
Timur, Kalimantan Selatan, Kalimantan Tengah, Kalimantan
Barat, Sumatra Selatan, Banka, Biliton and Pulau Bintan (of
the Riau Archipelago) (Witte & Schmidt, 1992; Tan & Tan,
1994; Kubota et al., 1996; M. Kottelat, pers. comm.). The
specimens from Pulau Bintan are slightly different from the
topotypes from Banjarmasin in the following characters: chin-
bar interrupted or joined by thin black mark in the middle,
end of chin-bar towards throat rounded (vs. uninterrupted
chin-bar without rounded ends); greater preanal length (48.2-
51.8% SL, vs. 45.6-47.2); and a smaller adult size (40 mm
SL, vs. 55 mm SL of holotype). From preliminary
examinations, the series from Palembang (Sumatra Selatan)
is similar to the Bankan population.
Betta foerschi species group
Betta rubra Perugia
(Fig. 1)
Betta rubra Perugia, 1893: 242; Volz, 1905: 127 (part); 1907: 127
(part); Regan, 1910: 781, pl. 77, Fig. 1; Kottelat et al., 1993:
163; Tan & Ng, 2005: 58 (present volume).
Material examined. – Lectotype - Sumatra Barat - MSNG 13019a
(lectotype), 28.6 mm SL.
Paralectotypes - MSNG 13019 (4 paralectotypes), 26.9-31.4 mm
SL; Siboga; E. Modigliani, 1886. Sumatra Utara - BMNH
1893.5.29:1 (1 paralectotype), 33.3 mm SL; Lake Toba; E.
Modigliani.
Others - Aceh - MZB 4784 (3), 23.4-35.0 mm SL, ZRC 42497 (2),
33.4-35.4 mm SL; Aceh Barat, Alur Sungai Iamueselatan; H. B.
Munaf & M. Toha, 13 Dec.1982.
Diagnosis. – Betta rubra can be differentiated from the other
members of B. foerschi group in having the following unique
set of characters: presence of 5-7 irregular dark vertical bars
on body; a pair of vertical parallel reddish bars on opercle;
presence of broad chin-bar, which merges with anterior part
of second postorbital stripe, forming a triangular dark mark
below eye; more anal fin rays than B. foerschi (27-29, vs.
26); fewer lateral scales than B. foerschi (281/2-30, vs. 31);
greater total length than B. foerschi (131.6-142.3% SL, vs.
127.2-133.5); smaller caudal peduncle depth than B. foerschi
and B. strohi (13.1-16.3% SL, vs. 17.5-20.1; 45.1-50.6% HL,
vs. 52-70); slightly smaller body depth at dorsal-fin origin
than B. foerschi and B. strohi (23.2-27.1% SL, vs. 26.6-28.9;
74.2-86.5% HL, vs. 97-102); greater pelvic fin length than
B. foerschi (28.1-38.4% SL, vs. 21.4-28.5); smaller dorsal-
fin base length than B. strohi (9.6-12.5% SL, vs. 12.6-15.8);
and a smaller interorbital width than B. foerschi and B. strohi
(6.3-9.9% SL, vs. 9.6-13.4; 22.7-29.5% HL, vs. 30-44).
Description. – General appearance as illustrated in Figs. 1a,
b; meristic and morphometric data of B. rubra is listed in
Table 2. Body relatively slender (body depth 23.2-27.1% SL),
head relatively short (head length 27.3-33.9% SL). Dorsal
and anal fins pointed, caudal fin lanceolate; dorsal fin placed
relatively far back (predorsal length 62.9-67.9% SL); anal-
fin base length about half of standard length (48.3-55.9% SL);
Fig. 1. Betta rubra - a. MSNG 13019 (lectotype), 29.4 mm SL,
Siboga; b. ZRC 42497, 33.4 mm SL, Aceh Barat (right side,
reversed).
a
b
pelvic fin rounded with short filamentous ray (28.1-38.4%
SL); pectoral fin rounded.
Preserved coloration. – See Figs. 1a, b for general
appearance. Dorsum of body dark brown, ventrum lighter
brown. Dorsum of head brown with black spots. Mouth with
lower lip black. Distinct pre- and postorbital dark brown
stripes on head, postorbital stripe interrupted by vertical
parallel reddish bars on opercle, opercle with distal dark
brown margin. Broad dark brown chin-bar present, which
merges with anterior part of second postorbital stripe, forming
a triangular dark brown mark below eye. Body with
irregularly spaced blackish vertical 5-7 bars, more distinct
on ventral half of body. Caudal peduncle spot faint. Fins
brownish, sometimes reddish. Dorsal, caudal and anal fins
without transverse bars on interradial membrane. Pectoral fin
rounded, and hyaline. Pelvic fin rounded, with reddish
filamentous second ray.
Distribution. – Betta rubra is currently known only from Lake
Toba area near Medan in Sumatra Utara; Aceh Barat in Aceh;
and Siboga in northern part of Sumatra Barat.
Remarks. – The recent collection (MZB 4784) had been
mislabeled as B. picta and had been badly preserved. All
specimens were darkly stained, fins broken and folded (Fig.
1b). As such, no colour information on relatively fresh
specimen can be made. The placing of B. rubra in the B.
foerschi group is tentative pending properly preserved fresh
collections (viz., Tan & Ng, 2005) (present volume). The
specimens from Genova (MSNG 13019) were in good
condition, but the opercle and pelvic, anal, caudal and dorsal
fins were tinged pinkish-red. On closer examination under a
microscope, the seemingly red coloration was found to be
due to either a side effect from early preservation chemicals
or a dye used previously. Nevertheless, this taxon is valid
from its unique suite of characters.

120
Tan & Ng: Anabantoidei and Channoidei of Sumatra
Betta picta species group
Betta falx Tan & Kottelat
Betta trifasciata (non Bleeker, 1850) - Günther, 1861: 388 (part);
Volz, 1905: 127; 1907: 127.
Betta pictum (non Valenciennes, in Cuvier & Valenciennes, 1846)
- Weber & de Beaufort, 1912: 541.
Betta picta (non Valenciennes, in Cuvier & Valenciennes, 1846) -
Bleeker, 1879: 26 (part); Weber & de Beaufort, 1922: 360 (part);
Kottelat et al., 1993: 163 (part).
Betta falx Tan & Kottelat, 1998b: 564.
Material examined. – Holotype - Jambi - MZB 9308 (holotype),
32.7 mm SL.
Paratypes - Jambi - ZRC 40974 (20 paratypes); ZMA 121.673 (6
paratypes); MZB 9307 (4 paratypes); RMNH 33087 (5 paratypes);
ZRC 38571 (44 paratypes); CMK 11119 (44 paratypes); ZRC 38254
(3 paratypes); ZRC 42496 (10 paratypes); MHNG 2593.95 (5
paratypes).
Others - Sumatra Utara - ZMA 121.590 (29), ZRC 40975 (2); ZMA
121.692 (5); ZMA 121.688 (2). Jambi - ZRC 42511 (3).
Remarks. – Betta falx is closely related to B. picta from Java
(see Tan & Kottelat, 1998b). It differs from B. picta in the
following characters: lower dorsal fin ray count (mode 8 vs.
9); lower lateral scale count (mode 27 vs. 28); dorsal fin origin
above 11-12th lateral scale (vs. 12-14th); anal fin origin below
modal 6th lateral scale (vs. 7th); lower predorsal scale count
(mode 19, vs. 20); slightly greater anal-fin base length (46.5-
50.3% SL, vs. 42.6-48.4); in life - male with distal margins
of anal and caudal fins reddish (vs. bluish); iridescent
greenish-blue opercle scales (vs. yellow-gold); preserved
material - male with distinct dorsal transverse bars (vs. faint);
male with distinct dark anal distal margin wide (vs. narrow);
male without elongated median caudal fin rays (vs. presence);
female with distinct caudal transverse bars (vs. very faint or
absent); dorsal head view narrow (vs. broad); thick preorbital
black stripe (vs. narrow); and the distance between posterior
part of anal fin to lower part of caudal narrow (vs. wide).
Betta pugnax species group
Betta cracens, new species
(Fig. 2)
Material examined. – Holotype - Jambi - MZB 9309 (holotype),
47.4 mm SL; Sungai Berliung Bata, Bertam, ca. 1 km into turnoff
to Permata Biru Indah, 10 km from Jambi towards Palembang after
main bus terminus, from fish collectors, coll. H. H. Tan & H. H.
Ng, 18 Jul.1997.
Paratypes - ZRC 41717 (6 paratypes), 32.8-57.4 mm SL, CMK
14415 (2 paratypes), 38.2-39.8 mm SL; same data as holotype. —
ZRC 42498 (2 paratypes), 31.2-51.3 mm SL; Sungai Berliung Bata,
Bertam, ca. 1 km into turnoff to Permata Biru Indah, 10 km from
Jambi towards Palembang after main bus terminal, coll. H. H. Tan,
29 Jul.1997.
Diagnosis. – Betta cracens can be differentiated from the
Fig. 2. Betta cracens - a. ZRC 41717 (paratype), 57.4 mm SL, freshly
preserved, Jambi (Bertam); b. ZRC 41717 (paratype), 57.4 mm SL,
Jambi (Bertam).
a
b
other members of the B. pugnax group in having the following
unique combination of characters: most slender body in the
species group (body depth at dorsal-fin origin 21.2-24.2%
SL); anal fin rays 27-29 (mode 28); lateral scales 32-33 (mode
33); thin dark blue edge on anal fin in male and female;
predorsal length 65.0-67.1% SL; head length 30.8-31.7% SL.
Description. – The general appearance is illustrated in Figs.
2a, b; meristic and morphometric data of B. cracens in Table
2. Body slender (body depth 21.2-24.2% SL), head short and
blunt (head length 30.8-31.7% SL; head width 17.1-19.9%
SL and 53.8-64.0% HL). Dorsal, caudal and anal fins pointed;
dorsal fin placed relatively far back (predorsal length 65.0-
67.1% SL); caudal fin with distal part of rays extended slightly
beyond interradial membrane, median rays elongated; anal-
fin base length more than half of standard length (53.4-55.7%
SL); pelvic fin rounded with relatively long filamentous ray
(36.1-43.4% SL); pectoral fin rounded.
Live coloration. – See Fig. 2a for general appearance. Dorsum
of body brown, ventrum lighter brown. Dorsum of head
brown, male with iridescent green opercle, female without
iridescent opercle. Distinct pre- and postorbital black stripes
on head, lower lip black, chin-bar present in both sexes and
juvenile. Male with some body scales on dorsal half with
blue iridescence, rest of body scales with faint dark brown
posterior edge, more distinct on belly scales. Body with 5-
7 faint irregularly spaced vertical brown bars. Female and
juvenile with indistinct central and second central stripes, and
indistinct caudal peduncle spot. Fins brownish, dorsal and
caudal fins with dark brown dorsal transverse bars in male,
only dorsal fin with transverse bars in female. Both male and
female with thin dark blue edge on anal fin. Pectoral fin
hyaline. Pelvic with bright whitish distal tip.
Preserved coloration. – See Fig. 2b for general appearance.
Dorsum of body dark brown, ventrum lighter brown to
whitish. Dorsum of head dark brown, male with distal part
of opercle after eye black. Distinct pre- and postorbital black

121
THE RAFFLES BULLETIN OF ZOOLOGY 2005
Table 2. Meristics and morphometrics of Betta cracens, B. fusca, B. raja and B. rubra.
Betta cracens Betta fusca Betta raja Betta rubra
Sample size 6 2 20 4
Standard length (mm) 38.2-57.4 53.1-59.7 45.9-63.9 33.1-35.0
MERISTICS
Vertebrae 2-3 + 8 + 20-21 2-3 + 8 + 19 2-3 + 8 + 18-19 2 + 8 + 18-19
(total 31-32, 31) (total 29-30, 29 or 30) (total 28-30, 29) (total 28-29, 29)
(n = 11) (n = 2) (n = 20) (n = 4)
Anal fin rays I-II, 26-27 0-I, 23-25 II, 23-26 II-III, 24-26
(total 27-29, 28) (total 24-25, 24 or 25) (total 25-28, 26) (total 27-29, 27)
Dorsal fin rays 0-I, 8-9 0-II, 7-8 I-II, 7-10 0-II, 7-8
(total 8-9, 8 or 9) (total 8-9, 8 or 9) (total 8-11, 9) (total 8-9, 9)
Caudal fin rays ii,5+6,ii ii,6+6,ii ii,5+6-7,i (ii,5+7,i) ii-iii,5+6,i-iii
Pelvic fin rays I, 1, 4 I, 1, 4 I, 1, 4 I, 1, 4
Pectoral fin rays 13 12 12-13 (13) 13
Anal fin rays below dorsal-fin origin - 9 10-13 (11) -
Anal fin rays at pectoral fin end - 5-6 6-9 (8) -
Anal fin rays at pelvic fin end - 11-12 11-17, females -
15 and above, males
Subdorsal scales 51/2-6 (51/2 or 6) 51/2-6 6-7 (61/2) 5-6 (6)
Transverse scales at dorsal-fin origin 91/2-10 (91/2) 10 10-101/2 (10) 81/2-91/2 (91/2)
Lateral scales 32-33 (33) 29 30-32 (31) 281/2-30 (29)
Lateral scales below dorsal-fin origin 16-19 (17 or 18) 16 14-16 (15) 15-16
Lateral scales above anal-fin origin 6-7 (7) 6 5-7 (6) 6-8 (6)
Predorsal scales 23-24 (23 or 24) 22-23 20-22 (22) 22-24 (24)
Postdorsal scales 11-12 (11 or 12) 11 10-11 (11) 10-11
MORPHOMETRICS - % SL
Total length 138.9-145.8 138.6 138.5-159.4 131.6-142.3
Predorsal length 65.0-67.1 68.5-70.2 63.2-69.9 62.9-67.6
Postdorsal length 20.4-24.6 22.2-23.5 20.0-24.6 20.3-23.3
Caudal peduncle depth 15.1-16.7 19.2-20.4 18.3-23.2 13.1-15.3
Preanal length 42.0-46.1 49.0-50.3 46.1-52.4 41.7-47.4
Head length 30.8-31.7 33.2-33.8 33.9-37.4 27.3-33.6
Body depth at dorsal-fin origin 21.2-24.2 30.9-31.7 28.1-33.5 23.2-25.8
Pelvic fin length 36.1-43.4 38.6-39.2 37.4-69.8 28.1-38.4
Anal-fin base length 53.4-55.7 47.8-51.4 48.9-54.8 49.2-52.8
Dorsal-fin base length 10.5-11.6 13.7-13.9 11.0-15.3 10.0-12.2
Body length 68.5-71.3 70.4-70.8 62.8-70.7 -
Head width 17.1-19.9 19.8-23.6 18.5-22.3 -
% HL
Orbit diameter 24.7-28.2 23.8-26.7 24.0-28.7 24.5-25.8
Postorbital length 46.3-50.7 48.3-50.5 45.1-53.9 49.5-57.1
Interorbital width 32.7-37.1 33.5-34.2 32.2-39.6 22.7-29.5
Snout length 24.0-25.8 25.0-26.7 26.3-29.8 17.9-24.7
Head width 53.8-64.0 59.7-69.8 51.8-62.0 -
% Postorbital length
Orbit diameter 50-58 47-55 46-64 -
Interorbital width 67-75 68-70 67-78 -
% Predorsal length
Head length 46.3-48.4 47.2-49.4 50.4-55.7 -
stripes on head, chin-bar present but may be obscured by
blackened opercle scales. Body scales with black posterior
edge, more prominent on belly scales. Body with irregularly
spaced 5-7 vertical dark brown bars. Female and juvenile
with indistinct central and second central stripes with caudal
peduncle spot indistinct. Fins brownish, dorsal and caudal
fins with dorsal transverse black bars in male, dorsal with
transverse black bars in female, anal fin with thin distal black
edge. Pectoral fin hyaline. Pelvic fin dark brownish, with
hyaline filamentous second ray.
Distribution. – Betta cracens is currently known only from
the swamp forest habitat in the Bertam area south of Jambi
town.
Field notes. – The type locality is a freshwater swamp forest,
part of it having been converted to rubber tree plantation.
The water lily, Barclaya motleyi (Nymphaeaceae) grows in
large mats at certain stretches of the stream line. Water depth
varied from 5 to 80 cm, pH 5.8. Syntopic species present:
Puntius banksi, Rasbora einthoveni, R. pauciperforata
(Cyprinidae), Hemirhamphodon pogonognathus
(Hemiramphidae), Betta pugnax, Sphaerichthys
osphromenoides, Trichogaster trichopterus
(Osphronemidae), Channa gachua and C. lucius (Channidae).

122
Tan & Ng: Anabantoidei and Channoidei of Sumatra
Etymology. – From the Latin cracens meaning slender,
graceful; alluding to the slender body, as compared with the
other members of the B. pugnax group.
Remarks. – Betta cracens can be differentiated from the other
members of the B. pugnax species group in the following
characters: most slender body (body depth at dorsal-fin origin
21.2-24.2% SL, vs. 24.8-36.9); higher anal fin ray count than
all members (mode 28, vs. 24-27); higher lateral scale count
than all members (mode 33, vs. 27-31); lower dorsal fin ray
count than B. pugnax, B. schalleri and B. prima (mode 8-9,
vs. 10-11); lower subdorsal scale count than B. schalleri, B.
pulchra, B. breviobesus (mode 51/2-6, vs. 61/2-7); higher
predorsal scale count than all members except B. fusca (mode
23-24, vs. 17-22); more postdorsal scales than B. prima (11-
12, vs. 9-10); smaller total length than B. enisae (138.9-
145.8% SL, vs. 143.0-155.2); smaller predorsal length than
B. fusca, B. prima and B. enisae (65.0-67.1% SL, vs. 67.3-
70.3); smaller caudal peduncle depth than B. fusca and B.
raja (15.1-16.7% SL, vs. 18.3-23.2); smaller preanal length
than all members except B. pugnax and B. pulchra (42.0-
46.1% SL, vs. 46.1-54.8); smaller head length than all
members except B. pugnax (30.8-31.7% SL, vs. 32.7-37.7);
greater anal-fin base length than B. fusca, B. prima, B. enisae,
B. schalleri and B. pulchra (53.4-55.7% SL, vs. 47.8-53.5);
smaller dorsal-fin base length than B. fusca, B. schalleri and
B. pulchra (10.5-11.6% SL, vs. 12.3-15.9); smaller head width
than B. fusca (17.1-19.9% SL, vs. 19.8-23.6); smaller snout
length than B. raja (24.0-25.8% HL, vs. 26.3-29.8); smaller
orbital diameter than B. prima, B. enisae and B. schalleri (50-
58% postorbital length, vs. 57-76); and a smaller head length
than B. prima, B. breviobesus and B. raja (46.3-48.4%
predorsal length, vs. 49.8-58.5).
Betta cracens can be differentiated from its syntopic species
B. pugnax in having the following characters: more anal fin
rays (27-29, vs. 23-24); fewer dorsal fin rays (8-9, vs. 10);
more lateral scales (32-33, vs. 29-30); dorsal-fin origin above
16-19th lateral scale (vs. 13-14th); more predorsal scales (23-
24, vs. 18-19); more postdorsal scales (11-12, vs. 10); greater
predorsal length (65.0-67.1% SL, vs. 63.0-64.2); smaller
caudal peduncle depth (15.1-16.7% SL, vs. 17.6-19.0);
smaller preanal length (42.0-46.1% SL, vs. 50.4-52.6); smaller
head length (30.8-31.7% SL, vs. 34.1-34.9); smaller body
depth (21.2-24.2% SL, vs. 27.8-27.9); greater anal-fin base
length(53.4-55.7% SL, vs. 46.8-46.9); smaller dorsal-fin base
length (10.5-11.6% SL, vs. 14.4-14.7); and a greater body
length (68.5-71.3% SL, vs. 65.3-67.2).
Betta fusca Regan
(Fig. 3)
Betta fusca Regan, 1910: 780, pl. 76, Fig. 2; Weber & de Beaufort,
1922: 356 (part); Kottelat et al., 1993: 162.
Betta pugnax (non Cantor, 1850) - Perugia, 1893: 243 (part); Volz,
1905: 127 (part); 1907: 127 (part).
Material examined. – Lectotype - Sumatra Utara - BMNH
1908.17.13:18 (lectotype), 59.7 mm SL male; Medan; coll. Morton. Fig. 3. Betta fusca - BMNH 1908.17.13:18 (lectotype), 59.7 mm
SL, Medan (caudal fin missing).
Paralectotype - Sumatra Utara - BMNH 1908.17.13:19 (1
paralectotype), 53.1 mm SL; sama data as lectotype.
Diagnosis. – Betta fusca can be differentiated from the other
members of the B. pugnax group by the following set of
characters: large head (head length 33.2-33.8% SL); anal fin
rays 24-25; lateral scales 29; body depth 30.9-31.7% SL;
predorsal length 68.5-70.2% SL; preanal length 49.0-50.3%
SL; body length 70.4-70.8% SL.
Description. – The general appearance is illustrated in Fig.
3; meristic and morphometric data of B. fusca in Table 2.
Body relatively stocky (body depth 30.9-31.7% SL), head
long and stocky (head length 33.2-33.8% SL; head width
19.8-23.6% SL and 59.7-69.8% HL). Dorsal, caudal and anal
fins pointed; dorsal fin placed relatively posteriorly (predorsal
length 68.5-70.2% SL); caudal fin with distal part of rays
extended slightly beyond interradial membrane, median rays
elongated; anal-fin base length about half of standard length
(47.8-51.4% SL); pelvic fin rounded with relatively long
filamentous ray (38.6-39.2% SL); pectoral fin rounded.
Preserved coloration. – See Fig. 3 for general appearance.
Dorsum of body dark brown, ventrum lighter brown to yellow.
Dorsum of head dark brown, male with distal part of opercle
after eye dark brown, almost black. Distinct pre- and
postorbital dark brown stripes on head, chin-bar absent. Body
scales with dark brown to black posterior edge, more
prominent on belly scales. Body with central and second
central stripes, and caudal peduncle spot indistinct. Fins
brown, dorsal fin with dorsal transverse bars, anal and caudal
transverse bars absent. Pectoral fin proximal part darkish, rest
hyaline. Pelvic fin brownish, with hyaline filamentous second
ray.
Distribution. – Betta fusca is currently known only from the
type series. Regan (1910) reported the type locality as
‘Sumatra’ and that they had been collected by W. Morton.
Their exact locality is not known, but Morton collected around
Medan. In the narrative of his travel (Morton, 1908: 165-
174), he mentioned having collected in ‘Tanjong Slamat’,
‘Bahsoemboe near Tebing Tinggi’, and foothills of ‘Mt. Surbo
Dolok’. The bulk of Morton’s collection is preserved in Musée
d’Histoire Naturelle, Lausanne, Switzerland; from Morton’s
text, it seems that some material was also deposited in
Naturhistorisches Museum, Zürich, Switzerland, but no
Morton fish material could be located in these two collections

123
THE RAFFLES BULLETIN OF ZOOLOGY 2005
(M. Kottelat, pers. comm.). Attempts had been made to collect
fresh material from Medan area, but we were not successful.
Remarks. – Betta fusca can be differentiated from the other
members of the species group in having the following
characters: fewer anal fin rays and dorsal fin rays than B.
schalleri (24-25, vs. 27; and 8-9, vs. 10-11); fewer anal fin
rays than B. pugnax, B. pulchra, B. breviobesus and B. cracens
(mode 24-25, vs. 26-29); fewer dorsal fin rays than B. pugnax
and B. prima (mode 8-9, vs. 10); fewer subdorsal scales than
B. schalleri (51/2-6, vs. 61/2-7); fewer subdorsal scales than B.
pulchra and B. breviobesus (mode 51/2-6, vs. 7); fewer lateral
scales than B. schalleri and B. cracens (29, vs. 31-33); more
lateral scales than B. enisae (29, vs. 26-28); more predorsal
scales than B. pugnax, B. schalleri, B. pulchra and B.
breviobesus (22-23, vs. 17-21); more postdorsal scales than
B. prima (11, vs. 9-10); fewer vertebrae than B. prima (total
29-30, vs. 28); greater predorsal length than B. schalleri, B.
pulchra, B. breviobesus and B. cracens (68.5-70.2% SL, vs.
60.8-67.5); smaller preanal length than B. prima and B. enisae
(49.0-50.3% SL, vs. 50.1-54.8); greater preanal length than
B. cracens (49.0-50.3% SL, vs. 42.0-46.1); smaller head
length than B. schalleri, B. prima and B. enisae (33.2-33.8%
SL, vs. 34.5-37.7) and smaller head length than B. breviobesus
(47.2-49.4% predorsal length, vs. 51.2-56.1); greater head
length than B. cracens (33.2-33.8% SL, vs. 30.8-31.7);
smaller body depth at dorsal-fin origin than B. schalleri and
B. enisae (30.9-31.7% SL, vs. 26.2-28.9); greater body depth
than B. cracens (30.9-31.7% SL, vs. 21.2-24.2); greater pelvic
fin length than B. prima (38.6-39.2% SL, vs. 30.1-37.2);
greater dorsal-fin base length than B. enisae and B. cracens
(13.7-13.9% SL, vs. 8.7-12.1); smaller anal-fin base length
than B. schalleri, B. prima, B. enisae, B. breviobesus and B.
cracens (47.8-51.4% SL, vs. 50.0-55.7); smaller orbital
diameter than B. pugnax, B. prima and B. pulchra (23.8-
26.7% HL, vs. 26.0-38.5) and smaller orbital diameter than
B. schalleri, B. prima and B. enisae (47-55% postorbital
length, vs. 57-76); smaller interorbital width than B. pulchra
(33.5-34.2% HL, vs. 34.3-41.8) and smaller interorbital width
than B. prima and B. enisae (68-69% postorbital length, vs.
70-84). Betta fusca shares with both B. schalleri and B. raja
in not having the following three characters: chin-bar, caudal
transverse bars, anal and caudal fin distal dark margins.
The larger specimen (BMNH 1908.17.13:18, 59.7 mm SL)
has been chosen as the lectotype although its caudal fin is
broken off. This is because the seemingly complete smaller
specimen (BMNH 1908.17.13:19, 53.1 mm SL) has damage
to the posterior half of its anal fin base, and it is scarred and
may lack several anal fin rays.
Betta raja, new species †
(Fig. 4)
Betta pugnax (non Cantor, 1850) - Volz, 1905: 127 (part); 1907:
127 (part); Kubota et al., 1996: 40.
Betta taeniata (non Regan, 1910) - Volz, 1905: 127 (part); 1907:
127 (part).
Material examined. – Holotype - Jambi - MZB 9315 (holotype),
56.3 mm SL male; Sungai Ayer Merah, feeder stream to Danau
Souak Padang ca. 15 mins. by boat upstream (1°36’45.7”S
103°27’00.0”E), coll. H. H. Tan & S. H. Tan, 21 Nov.1996.
Paratypes - Jambi - ZRC 42499 (12 paratypes), 19.8-58.1 mm SL,
MZB 9314 (4 paratypes), 32.5-57.5 mm SL; same locality as
holotype. — CMK (9 paratypes), ZRC 42501 (10 paratypes), 27.8-
57.6 mm SL; Sungai Bakong, tributary to Danau Arang Arang
(1°37’31.0”S 103°47’20.6”E), coll. H. H. Tan & H. H. Ng, 25
Jul.1997. — ZRC 42502 (14 paratypes), 47.5-63.8 mm SL; Danau
Rasau, opposite Kampung Rantau Panjang, coll. H. H. Tan et al.,
Jun.1996. Sumatra Selatan - ZRC 42500 (20 paratypes), 35.1-63.9
mm SL, ZMA 121.823 (5 paratypes), 36.3-50.7 mm SL, BMNH (5
paratypes), 37.5-50.7 mm SL; Sungai Sentang, ca. 20 mins. walk
through rubber plantation and swamp forest after 4.8 km drive in,
12 km from Jambi to Bayung Lencir (216 km to Palembang), near
Desa Sukajaya, coll. H. H. Tan, 27 Jul.1997.
Others - Riau - ZRC 9074 (1), 46.7 mm SL; Sungai Bengkwan,
tributary of Batang Kuantan (Indragiri), 4 hours downstream from
Rengat; H. H. Ng et al., 15 June 1995 [JMB9504]. — ZRC 42503,
7 ex., 46.2-55.9 mm SL; [JMBAQMT]. — ZMA 121.824 (1); Taluk;
J. P. Kleiweg de Zwaan, 1907. — ZMA 121.686 (22), 29.3-54.2
mm SL; Taluk; J. P. Kleiweg de Zwaan. Sumatra Barat - ZMA
121.687 (2), 33.7-41.5 mm SL; Sidjoengjoeng (=Sijunjung); J. P.
Kleiweg de Zwaan, 1907. Jambi - CMK 11033 (2); Danau Pinang,
lake connected to Sungai Pijoan, 1 hour by boat upriver of Pijoan
(19 km west of Jambi on road to Bungo); M. Kottelat & H. H. Tan,
28 May.1994. — ZRC 38529 (3), 28.3-45.4 mm SL, CMK 11087
(1); Sungai Bakung, north tributary of Sungai Kembang which joins
Danau Arang Arang and Kampung Kumpeh Hulu in Arang Arang;
M. Kottelat & H. H. Tan, 29 May.1994. — ZRC 38612 (6), 36.0-
63.8 mm SL, CMK 11159 (7); Danau Rasau, a blackwater lake
draining to Batang Hari, opposite Kampung Rantau Panjang; M.
Fig. 4. Betta raja - a. not preserved, ca. 50 mm SL, Jambi (Pijoan);
b. ZRC 38612, 64.2 mm SL, Jambi (Danau Rasau) (right side,
reversed).
a
b

124
Tan & Ng: Anabantoidei and Channoidei of Sumatra
Kottelat & H. H. Tan, 1-2 Jun.1994. — ZRC 42504 (16), 54.3-65.6
mm SL; Rantau Panjang, from aquarium trade, coll. H. H. Tan &
S. H. Tan, 21 Jul.1993. — ZRC 42505 (16), 50.2-62.8 mm SL;
Rantau Panjang, from fish collectors, coll. H. H. Tan et al., Jun.1996.
— ZRC 42506 (2), 48.9-50.2 mm SL; Darkie water reserve
(1°17’54.8”S 104°08’30.4”E), coll. H. H. Tan et al., 6 Jun.1996. —
ZRC 42507 (4), 28.0-48.1 mm SL; second stream on road towards
Muara Bulian, after interjunction at crossroads towards Palembang,
Muara Bulian and Jambi (1°47’44.4”S 103°25’16.1”E), coll. H. H.
Tan et al., 7 Jun.1996. — ZRC 42508 (2), 44.6-46.7 mm SL; Sungai
Pijoan (1°35’35.0”S 103°27’07.2”E), coll. H. H. Tan et al., 8
Jun.1996. — ZRC 42509 (17), 30.1-54.2 mm SL; Pijoan, Danau
Souak Padang (1°36’34.4”S 103°26’55.1”E), coll. H. H. Tan et al.,
8 Jun.1996. — ZRC 42510 (7), 21.3-57.7 mm SL; Leibong Sepbaju,
stream in swampforest near Pijoan, coll. H. H. Tan & S. H. Tan, 21
Nov.1996. — ZMA 114.368 (2), 45.6-51.6 mm SL; Jambi; P. H.
Moolenburgh, 19 Jul.1909. Sumatra Selatan - CMK 11447 (1),
24.6 mm SL; Sumatra Selatan: Sungai Simpang, small tributary
entering Sungai Niru from east immediately upriver of Muara Niru;
M. Kottelat, 2 Feb.1995 (MK 95-013).
Diagnosis. – Betta raja can be differentiated from the other
members of the B. pugnax group by the following set of
characters: mature males with very long pelvic fin (15-23
anal fin rays at pelvic fin end, even exceeding anal-fin base
in some specimens); dark margins on anal fin and lower half
of caudal fin; anal fin rays 25-28 (mode 26); lateral scales
30-32 (mode 31); head length 33.9-37.4% SL; predorsal
length 20.0-24.6% SL; preanal length 46.1-52.4% SL; body
length 62.8-70.7% SL.
Description. – The general appearance is illustrated in Figs.
4a, b; meristics and morphometrics of B. raja is listed in Table
2. Body relatively stocky (body depth 28.1-33.5% SL), head
long and stocky (head length 33.9-37.4% SL; head width
18.5-22.3% SL and 51.8-62.0% HL). Dorsal, caudal and anal
fins pointed; dorsal fin placed relatively far back (predorsal
length 63.2-69.9% SL); caudal fin with distal part of rays
extended slightly beyond interradial membrane, median rays
elongated; anal-fin base length about half of standard length
(48.9-54.8% SL); pelvic fin rounded with long filamentous
ray (37.4-69.8% SL); pectoral fin rounded.
Live coloration. – See Fig. 4b for general appearance. Dorsum
of body brown, ventrum lighter brown to yellow. Dorsum of
head brown, male with iridescent green opercle, female with
gold opercle. Distinct pre- and postorbital black stripes on
head, lower lip black. Male with some body scales on dorsal
half with blue, other scales near belly gold, female without
gold scales, rest of body scales with faint dark brown posterior
edge. Body with faint irregularly spaced 3-5 vertical brown
bars. Female and juvenile with more distinct central and
second central stripes with caudal peduncle spot. Fins
brownish, dorsal fin with dark brown dorsal transverse bars.
Both male and female with diffuse blue colour on anal fin
distal margin, small blackish irregularly spaced spots may
be present on anal fin in some specimens. All fins with thin
whitish distal edges, except hyaline pectoral fin. Pelvic fin
with bright whitish distal tip.
Preserved coloration. – See Fig. 4a for general appearance.
Dorsum of body dark brown, ventrum lighter brown to
whitish. Dorsum of head dark brown, male with distal part
of opercle black. Distinct pre- and postorbital dark brown
stripes on head, chin-bar absent. Body scales with black
posterior edge, more prominent on belly scales. Body with
irregularly spaced 3-5 vertical dark brown bars. Male, female
and juvenile with distinct central and second central stripes
with caudal peduncle spot distinct. Fins brownish to greyish,
dorsal fin with dorsal transverse bars, anal fin sometimes with
transverse bars, caudal transverse bars absent. Pectoral fin
hyaline. Pelvic fin brownish, with whitish filamentous second
ray.
Distribution. – Betta raja is found in the lowland swamp
forests of the Batang Hari basin in Jambi, the northern part
of Sumatra Selatan, in the Indragiri basin in Riau and eastern
part of Sumatra Barat in Sijunjung.
Field notes. – In Danau Rasau, a blackwater lake draining
to Batang Hari, fisherfolk catch them with hook-and-line and
use them as bait for catching Channa. Syntopic species
present: Osteochilus spilurus, Parachela oxygastroides,
Puntius johorensis, Rasbora cephalotaenia (Cyprinidae),
Mystus bimaculatus, Pseudomystus leiacanthus (Bagridae),
Kryptopterus macrocephalus, Silurichthys indragirensis
(Siluridae), Pseudeutropius brachypopterus, P.
moolenburghae (Schilbeidae), Chaca bankanensis
(Chacidae), Nandus nebulosus (Nandidae), Helostoma
temminckii (Helostomatidae), Belontia hasseltii, Betta
coccina, Betta renata, Luciocephalus pulcher,
Parosphromenus sumatranus, Sphaerichthys
osphromenoides, Trichogaster leerii (Osphronemidae),
Channa lucius, C. striata (Channidae). Betta raja is a paternal
oralbrooder (pers. observ.; Kubota et al., 1996: 40).
Etymology. – From the Malay vernacular Raja, meaning king
or prince, also from the Indonesian vernacular for this species
- Chupang Raja. A noun in apposition.
Remarks. – Betta raja can be differentiated from the other
members of the species group by the following characters:
mature males with very long pelvic fin (15-23 anal fin rays
at pelvic fin end, even exceeding anal-fin base in some
specimens); more anal fin rays than B. fusca (25-28, vs. 24-
25); fewer anal fin rays than B. schalleri (mode 26, vs. 27);
fewer dorsal fin rays than B. pugnax, B. schalleri and B. prima
(mode 9, vs. 10-11); more subdorsal scales than B. enisae
(6-7, vs. 5-51/2); more lateral scales than B. fusca, B. prima
and B. enisae (30-32, vs. 26-29); fewer lateral scales than B.
cracens (mode 31, vs. 33); more predorsal scales than B.
schalleri (20-22, vs. 17-19); fewer predorsal scales than B.
cracens (20-22, vs. 23-24); smaller postdorsal length than B.
prima (20.0-24.6% SL, vs. 24.1-26.3); greater head length
than B. fusca and B. cracens (33.9-37.4% SL, vs. 30.8-33.8;
50.4-55.7% predorsal length, vs. 46.3-49.4); greater body
depth at dorsal-fin origin than B. schalleri, B. enisae and B.
cracens (28.1-33.5% SL, vs. 21.2-28.9); greater caudal
peduncle depth than B. cracens (18.3-23.2% SL, vs. 15.1-
16.7); greater preanal length than B. cracens (42.0-46.1% SL,
vs. 46.1-52.4); greater pelvic fin length than B. prima (37.4-
69.8% SL, vs. 30.1-37.2); greater dorsal-fin base length than

125
THE RAFFLES BULLETIN OF ZOOLOGY 2005
B. enisae (37.4-69.8% SL, vs. 8.7-12.1); slightly smaller head
width than B. fusca (51.8-62.0%HL, vs. 59.7-69.8); greater
snout length than B. cracens (26.3-29.8% HL, vs. 24.0-25.8);
smaller orbital diameter than B. pugnax (24.0-28.7% HL, vs.
27.9-38.5) and B. schalleri (46-64% postorbital length, 61-
76).
Betta pugnax (Cantor)
Betta pugnax - Volz, 1905: 127 (part); 1907: 127 (part); SH Tan &
HH Tan, 1994: 357.
Material examined. – Riau - ZRC 42516 (11). Riau Archipelago
- ZMA 121.587 (2); ZRC 31452-57 (6); ZRC 14014-27 (10); ZRC
22294-95 (2); ZRC 38262 (11); ZRC 34831-54 (24); ZRC 38331
(28); ZRC 37536 (8). Jambi - ZRC 42517 (6).
Remarks. – Betta pugnax is found in Pulau Batam, Pulau
Bintan, Pulau Lingga and Pulau Singkep of the Riau
Archipelago and Sumatra. Betta pugnax has a wide
distribution range and can be found throughout the Malay
Peninsula (Tan & Tan, 1996).
Betta cf. pugnax
Material examined. – Nias - ZMA 121.592 (11).
Remarks. – The specimens from Nias (ZMA 121.592) are
tentatively identified as B. pugnax and have the following
characters: dark opercle, dorsal and caudal transverse bars
present in male, absent in female; distinct chin-bar in juvenile,
faint in adults; total anal fin rays 25-27; total dorsal fin rays
9-10; lateral scales 29-30; predorsal scales 21-22. To ascertain
their true identity, fresh specimens would be required.
Betta schalleri Kottelat & Ng
Betta schalleri Kottelat & Ng, 1994: 74, figs. 9-11.
Material examined. – Holotype - Banka - ZRC 35170 (holotype),
44.8 mm SL.
Paratypes - Banka - ZRC 35171-73 (3 paratypes); CMK 9639 (3
paratypes); ZRC 35174-78 (5 paratypes), CMK 9585 (4 paratypes).
Remarks. – Betta schalleri is closely allied to both B. fusca
and B. raja. It is restricted to the island of Banka. Betta
schalleri can be differentiated from the other members from
Sumatra in the following characters: more anal fin rays than
B. fusca (27, vs. 24-25); more dorsal fin rays than B. cracens
and B. fusca (10-11, vs. 8-9); more subdorsal scales than B.
cracens and B. fusca (61/2-7, vs. 51/2-6); more lateral scales
than B. fusca (31, vs. 29); fewer lateral scales than B. cracens
(31, vs. 32-33); fewer predorsal scales than B. cracens, B.
fusca and B. raja (17-19, vs. 20-24); greater head length than
B. pugnax, B. cracens and B. fusca (35.5-36.5% SL, vs. 27.5-
35.2); smaller predorsal length than B. fusca (62.7-66.3% SL,
vs. 68.5-70.2); greater preanal length than B. cracens (47.8-
50.9% SL, vs. 42.0-46.1); greater body depth than B. cracens
(26.7-27.6% SL, vs. 21.2-24.2); greater dorsal fin base length
than B. cracens (12.4-13.9% SL, vs. 10.5-11.6); smaller anal
fin base length than B. cracens (50.4-52.0% SL, vs. 53.4-
55.7); greater orbital diameter than B. cracens and B. fusca
(61-76% postorbital length, vs. 47-58).
BETTA SPELENDENS SPECIES GROUP
Betta imbellis Ladiges
Betta imbellis - Schaller & Kottelat, 1990: 36; Kottelat et al., 1993:
162; Tan & Ng, 2005: 78 (present volume).
Material examined. – Sumatra Utara - ZRC 40966 (17); CMK
13126 (5).
Betta splendens Regan
Material examined. – Jambi - ZRC 42281 (5); ZRC 42461 (22).
Remarks. – The population of Betta splendens found in the
highlands of Kerinci is most probably introduced. This
population has adapted to the colder waters and did not take
well to the warmer temperatures of captivity in the lowlands.
BETTA WASERI SPECIES GROUP
Betta chloropharynx Kottelat & Ng
Betta chloropharynx Kottelat & Ng, 1994: 72, figs. 6-8.
Material examined. – Holotype - Banka - ZRC 35166 (holotype),
58.6 mm SL.
Paratypes – Banka - ZRC 35167 (1 paratype); CMK 9761 (1
paratype).
Remarks. – Betta chloropharynx belongs to the B. waseri
group, as had been discussed by Tan (1998) and Tan &
Kottelat (1998a). It is restricted to the island of Banka. Betta
chloropharynx can be separated from B. hipposideros in the
following characters: µ-shaped black throat mark (vs. horse-
shoe shape black mark); absence of both dorsal and caudal
transverse bars (vs. presence); fewer subdorsal scales (mode
5-6, vs. 61/2). Betta chloropharynx can be differentiated from
B. renata in the following characters: µ-shaped black throat
mark (vs. kidney-shaped); absence of both dorsal and caudal
transverse bars (vs. presence); non-spotted opercle (vs. spotted
opercle); opercle with yellow lower margin (vs. black
margin); fewer subdorsal scales (mode 5-6, vs. 61/2). Betta
chloropharynx can be distinguished from B. spilotogena in
the following characters: µ-shaped black throat mark (vs.
black median spot); non-spotted opercle (vs. spotted opercle);
and an opercle with yellow lower margin (vs. black margin).
Betta hipposideros Ng & Kottelat
Betta anabatoides (non Bleeker, 1851) - Weber & de Beaufort, 1922:
357 (part).

126
Tan & Ng: Anabantoidei and Channoidei of Sumatra
Betta hipposideros - Tan & Ng, 2005: 84 (present volume).
Betta waseri (non Krummenacher, 1986) - Kottelat et al., 1993: 163
(part).
Material examined. – Riau - ZRC 39630 (2); ZMA 121.683 (3).
Riau Archipelago - ZRC 44058 (14).
Remarks. – Betta hipposideros is a stenotopic blackwater
species and its range extension had been noted by Tan & Ng
(2005). Betta hipposideros can be separated from B.
chloropharynx in the following characters: horseshoe-shaped
black throat mark (vs. µ-shaped black throat mark); presence
of both dorsal and caudal transverse bars (vs. absence); more
subdorsal scales (mode 61/2, vs. 5-6). Betta hipposideros can
be differentiated from B. renata in the following characters:
horseshoe-shaped black throat mark (vs. kidney-shaped); non-
spotted opercle (vs. spotted opercle); opercle with yellow or
brown lower margin (vs. black margin); fewer postdorsal
scales (mode 12, vs. 91/2); more anal fin rays (mode 30, vs.
29); more lateral scales (mode 32, vs. 31). Betta hipposideros
can be distinguished from B. spilotogena in the following
characters: horseshoe-shaped black throat mark (vs. black
median spot); non-spotted opercle (vs. spotted opercle);
opercle with yellow or brown lower margin (vs. black
margin); presence of both dorsal and caudal transverse bars
(vs. absence); and more subdorsal scales (mode 61/2, vs. 51/2).
Betta renata Tan †
Betta anabatoides (non Bleeker, 1851) - Weber & de Beaufort, 1922:
357 (part); Hardenberg, 1931: 125; 1933: 308.
Betta renata Tan, 1998: 282, figs. 1-2.
Material examined. – Holotype - Jambi - MZB 9310 (holotype),
70.8 mm SL.
Paratypes - Jambi - ZRC 39265 (1 paratype); ZRC 39117 (3
paratypes); ZRC 40285 (7 paratypes); CMK 13024 (3 paratypes);
ZRC 40286 (5 paratypes).
Others - Jambi - ZRC 40287 (1); ZRC 40288 (1); ZRC 42519 (1);
ZMA 121.684 (4). Sumatra Selatan - NMBA 2298-2300 (3);
NMBA 2303-04 (2); ZMA 121.682 (9).
Remarks. – Betta renata is a stenotopic blackwater species
and can be differentiated from B. chloropharynx in the
following characters: kidney-shaped black throat mark (vs.
µ-shaped black throat mark); presence of both dorsal and
caudal transverse bars (vs. absence); spotted opercle (vs. non-
spotted); opercle with black lower margin (vs. yellow margin);
more subdorsal scales (mode 61/2, vs. 5-6). Betta renata can
be separated from B. hipposideros in the following characters:
kidney-shaped black throat mark (vs. horse-shoe shape);
spotted opercle (vs. non-spotted); opercle with black lower
margin (vs. yellow or brown); more postdorsal scales (mode
12, vs. 91/2); fewer anal fin rays (mode 29, vs. 30); fewer
lateral scales (mode 31, vs. 32). Betta renata can be
distinguished from B. spilotogena in the following characters:
kidney-shaped black throat mark (vs. black median spot);
more subdorsal scales (mode 61/2, vs. 51/2); more postdorsal
scales (mode 12, vs. 10); fewer anal fin rays (mode 29, vs.
30); and fewer lateral scales (mode 31, vs. 32).
Betta cf. renata
Betta anabatoides - Bleeker, 1852a: 89; de Beaufort, 1939: 193.
Betta picta - Bleeker, 1879: 26.
Material examined. – Biliton - ZMA 121.695 (46, but only 10
largest specimens were examined), 56.9-76.8 mm SL.
Remarks. – The present series of specimens from Biliton are
in poor condition. They are stained dark brown, had been
previously dried out, the bodies are twisted and had clamped
and rigid fins. They are tentatively identified as B. renata
based on the following characters: a faint black rounded mark
on throat below lower jaw, not joined to black lower jaw;
faint spots on opercle; dorsal and caudal transverse bars; 2
rows of 8-12 black spots in the belly region above the anal
fin; total anal fin rays 29-31; lateral scales 30-32; and
subdorsal scales 6-61/2. Fresh material would be necessary to
ascertain their true identity.
Betta spilotogena Ng & Kottelat
Betta spilotogena Ng & Kottelat, 1994: 606, fig. 8; SH Tan & HH
Tan, 1994: 357.
Betta anabatoides - Weber & de Beaufort, 1922: 357 (part).
Material examined. – Holotype - Riau Archipelago - ZRC 35417
(holotype), 69.7 mm SL.
Paratypes - Riau Archipelago - ZRC 35418-20 (3 paratypes); CMK
10021 (2 paratypes); ZRC 35421-31 (11 paratypes); MZB (3
paratypes); ZRC 34783 (1 paratype); ZRC 34748 (1 paratype).
Others - Riau Archipelago - ZMA 121.586 (1).
Remarks. – Betta spilotogena is restricted to the Pulau Bintan
and Pulau Singkep of the Riau Archipelago. It is closely
related to B. renata (see Tan, 1998). Betta spilotogena can
be differentiated from B. chloropharynx in the following
characters: median black throat spot (vs. two rounded black
marks); spotted opercle (vs. non-spotted); opercle with black
lower margin (vs. yellow margin). Betta spilotogena can be
separated from B. hipposideros in the following characters:
median black throat spot (vs. horse-shoe shape mark); spotted
opercle (vs. non-spotted); opercle with black lower margin
(vs. yellow or brown); more postdorsal scales (mode 10, vs.
91/2). Betta spilotogena can be distinguished from B. renata
in the following characters: black median spot (vs. kidney-
shaped mark); absence of dorsal and caudal transverse bars
(vs. presence); fewer subdorsal scales (mode 51/2, vs. 61/2);
fewer postdorsal scales (mode 10, vs. 12); more anal fin rays
(mode 30, vs. 29); and more lateral scales (mode 32, vs. 31).
Luciocephalus pulcher (Gray) †
(Fig. 5)
Diplopterus pulcher Gray, 1830: Pl. 87; 1831: 8.
Luciocephalus pulcher - Bleeker, 1851: 273; 1852a: 99; 1879: 29;
Günther, 1861: 390; Károli, 1881: 172; Steindachner, 1901: 434;
Vaillant, 1902: 19; Duncker, 1904: 165; Volz, 1904: 459; 1905:

127
THE RAFFLES BULLETIN OF ZOOLOGY 2005
127; 1907: 127; Weber & de Beaufort, 1912: 541; 1922: 369,
fig. 94; Vinciguerra, 1926: 566; Hardenberg, 1935-36: 244;
Herre & Myers, 1937: 70; Fowler, 1938: 124; de Beaufort, 1939:
194; Tweedie, 1952: 65; 1953: 48, pl. 9, fig. 1; Klausewitz, 1955:
322; Alfred, 1966: 54; Imaki et al., 1981: 43, fig. 106; Mizuno
& Furtado, 1982: 324; Mohsin & Ambak, 1983: 176, fig. 133;
Brown & Brown, 1987: 163; Kottelat, 1989: 20; Roberts, 1989:
178; Lim & Ng, 1990: 102; Lim et al., 1990: 50; Kottelat et al.,
1992: 11; Ng et al., 1992: 29, fig. 103; Kottelat et al., 1993:
159, pl. 74; Ng et al., 1994: 210; SH Tan & HH Tan, 1994: 357;
Kottelat & Lim, 1995: 247; Vidthayanon et al., 1997: 55; Sim,
2002: 71; Vidthayanon, 2002: 115; Tan & Lim, 2004: 111.
Material examined. – Neotype - PENINSULAR MALAYSIA:
ZRC 42520, neotype, 106.2 mm SL; Johor: Mersing area, stream
ca. km 66 to Kluang, km 166 to Batu Pahat, coll. H. H. Tan et al.,
9 May.1995 .
Others - Terengganu - ZRC 23980-84 (5), 47.2-102.3 mm SL; ZRC
40204 (3), 50.4-56.2 mm SL. Perak - ZRC 39919 (4), 35.8-81.3
mm SL. Selangor - ZRC 28530-31 (2), 84.8-106.4 mm SL; ZRC
15054 (1), 128.3 mm SL. Pahang - ZRC 25117-19 (3), 59.0-67.8
mm SL; ZRC 420 (2), 84.6-94.0 mm SL. Johor - ZRC 39996 (1),
80.9 mm SL; ZRC 42521 (1), 84.3 mm SL; ZRC 42522 (1), 73.7
mm SL; ZRC 42523 (1), 85.8 mm SL; ZRC 42524 (6), 23.9-82.1
mm SL. Singapore: ZRC 19140 (1), 82.4 mm SL; ZRC 9600-04
(5), 58.7-97.6 mm SL; ZRC 1210 (1), 74.8 mm SL; ZRC 1209 (1),
105.0 mm SL; ZRC 1254 (1), 100.1 mm SL; ZRC 1253 (1), 106.1
mm SL. SUMATRA: Sumatra Utara - ZMA 121.701 (2), 43.4-
54.4 mm SL. Riau - ZRC 39077 (5), 54.8-99.2 mm SL; ZRC 42531
(1), 31.4 mm SL; ZRC 39048 (1), 66.6 mm SL. Riau Archipelago
- ZRC 37534 (1), 122.9 mm SL; ZRC 37552 (7), 12.0-118.4 mm
SL; ZRC 42532 (5), 47.5-80.0 mm SL; ZRC 42533 (1), 80.5 mm
SL; ZRC 42534 (6), 39.3-95.4 mm SL; ZRC 42535 (1), 103.8 mm
SL; ZRC 42536 (4), 77.8-93.9 mm SL; ZRC 42537 (1), 99.2 mm
SL; ZRC 38272 (2), 12.1-44.3 mm SL. Jambi - SMF 3364 (12),
42-114 mm TL; SMF 3605 (33), 61-124 mm TL; ZMA 121.700
(5), 47.0-106.0 mm SL; ZRC 37534 (4), 39.2-130.7 mm SL; ZRC
38541 (3), 21.6-69.4 mm SL; ZRC 42525 (10), 40.3-67.3 mm SL;
ZRC 42526 (8), 9.2-86.9 mm SL; ZRC 42527 (1), 85.6 mm SL;
ZRC 42528 (2), 109.4-138.1 mm SL; ZRC 42529 (8), 49.2-90.0
mm SL. Sumatra Selatan - ZRC 42530 (1), 44.0 mm SL. Banka
- ZRC 31006-08 (3), 40.6-58.5 mm SL. Biliton - ZRC 46302 (1),
33.6 mm SL. BORNEO: Sarawak: ZRC 37968 (1), 134.2 mm
SL; ZRC 37924 (2), 88.3-103.9 mm SL; ZRC 37956 (3), 58.4-110.5
mm SL; ZRC 39505 (3), 17.5-96.7 mm SL; ZRC 39501 (1), 69.7
mm SL; ZRC 42538 (6), 68.3-126.3 mm SL; ZRC 422 (1), 141.9
mm SL. Brunei: ZRC 31818-19 (2), 35.5-44.6 mm SL; ZRC 42539
(5), 19.6-76.9 mm SL; ZRC 42540 (2), 15.4-123.7 mm SL.
Diagnosis. – Luciocephalus pulcher differs from L. aura in
lacking numerous iridescent green spots on central stripe
when live (vs. presence); presence of black irregular pattern
on area below preorbital to central stripe (vs. absence); area
just above anal-fin base with 3-4 dark brown blotches, which
appear circular when body is viewed ventrally (vs. absence
of such markings); yellowish caudal fin with 3-5 broad
vertical black bars (vs. brownish caudal fin with 3-7 narrow
iridescent vertical gold bars); presence of markings on anal
and pelvic (vs. absence); smaller caudal-fin length (total
length 120.0-125.6% SL, vs. 123.7-127.1); smaller postdorsal
length (10.4-12.4% SL, vs. 12.0-13.2); greater head length
(44.1-48.6% SL, vs. 43.3-45.9); smaller caudal peduncle
depth (8.9-10.6% SL, vs. 10.7-11.2); and relatively shorter
anal-fin base length (19.3-23.4% SL, vs. 23.6-24.5).
Description. – General body form as in Figs. 5a, b; meristics
and morphometrics listed in Table 3. Head long and slender,
almost half of standard length (head length 44.1-48.6% SL,
head width 9.5-13.4% SL); elongated snout with long
protrusible jaws (snout length 18.8-21.9% SL and 41.6-45.1%
HL, upper jaw length 35.8-39.5% HL, lower jaw length 45.7-
52.2% HL); body slender (body depth 13.6-17.6% SL); dorsal
fin oval-shaped, situated posteriorly, near caudal fin
(predorsal length 77.5-81.4% SL), dorsal-fin base about half
of anal-fin base (40-50% anal-fin base length); caudal-fin
asymmetrically pointed; a deep notch on anal fin at proximal
6-8th anal ray, notch at about one third of anal-fin base (29.1-
43.8% anal-fin base length); pelvic fin with filamentous first
ray, with extension about twice or more as long as other pelvic
rays; pectoral fin rounded.
Live coloration. – Body pattern consists of a dark brown
dorsum, whitish ventrum; preorbital and postorbital broad
dark brown stripe joining with broad central dark brown
stripe, having 6-17 black spots on either dorsal or ventral
borders of central stripe, area just above central stripe golden;
opercle area below preorbital and postorbital stripes with
interrupted black stripe; pectoral fin hyaline, area just below
pectoral fin with black stripe which may be interrupted; pelvic
fin with 2-4 proximal black marks, rest hyaline; anal fin with
very faint to distinct black marks near base, rest hyaline, area
just above anal-fin base with 3-4 dark brown semi-circular
blotches, which appear circular when body is viewed
ventrally; dorsal fin with anterior 3-4 rays proximally
pigmented dark brown, rest hyaline; caudal fin yellowish with
3-5 broad vertical black bars, either in a regular or irregular
pattern.
Preserved coloration. – Preserved coloration illustrated in
Fig. 5. a. Diplopterus pulcher – reproduction of Gray, 1830: pl. 87;
b. Luciocephalus pulcher - ZRC 42520 (neotype), 106.2 mm SL,
Malaysia: Johor.
a
b

128
Tan & Ng: Anabantoidei and Channoidei of Sumatra
Table 3. Meristics and morphometrics of Luciocephalus pulcher and L. aura.
Luciocephalus pulcher Luciocephalus aura
sample size, n 23 6
MERISTICS
vertebrae 18-19 + 20-22 17-19 + 20-22
(total 38-40, 39) (n = 28) (total 38-40, 39) (n = 13)
dorsal-fin ray 10-12 (11) 11-12 (11)
anal-fin ray 0-I, 19-21 (total 19-21, 19) 0, 18-21 (total 18-21, 20)
pelvic-fin ray I, 5 I, 5
pectoral-fin ray 16-18 (17) 16-17 (17)
subdorsal scales 71/2-8 (81/2)8
1/2
transverse scales 141/2-161/2 (151/2)15
1/2
lateral scales 42-45 (43) 43
predorsal scales 60-65 (61) 60-63 (60)
postdorsal scales 11-14 (13) 11-12 (12)
caudal peduncle scales 91/2-12 (11) 10-11 (11)
lateral scales below dorsal-fin origin 25-27 (26) 25-28 (26)
lateral scales above anal-fin origin 15-17 (15) 15-16 (15)
MORPHOMETRICS - % SL
total length 120.0-125.6 123.7-127.1
body length 50.8-56.8 54.9-56.9
predorsal length 77.5-81.4 77.1-79.1
postdorsal length 10.4-12.4 12.0-13.2
preanal length 62.9-68.2 62.5-63.8
caudal peduncle length 11.1-15.2 11.4-14.0
head length 44.1-48.6 43.3-45.9
body depth 13.6-17.6 15.4-17.7
caudal peduncle depth 8.9-10.6 10.5-11.2
anal-fin base length 19.3-23.4 23.3-24.9
dorsal-fin base length 8.3-10.4 10.1-11.0
distance between pelvic- and anal-fin origins 18.1-22.0 18.1-20.3
pelvic-fin length 38.7-48.2 39.9-46.9
head width 9.5-13.4 9.8-10.7
snout length 18.8-21.9 18.7-20.2
% HL
orbit diameter 13.7-17.0 13.3-14.1
postorbital length 38.2-43.2 40.0-42.9
head width 21.5-28.9 22.3-24.7
interorbital width 16.4-19.6 15.8-18.6
snout length 41.6-45.1 42.6-45.3
upper jaw length 35.8-39.5 34.9-38.2
Fig. 5b. Body with dark brown dorsum; whitish ventrum
covered faintly or strongly with dark brown irregular pattern,
often with 3-4 distinct dark brown semi-circular blotches just
above anal-fin base, which appear as circular markings when
body is viewed ventrally; broad dark brown preorbital and
postorbital stripe joining with central stripe, ending at caudal
peduncle with a black largish spot; middle of central stripe
strongly pigmented or faint with dorsal and ventral borders
covered with 6-17 black spots; area just above central stripe
cream with 0-10 black spots on dorsal edge; area beneath
preorbital to central stripe covered with faint or strong dark
brown to black irregular markings; lower jaw whitish with
black edge; pectoral fin hyaline; pelvic fin with 2-4 proximal
black marks, rest hyaline; anal fin with very faint to distinct
black marks near base, rest hyaline; dorsal fin with anterior
3-4 rays proximally pigmented black, rest hyaline; caudal fin
whitish with 3-5 vertical black bars, either in a regular or
irregular pattern.
Distribution. – Luciocephalus pulcher is known from
Narathiwat province (Peninsular Thailand); Terengganu,
Perak, Selangor, Pahang, Johor (Peninsular Malaysia); Pulau
Bintan, Pulau Batam, Pulau Lingga, Riau, Jambi, Palembang,
Banka, Biliton (Sumatra); Brunei Darussalam; north, central
and south Sarawak; Kalimantan Barat; Kalimantan Tengah;
and the island of Natuna Besar.
Field notes on biology. – Luciocephalus pulcher is found
only in freshwater swamp forest and peat swamp habitats. It
is usually found among submerged vegetation along the
banks. It is an ambush predator, suddenly extending its funnel-
like mouth and engulfing its prey. The use of suction to ingest
prey is negligible (Lauder & Liem, 1981). It is a paternal
oralbrooder, up to 90 eggs (3-4 mm diameter) have been found
in the gular pouch (Kottelat et al., 1993: 159). Details on its
spawning as documented by Britz (1994b).

129
THE RAFFLES BULLETIN OF ZOOLOGY 2005
Remarks. – Further to the characters mentioned in the
diagnosis, L. pulcher can be differentiated from L. aura in
having slightly greater preanal length (62.9-68.2% SL, vs.
62.5-63.8); slightly smaller postdorsal length (18.6-23.4%
body length, vs. 21.0-24.1); slightly greater caudal peduncle
length (21.9-28.6% body length, vs. 20.0-24.7); smaller anal-
fin base length (35.4-42.3% body length, vs. 41.2-44.4); and
greater orbital diameter (13.7-17.0% HL, vs. 13.3-14.1). As
the populations of L. pulcher are many and widespread, there
is a fair amount of variation in morphometrics to be expected
from L. pulcher.
The name Diplopterus pulcher was first used by Gray in the
caption of a plate (1830: pl. 87; Fig. 5a) and was later
described briefly by Gray (1831: 8) based on specimen(s)
collected by Hardwicke from the British territory of India
(which included Peninsular Malaysia, which is near the
northernmost known limit of the distribution of L. pulcher).
No types were designated, but a colour plate was published
(Gray, 1830: Pl. 87). Bleeker (1851) replaced the name
Diplopterus with Luciocephalus, as Diplopterus was
preoccupied by Boie (Aves). Eschmeyer (1998) wrote that
no types were designated, although there is one specimen
(NMV 46359) from Bleeker’s collection in the National
Museum of Victoria. Attempts at finding the types or locality
of Gray’s specimen(s) have all failed (W. N. Eschmeyer, D.
Catania, M. Kottelat, A. Wheeler, and R. Pethiyagoda, pers.
comm.). The specimen or specimens on which Gray (1830)
based his figure is thus almost certainly no longer extant. As
Gray’s material came from many areas, it is not possible to
ascertain the precise locality from which the specimen in the
drawing is based (see Dawson (1946) for a history behind
Gray and Hardwicke’s illustrations). This uncertainty has not
been a problem as it has long been accepted that
Luciocephalus is a monospecific genus. There has also never
been any doubt as to the identity of the fish depicted by Gray,
although the plate actually does not depict the taxon accurately
as it shows an individual with two separate anal fins, instead
of an anal fin with a deep notch. This can easily be explained
as an error on the part of the artist or perhaps the original
specimen was damaged. The specimen depicted on Gray’s
figure (1830: Pl. 87) has a very uniform body coloration and
patterning, and while this does not really match what is now
known as L. pulcher, it has been assumed that this was merely
variation or artistic inaccuracy. The present discovery of a
second species of Luciocephalus, however, opens up a whole
new set of problems. It is quite possible that Gray’s figure
was actually based on a specimen from Sumatra, and that it
may even have been based on what is here named L. aura.
This is especially confusing considering the uniform body
patterning which is superficially closer to L. aura than L.
pulcher. However, in view of the fact that L. pulcher is the
type species of Luciocephalus Bleeker, 1851, and that the
current understanding of the name refers to the more
widespread species that occurs in Borneo, Malaysia and
Sumatra, there is a clear need to designate a neotype for L.
pulcher (Gray, 1830) that maintains this usage. This will
ensure there is nomenclatural stability, especially with the
present discovery of a second species which occurs
sympatrically with L. pulcher in Sumatra, and the fact that
future studies may even reveal the existence of additional
taxa. To this effect, we hereby designate a neotype for L.
pulcher (Gray, 1830). The specimen is a recently collected
specimen from Peninsular Malaysia and carries the catalogue
number ZRC 42520.
From recent collections in Sumatra, a separate and distinct
syntopic species was obtained. This taxon is described here
as a new species below.
Luciocephalus aura, new species
(Fig. 6)
Material examined. – Holotype - Jambi - MZB 9311 (holotype),
82.6 mm SL; Jambi: Pijoan area, coll. H. H. Tan & S. H. Tan, 23
Nov.1996.
Paratypes - Jambi - CMK 11054 (2 paratypes), 76.8-80.8 mm SL,
ZRC 41714 (1 paratype), 80.5 mm SL; Jambi: Sungai Pijoan,
downriver of confluence with stream draining Danau Pinang (ca.
half-hour upriver of Pijoan); M. Kottelat & H. H. Tan, 28 May.1994.
— CMK 11257 (3 paratypes), 86.8-105.1 mm SL, MZB 9312 (1
paratype), 86.8 mm SL, ZRC 41715 (3 paratypes), 74.2-98.4 mm
SL; Jambi: Danau Semangkat, a lake connected to Batang Hari by
Fig. 6. Luciocephalus aura - a. ZRC 46203 (paratype), 81.2 mm
SL, Jambi (Pijoan); b. freshly preserved specimen, CMK 11054
(paratype), 80.8 mm SL, Jambi (Pijoan); c. ZRC 47259, 111.4 and
102.2 mm SL, Jambi (Sungai Sentang).
a
b
c

130
Tan & Ng: Anabantoidei and Channoidei of Sumatra
Sungai Bangko opposite Kampung Senaning; M. Kottelat, 6
Jun.1994. — ZRC 41716 (3 paratypes), 78.5-89.4 mm SL; Jambi:
Pijoan area, coll. H. H. Tan et al., 7 Jun.1996. — ZRC 46203 (8
paratypes), 72.0-83.2 mm SL; Jambi: Pijoan area; don. Patrick Yap,
16 Mar.2001.
Others - Jambi - ZRC 47259, 2 ex., 102.2-111.4 mm SL; Jambi:
Sungai Sentang; don. Patrick Yap, Jun.2002.
Diagnosis. – Luciocephalus aura differs from L. pulcher in
having numerous iridescent green spots on central stripe when
live (vs. absence); absence of black irregular pattern on area
below preorbital to central stripe (vs. presence); area just
above anal-fin base without 3-4 dark brown semi-circles,
which appear circular when body is viewed ventrally (vs.
presence of dark markings); brownish caudal fin with 3-7
narrow iridescent vertical gold bars (vs. yellowish caudal fin
with 3-5 broad vertical black bars); absence of markings on
anal and pelvic fins (vs. presence); greater caudal-fin length
(total length 123.7-127.1% SL, vs. 120.0-125.6); greater
postdorsal length (12.0-13.2% SL, vs. 10.4-12.4); smaller
head length (43.3-45.9% SL, vs. 44.1-48.6); greater caudal
peduncle depth (10.5-11.2% SL, vs. 8.9-10.6); greater dorsal-
fin base length (10.1-11.0% SL, vs. 8.3-10.4); and greater
anal-fin base length (23.3-24.9% SL, vs. 19.3-23.4).
Description. – General body form as in Figs. 6a-c; meristics
and morphometrics listed in Table 3. Head long and slender,
almost half of standard length (head length 43.3-45.9% SL,
head width 9.8-10.7% SL); elongated snout with long
protrusible jaws (snout length 18.7-20.2% SL and 42.6-45.3%
HL, upper jaw length 34.9-38.2% HL, lower jaw length 46.1-
51.2% HL); body slender (body depth 15.4-17.7% SL); dorsal
fin elongate rounded, situated posteriorly, near caudal fin
(predorsal length 77.1-79.1% SL), dorsal-fin base less than
half of anal-fin base (43-44% anal-fin base length); caudal-
fin rounded; a deep notch on anal fin at proximal 6-8th anal
ray, about one third of anal-fin base (36.0-45.0% anal-fin base
length); pelvic fin with filamentous first ray, with extension
about twice or more as long as other pelvic rays; pectoral fin
rounded.
Live coloration. – Live and freshly captured coloration
illustrated in Figs. 6a, b. Body pattern consists of a dark brown
dorsum, whitish ventrum; distinct yellow narrow stripe on
dorsal of body when viewed dorsally; preorbital and
postorbital dark brown broad stripe joining with central dark
brown broad stripe, having numerous iridescent green spots,
area just above central stripe iridescent gold; opercle area
below preorbital and postorbital stripes without markings;
pectoral fin hyaline, area just below pectoral fin without
markings; pelvic fin with filamentous ray brownish, rest
hyaline; anal fin hyaline; dorsal fin with anterior 3-4 rays
proximally pigmented brown, rest hyaline; caudal fin
brownish with 3-7 narrow iridescent vertical gold bars, in a
regular pattern.
Preserved coloration. – Preserved coloration illustrated in
Fig. 6c. Body with brown dorsum, yellow narrow stripe on
dorsal of body when viewed dorsally lost due to preservation;
whitish ventrum with or without very faint brown stripe; broad
dark brown preorbital and postorbital stripe joining with
central stripe, ending at caudal peduncle with black spot;
middle of central stripe uniformly pigmented, green iridescent
spots lost due to preservation; area just above central stripe
cream with no pattern; area beneath preorbital to central stripe
cream with no pattern; lower jaw brown without black edge;
pectoral fin hyaline; pelvic fin with filamentous ray brownish,
rest hyaline; anal fin with very faint brown marks near base,
rest hyaline; dorsal fin with anterior 3-4 rays proximally
pigmented black, rest hyaline; caudal fin blackish with 3-7
narrow vertical cream bars, in a regular pattern.
Distribution. – Luciocephalus aura is currently only known
from the freshwater swamp forest habitats from the middle
reaches of the Batang Hari basin in Jambi, central Sumatra.
Field notes. – Luciocephalus aura is found only in freshwater
swamp forest habitats. Syntopic species include: Chela
maassi, Crossocheilus cobitis, Epalzeorhynchos kalopterus,
Osteochilus microcephalus, O. spilurus, Pectenocypris aff.
korthausae, Puntius gemellus, P. tetrazona, P. lineatus,
Rasbora brittani, R. dorsiocellata, R. dusonensis, R. gracilis,
R. pauciperforata, R. subtilis, Thynnichthys thynnoides
(Cyprinidae), Nemacheilus selangoricus (Balitoridae), Pangio
semicinctus (Cobitidae), Chaca bankanensis (Chacidae),
Doryichthys deokhatoides (Syngnathidae), Gymnochanda sp.
(Ambassidae), Pristolepis grooti (Pristolepididae), Betta falx,
B. raja, Parosphromenus sumatranus, Luciocephalus pulcher,
Sphaerichthys osphromenoides, Trichogaster trichopterus,
Trichopsis vittata (Osphronemidae), Channa micropeltes
(Channidae). Recently, L. aura has become available in
limited numbers in the aquarium fish trade (pers. observ.).
Etymology. – From the Latin aura, meaning glow, alluding
to the glowing iridescence of the green spots on its body. A
noun in apposition.
Remarks. – Luciocephalus aura can be differentiated from
L. pulcher in having slightly smaller preanal length (62.5-
63.8% SL, vs. 62.9-68.2); slightly greater postdorsal length
(21.0-24.1% body length, vs. 18.6-23.4); slightly smaller
caudal peduncle length (20.0-24.7% body length, vs. 21.9-
28.6); greater anal-fin base length (41.2-44.4% body length,
vs. 35.4-42.3); and smaller orbital diameter (13.3-14.1% HL,
vs. 13.7-17.0).
Luciocephalus aura is found syntopically with L. pulcher in
Jambi. There may be habitat differentiation for the two species
in Jambi, with L. pulcher seeming to prefer more stagnant
parts, whereas L. aura inhabits the more fast flowing sections
(M. Kottelat, pers. comm.). Riehl & Kokoscha (1993)
discussed the egg surface of a similar taxon from Kalimantan
Tengah, but without examining their specimens, we cannot
be sure if they are conspecific with the taxon from Sumatra.
Osphronemus goramy La Cepède *†
Osphromenus olfax - Günther, 1861: 382; Bleeker, 1879: 17;
Vinciguerra, 1879: 385; Volz, 1905: 126; 1907: 126; Weber &
de Beaufort, 1912: 540.

131
THE RAFFLES BULLETIN OF ZOOLOGY 2005
Osphronemus gourami - Regan, 1910: 774.
Osphronemus goramy - Weber & de Beaufort, 1922: 344; Roberts,
1992: 352.
Material examined. – Sumatra Utara - ZRC 42462 (2). Jambi -
ZMA 120.496 (5); ZRC 42364 (1); ZRC 43174 (1).
Remarks. – Roberts (1992) revised Osphronemus and
described two other species, with Sumatra having only one
recognised species - O. goramy. This is a commonly
cultivated food fish, with good tasting flesh.
Parosphromenus bintan Kottelat & Ng
Parosphromenus sp. – SH Tan & HH Tan, 1994: 357.
Parosphromenus bintan Kottelat & Ng, 1998: 269, figs. 4-6.
Material examined. – Holotype - Riau Archipelago - Pulau Bintan
- MZB (holotype), 23.6 mm SL.
Paratypes - Riau Archipelago - Pulau Bintan - ZRC 41403 (15
paratypes); CMK 11925 (6 paratypes); ZRC 30815-33 (19
paratypes).
Others - Riau Archipelago - Pulau Bintan - ZRC 31377-83 (7);
ZRC 30815-33 (19); ZRC 31327-31 (5); ZRC 31162-64 (3); Pulau
Lingga - ZRC (1).
Remarks. – Parosphromenus bintan was recently described
by Kottelat & Ng (1998), and can be found in Pulau Bintan,
Pulau Lingga and the island of Banka (pers. observ.). It can
be distinguished principally from P. deissneri in having a
rounded caudal fin and the median caudal rays branched (vs.
lanceolate caudal fin in male, and median caudal ray simple
in both sexes and filamentous in male); white pelvic fin
filament (vs. dark grey to black). Parosphromenus bintan can
be differentiated from P. sumatranus in missing the
conspicuous black spot at the posterior extremity of the dorsal-
fin base; and a white pelvic fin filament (vs. black) (Kottelat
& Ng, 1998).
Parosphromenus deissneri (Bleeker)
Osphronemus deissneri Bleeker, 1859: 376 (type locality:
Baturussak, Banka).
Polyacanthus deissneri - Günther, 1861: 381.
Parosphromenus deissneri - Bleeker, 1879: 20; Weber & de
Beaufort, 1922: 348; Kottelat et al., 1993: 164 (part); Kottelat
& Ng, 1998: 265, figs. 1-3.
Material examined. – Neotype - Banka - ZRC 31377, neotype,
20.2 mm SL.
Others - RMNH 6363 (1); ZRC 31017-28 (12); CMK 9613 (12);
ZRC 31378-83 (6); CMK 9651 (6). Biliton - ZRC 46184 (2).
Remarks. – Parosphromenus deissneri is found in the island
of Banka, but recently also found in Biliton. Males have
filamentous extension to the median caudal ray. All previous
records of this taxon from Malay Peninsula and Borneo refer
to other taxa. The holotype (RMNH 6363) is in very poor
condition, with missing caudal fin, translucent body and loss
of body pattern. Ng & Kottelat (1998) had applied to the
Commission to designate a neotype and had been successful
(ICZN, 2000). The neotype (ZRC 31377) is a freshly collected
male specimen from the island of Banka.
Parosphromenus deissneri can be differentiated from P.
bintan in having a lanceolate caudal fin in male and the
median caudal rays simple in both sexes, filamentous in male
(vs. rounded caudal fin in male, and median caudal ray
branched); and a dark grey to black pelvic fin filament (vs.
white). Parosphromenus deissneri is distinguished from P.
sumatranus in missing the conspicuous black spot at the
posterior extremity of the dorsal-fin base.
Parosphromenus sumatranus Klausewitz †
(Fig. 7)
Parosphromenus deissneri - Regan, 1910: 775 (part); Kottelat et
al., 1993: 164 (part).
Parosphromenus deissneri sumatranus - Klausewitz, 1955: 320.
Material examined. – Holotype - Jambi - SMF 3566 (holotype),
21.5 mm SL.
Paratypes – Jambi - SMF 3711 (4 paratypes).
Others - Jambi - ZRC 46208 (7); ZRC 42406 (50). Sumatra Selatan
- ZRC 43250 (6).
Remarks. – Parosphromenus sumatranus has been exported
for the aquarium trade identified as P. deissneri (T. Sim, pers.
comm.). It can be differentiated from P. bintan and P.
deissneri in having the conspicuous black spot at the posterior
extremity of the dorsal-fin base (vs. absence).
Parosphromenus sumatranus has black pelvic fin filament
(vs. white of P. bintan) (Kottelat & Ng, 1998).
Fig. 7. Parosphromenus sumatranus - a. not preserved male, ca. 20
mm SL, Jambi (Pijoan); b. SMF 3711 (paratype), 22.8 mm SL,
Jambi.
a
b

132
Tan & Ng: Anabantoidei and Channoidei of Sumatra
Sphaerichthys osphromenoides Canestrini †
Osphromenus malayanus - Weber & de Beaufort, 1912: 540.
Sphaerichthys osphromenoides - Weber & de Beaufort, 1922: 349;
Hardenberg, 1931: 125; 1933: 308; de Beaufort, 1939: 193;
Klausewitz, 1955: 322; Roberts, 1989: 176; Kottelat et al., 1993:
164.
Material examined. – Riau - ZRC 43114 (6); ZRC 39106 (4). Riau
Archipelago - ZRC 44051 (5). Jambi - SMF 3371 (46); SMF 3568
(15); ZRC 43126 (12); ZRC 43048 (3); ZRC 46207 (8); ZRC 46206
(38). Sumatra Selatan - ZRC 43249 (13). Banka - ZRC 31010
(2); ZRC 31160 (2).
Sphaerichthys selatanensis Vierke
(Fig. 8)
Material examined. – Biliton - ZRC 46183 (3).
Remarks. – The present series represents the first record of
a non-S. osphromenoides species out of Borneo. It resembles
S. selatenensis in having 4-5 cream or gold bars on body,
however, it differs in having a different body colour (reddish,
vs. brown). More material is required before its true identity
can be ascertained.
Trichogaster leerii (Bleeker) †
Osphromenus trichopterus - Volz, 1905: 126 (part); 1907: 126 (part).
Osphromenus trichopterus var. leerii - Günther, 1861: 384; Weber
& de Beaufort, 1912: 540.
Trichopodus leeri - Regan, 1910: 783, pl. 79, Fig. 2; Weber & de
Beaufort, 1922: 367.
Trichogaster leerii - Kottelat et al., 1993: 164.
Fig. 8. Sphaerichthys selatanensis – a. not preserved; b. ZRC 46183,
32.4 mm SL, Biliton.
a
b
Material examined. – Sumatra Utara - ZRC 42488 (3). Riau -
ZRC 39050 (6). Jambi - ZRC 39163 (2); ZRC 46209 (19). Sumatra
Selatan - ZRC 43248 (1).
Trichogaster pectoralis (Regan) *
Trichogaster pectoralis - Kottelat et al., 1993: 164.
Material examined. – Sumatra Utara - ZRC 42478 (1). Riau -
ZRC 39049 (2). Jambi - ZRC 46210 (1); ZRC 42323 (3).
Remarks. – Trichogaster pectoralis is most probably
introduced for aquaculture purposes.
Trichogaster trichopterus Pallas *
(Fig. 9)
Osphromenus trichopterus - Günther, 1861: 384; Volz, 1905: 126
(part); 1907: 126 (part); Weber & de Beaufort, 1912: 540.
Osphronemus trichopterus - Perugia, 1893: 242.
Trichopodus trichopterus - Bleeker, 1879: 21; Vinciguerra, 1879:
385; Regan, 1910: 783; Weber & de Beaufort, 1922: 366; Fowler
& Bean, 1927: 4; de Beaufort, 1939: 194.
Trichogaster trichopterus - Kottelat et al., 1993: 165; SH Tan &
HH Tan, 1994: 358.
Material examined. – Sumatra Utara - ZRC 42471 (1). Riau
Archipelago - ZRC 32600 (15). Jambi - ZRC 41739 (1); ZRC
46211 (4); ZRC 43258 (2).
Remarks. – Several captive strains of T. trichopterus have
been developed. Blue coloured strains are usually reported
in the aquarium literature as originating from Sumatra. In
the various samples we have have obtained across the whole
range of the species, we have never seen blue-coloured
individuals. The natural coloration of wild populations is as
follows (see Fig. 9): greyish body colour, darker at the dorsal,
almost white at ventral; head with dark grey dorsal and
silverish opercle scales with 2-3 blackish bars, eyes bright
red or orange when life; body with up to 20 blackish thin
bars, anterior bars complete, posterior 4-5 bars interrupted;
a large round black spot just after posterior edge of pectoral
fin on median of body, another black spot on middle of caudal
fin base; dorsal and caudal fins dark grey with rounded whitish
spots; dorsal fin distal margin yellowish; anal fin with anterior
part whitish and rest dark grey, broad orange distal margin,
Fig. 9. Trichogaster trichopterus - ZRC 43258, 57.8 mm SL, Jambi
(Bertam).

133
THE RAFFLES BULLETIN OF ZOOLOGY 2005
posterior of anal fin with a few rounded yellowish spots;
pectoral fin hyaline; pelvic fin elongated and exceeds anal
fin base length, whitish silver.
Trichopsis vittata (Cuvier & Valenciennes) †
Ctenops vittatus - Bleeker, 1879: 24; Regan, 1910: 776; Weber &
de Beaufort, 1922: 351.
Osphromenus vittatus - Weber & de Beaufort, 1912: 540.
Trichopsis vittata - Kottelat et al., 1993: 165.
Material examined. – Sumatra Utara - ZRC 42469 (60). Jambi
- ZRC 42584 (27).
Remarks. – This is a common and abundant species, most
often found in disturbed habitats (e.g. padi fields, ditches).
Specimens from Jambi have an iridescent green body colour.
SUB-ORDER CHANNOIDEI
FAMILY CHANNIDAE
The juvenile coloration pattern of all Channa species differ
markedly from its adult coloration (see Ng & Lim, 1990).
Typically, the juvenile coloration pattern consists of two
longitudinal black stripes on body, with alternating yellow
to bright red in between for C. bankanensis, C. lucius, C.
micropeltes and C. striata. For C. gachua, the young have
brownish transverse bars. For C. melasoma and C.
marulioides, the young is plain with orangish dorsal, with
ocellus on posterior part of dorsal fin and upper proximal
part of caudal fin respectively. For C. pleurophthalma, the
young have a reticulated pattern of four to six longitudinal
interrupted black stripes on body, with faint black patches
on lateral which would later develop into ocelli (Fig. 11b).
The juvenile pattern for C. cyanospilos is not known, but
probably similar to the young of C. melasoma due to their
close affinity. Ng & Lim (1990) and Courtenay & Williams
(2004) provide overviews of Channidae.
Channa bankanensis (Bleeker) *†
Ophiocephalus bankanensis Bleeker, 1852b: 726 (type locality:
Toboali, Banka); 1879: 51; Günther, 1861: 475; Weber & de
Beaufort, 1922: 323; de Beaufort, 1939: 192.
Channa bankanensis - Kottelat et al., 1993: 165; SH Tan & HH
Tan, 1994: 358.
Material examined. – Riau Archipelago - ZRC 32724 (2). Jambi
- ZRC 43125 (4); ZRC 41522 (4). Sumatra Selatan - ZRC 30530-
32 (3). Banka - ZRC 30848 (1); ZRC 30934-38 (5).
Remarks. – This is a stenotopic blackwater species. Channa
bankanensis can be differentiated from C. lucius, even at
juvenile stage, in having a rounder head vs. pointed; numerous
spots on unpaired fins vs. bars and stripes; reddish-brown
life coloration vs. brownish-black. For more details on
taxonomy and morphological changes during growth, see Ng
& Lim (1991) and Lee & Ng (1994) respectively.
Channa cyanospilos (Bleeker) *
Ophiocephalus cyanospilos Bleeker, 1853: 256 (type locality:
Telokbetong); Günther, 1861: 474.
Ophiocephalus cyanospilus - Bleeker, 1879: 45; Volz, 1905: 132;
1907: 132.
Ophiocephalus striatus - Weber & de Beaufort, 1922: 317 (part).
Channa cyanospilos - Ng & Lim, 1991: 120; Kottelat et al., 1993:
165.
Material examined. – Aceh - MZB 5611B (1); ZRC 14252 (1).
Riau — ZRC 39095 (2). Jambi - ZRC 41693 (5); MZB 9316 (1);
CMK 14416 (1).
Remarks. – This species is apparently quite rare, as it is not
encountered frequently. A detailed description of this species,
comparison with C. melasoma and C. striata is provided by
Ng & Lim (1991). The present series collected from the
market shows more variation. The spots on the throat can be
elongated and more blotchy. The spots on the lateral side of
body is present in some, but absent in others.
Channa gachua (Hamilton)
Ophiocephalus gachna – Volz, 1905: 131; 1907: 131.
Ophiocephalus gachua - Bleeker, 1879: 38; Perugia, 1893: 242;
Weber & de Beaufort, 1912: 539; 1915: 270; 1922: 321; de
Beaufort, 1939: 192.
Channa gachua - Kottelat et al., 1993: 165; SH Tan & HH Tan,
1994: 358.
Material examined. – Sumatra Utara - CMK 10804 (5); CMK
4514 (2); ZMA 121.681 (1). Riau Archipelago - ZRC 38314 (4).
Jambi - CMK 10792 (1); ZRC 42541 (3); ZRC 42542 (7). Sumatra
Selatan - NMBE 1020933 (1 - identified as C. mystax).
Remarks. – The specimens from Jambi possess numerous
tiny black spots on head and body, in contrary to Ng et al.
(1999: 60) who had stated that only specimens coming from
Indochina had black spots on the body. It appears that there
is a species complex and has a wide distribution range from
India to Bali (Kottelat et al., 1993; Ng et al., 1999; Kottelat,
2001).
Channa lucius (Cuvier & Valenciennes) *
Ophiocephalus bistriatus - Weber & de Beaufort, 1922: 322.
Ophiocephalus lucius - Günther, 1861: 475; Bleeker, 1879: 53; Volz,
1905: 133; 1907: 133; Weber & de Beaufort, 1912: 538; 1922:
326; de Beaufort, 1939: 193.
Ophiocephalus polylepis - Günther, 1861: 475; Bleeker, 1879: 50.
Ophiocephalus spiritalis Fowler, 1904: 530, Pl. 9.
Channa lucius - Kottelat et al., 1993: 165; SH Tan & HH Tan, 1994:
358.
Material examined. – Riau - ZRC 43111 (1). Riau Archipelago
- ZRC 37527 (1). Jambi - ZRC 43256 (1). Sumatra Selatan - ZRC
43252 (3). Banka - ZRC 30725 (1).
Remarks. – Channa lucius is sometimes confused with C.
bankanensis, but can be differentiated by its more flattened
and pointed head (vs. rounded), bars on fins (vs. spots) and
brownish black life colour (vs. reddish brown).

134
Tan & Ng: Anabantoidei and Channoidei of Sumatra
Channa marulioides (Bleeker) *
(Fig. 10)
Channa marulioides - Roberts, 1989: 170; Ng & Lim, 1990: 135.
Channa melanoptera - Roberts, 1989: 170.
Channa melanopterus - Ng & Lim, 1990: 135; Kottelat et al., 1993:
166.
Ophiocephalus marulius - Volz, 1905: 133; 1907: 133.
Ophiocephalus marulioides - Weber & de Beaufort, 1922: 315.
Ophiocephalus melanopterus - Bleeker, 1879: 41; Volz, 1905: 132;
1907: 132; Weber & de Beaufort, 1912: 539; 1922: 315.
Material examined. – Riau - ZRC 39029 (1). Jambi - ZRC 41508
(3).
Remarks. – Channa marulioides was described by Bleeker
(1851) from Sambas in Kalimantan Barat, based on a 270
mm TL specimen. This species was depicted (Bleeker, 1877-
78, Pl. 399, fig. 2) with an ocellus on the upper proximal part
of caudal fin. Bleeker (1855) described Channa melanoptera
based on a 601 mm TL specimen, from River Kapuas,
Pontianak in Kalimantan Barat. The specimen was depicted
(Bleeker, 1877, Pl. 399, fig. 2) as a yellowish brown fish with
striped pattern on dorsal, caudal and anal fins, with a
uniformly coloured body. Weber & de Beaufort (1922: 315,
fig. 84) depicted B. melanoptera as having irregular patches
of black on its lateral. We have come across and examined
specimens which have regularly spaced black patches of
lateral scales with white posterior edges (ZRC 41508). The
identity of C. marulioides is characterised by the presence of
the ocellus on the caudal fin, which is only distinct in young
individuals (ca. 50 mm SL). This ocellus later fades off and
the lateral black patches of scales becomes prominent. It
seems that C. melanoptera is either a junior synonym of C
marulioides (viz., Lee & Ng, 1994) or it is a species restricted
to the Sambas area in Kalimantan Barat. However, from
recent collections there, there is no species of Channa
resembling that of C. melanoptera as described by Bleeker
(unpublished data). Lee & Ng (1994) had earlier reported C.
melanoptera from Peninsular Malaysia and had arrived at the
earlier conclusion.
The present series obtained from central Sumatra had four or
five patches of white edged black scales on posterior three-
quarters of body (Fig. 10). The ocellus on the caudal fin is
obscured by dark pigmentation. It is not commonly
encountered in the market.
Channa melasoma (Bleeker, 1851) *
Ophiocephalus melanosoma - Günther, 1861: 473; Bleeker, 1879:
46; Weber & de Beaufort, 1912: 538; 1922: 319; de Beaufort,
1939: 192.
Channa melasoma - Kottelat et al., 1993: 166; SH Tan & HH Tan,
1994: 358.
Material examined. – Riau - ZRC 39094 (4). Riau Archipelago
- ZRC 37489-94 (6); ZRC 14012 (1). Sumatra Selatan - ZRC
43251 (1); NMBE 1020932 (1 – originally identified as C. mystax).
Remarks. – This species resembles C. cyanospilos but differs
in having a more pointed snout in lateral view (vs. blunt)
(see Ng & Lim, 1991, for more details).
Channa micropeltes (Cuvier & Valenciennes) *
Ophiocephalus micropeltes - Günther, 1861: 482; Bleeker, 1879:
55; Volz, 1905: 133; 1907: 133; Weber & de Beaufort, 1912:
538; 1922: 328; de Beaufort, 1939: 193.
Channa micropeltes - Roberts, 1989: 170; Kottelat et al., 1993:
166.
Material examined. – Riau - ZRC 39028 (2). Jambi - ZRC 38683
(1).—Sumatra Selatan - ZRC 30528-29 (2); NMBE 1020938 (1
- holotype of C. studeri).
Channa pleurophthalma (Bleeker) *†
(Fig. 11)
Channa pleurophthalmus - Roberts, 1989: 170; Kottelat et al.,
1993: 166.
Ophiocephalus pleurophthalmus - Günther, 1861: 479; Bleeker,
1879: 48; Volz, 1905: 133; 1907: 133; Weber & de Beaufort,
1922: 324.
Material examined. – Jambi - ZRC 41672 (20). Sumatra Selatan
- ZRC 43203 (7); ZRC 43253 (2).
Remarks. – The adult fish has a spectacular colour pattern
consisting of a metallic iridescent green base colour, with 3-
4 lateral yellow to orange ocelli with black centre on the body
Fig. 11. Channa pleurophthalma - a. ZRC 41672, 46.9 mm SL,
Jambi (Ayer Hitam); b. aquarium stock, ca. 250 mm SL, Jambi
(Ayer Hitam).
Fig. 10. Channa marulioides - ZRC 41508, ca. 420 mm SL, Jambi.
a
b

135
THE RAFFLES BULLETIN OF ZOOLOGY 2005
and an additional ocellus on both the opercle and caudal fin
base, and orange patches on the base of the anal and caudal
fins (Fig. 11b). The young fish has a reticulated pattern of 4-
6 longitudinal interrupted black stripes on body, with faint
black patches on lateral which would later develop into ocelli
(Fig. 11a).
Channa striata (Bloch) *
Ophiocephalus striatus - Bleeker, 1879: 42; Volz, 1905: 132; 1907:
132; Weber & de Beaufort, 1912: 538; 1922: 317 (part); Ahl,
1934: 238; Hardenberg, 1933: 308; de Beaufort, 1939: 192.
Ophicephalus striatus - Fowler & Bean, 1927: 4.
Channa striata - Kottelat et al., 1993: 166; SH Tan & HH Tan,
1994: 358.
Material examined. – Sumatra Utara - ZRC 42470 (25). Riau -
ZRC 43112 (2). Riau Archipelago - ZRC 32615-16 (2). Jambi -
ZRC 41694 (1). Sumatra Selatan - ZRC 43255 (1). Banka - ZRC
30866 (1).
Remarks. – See Ng & Lim (1991) for a comparison of C.
striata with C. cyanospilos and C. melasoma.
GENERAL DISCUSSION
The anabantoids and channoids of central Sumatra (consisting
of the Riau, Jambi and West Sumatra Provinces) includes
nearly all of the species previously recorded from the whole
of Sumatra (30 of 43 species = 70%) or from mainland
Sumatra (30 of 34 species = 88%); 21% (9 of 43 species) are
found exclusively in the smaller islands surrounding Sumatra.
Anabantoids and channids are a staple source of protein for
the local human population. They are found abundantly in
lakes, rivers and streams. The presence of the labyrinth and
accessory organs enables them to survive out of the water in
moist conditions for extended periods of time. This feature
facilitates their transport to far-off markets for live sale.
Channids have firm and odourless flesh which is much
favoured by locals (Ng & Lim, 1990). Trichogaster
trichopterus and T. pectoralis are often salted and dried,
commonly sold in the markets as salted fish. Osphronemus
goramy has a light creamy-textured flesh that is well
appreciated and is reflected by its high price in the market.
Large species of Betta are often collected and used as baits
for angling in Sumatra, and for Chitala lopis in the middle
Kapuas basin (M. Kottelat, pers. comm.).
Certain species of anabantoids and channids possess brightly
coloured and unusual head and body patterns, e.g. Betta
coccina, Parosphromenus spp., Sphaerichthys
osphromenoides, Trichogaster leerii, Channa gachua, C.
marulioides and C. pleurophthalma. This has made them
popular in the aquarium trade, and some species are even
bred commercially (e.g. Sphaerichthys osphromenoides and
Trichogaster trichopterus; see Ng & Tan, 1997).
All species of Osphronemidae exhibit paternal care for the
eggs and to a certain extent, to the fry. The genus Betta is
well noted for paternal care in oralbrooding and bubble nest
building species. There is also sibling brood care, as exhibited
by B. brownorum (from Sarawak) (Witte & Schmidt, 1992).
The genus Channa exhibit biparental care for eggs and fry,
sometimes causing grevious injury to unsuspecting humans
who wade into a school of fry, especially the largest species
- C. micropeltes (see Ng & Lim, 1990). Some taxa of
Channidae even exhibit oral-brooding, e.g. Channa gachua.
Species of Anabantidae and Helostomatidae are not known
to show any brood care.
ACKNOWLEDGEMENTS
We are grateful to Gunawan ‘Thomas’ and Verawaty Kasim,
for being such good hosts, providing logistic support and
donation of specimens; Maurice Kottelat, for his patience,
comments and information on Morton’s collections; Alwyn
Wheeler, Rohan Pethiyagoda (WHT), David Catania (CAS)
and W. N. Eschmeyer (CAS), for sharing information on
Gray’s type material; Argus Tjakrawidjaja, Renny Hadiaty,
Ike Rachmatika and Daisy Wowor (MZB), Giulianno Doria
(MSNG), Darrell J. Siebert (BMNH), Martien van Oijen
(RMNH), Isaäc Isbrücker (ZMA), Kelvin K. P. Lim (ZRC),
for access to specimens under their care; Oliver K. S. Chia,
for hand carrying specimens from Genova; Patrick Yap, for
providing fresh material; M. Kottelat, Tommy H. T. Tan, S.
H. Tan, Darren C. J. Yeo and H. H. Ng, for their help and
companionship in the field. This paper has been partially
supported by research grants RP 950326 and RP960314 to
PN from the National University of Singapore.
LITERATURE CITED
Ahl, E., 1934. Weitere Fische aus dem Toba-See in Sumatra.
Sitzungsberichte der Gesellschaft Naturforschende Freunde zu
Berlin, 1934: 235-237.
Alfred, E. R., 1966. The fresh-water fishes of Singapore. Zoologische
Verhandelingen, Leiden, 78: 1-68, 8 pls.
de Beaufort, L. F., 1939. On a collection of freshwater fishes of the
island of Billiton. Treubia, 17: 189-198.
Bleeker, P., 1850. Bijdrage tot de kennis der visschen met
doolhofvormige kieuwen van den Soenda-Molukschen Archipel.
Verhandenlingan Batavia Genootschap, 23: 1-15.
Bleeker, P., 1851. Nieuwe bijdrage tot de kennis der ichthyologische
fauna van Borneo, met beschrijving van eenige nieuwe soorten
van zoetwatervisschen. Natuurkundig Tijdschrift voor
Nederlandsch Indié, 1: 259-275.
Bleeker, P., 1852a. Bijdrage tot de kennis der ichthyologische fauna
van Blitong (Billiton), met beschrijving van eenige nieuwe
soorten van zoetwatervisschen. Natuurkundig Tijdschrift voor
Nederlandsch Indié, 3: 87-100.
Bleeker, P., 1852b. Nieuwe bijdrage tot de kennis der
Ichthyologische fauna van het eiland Banka. Natuurkundig
Tijdschrift voor Nederlandsch Indié, 3: 715-738.
Bleeker, P., 1853. Diagnostische beschrijvingen van nieuwe of
weinig bekende vischsoorten van Sumatra. Natuurkundig
Tijdschrift voor Nederlandsch Indié, 4: 243-302.

136
Tan & Ng: Anabantoidei and Channoidei of Sumatra
Bleeker, P., 1855. Nalezingen op de vischfauna van Sumatra.
Visschen van Lahat en Sibogha. Natuurkundig Tijdschrift voor
Nederlandsch Indiè 9, 257-280.
Bleeker, P., 1859. Negende bijdrage tot de kennis der vischfauna
van Banka. Natuurkundig Tijdschrift voor Nederlandsch Indié,
18: 359-378.
Bleeker, P., 1877-78. Atlas ichthyologique des Indes orientales
néêrlandaises. Tome IX. Percoïdes III. Muller, Amsterdam, 80
pp., pls. 355-420.
Bleeker, P., 1879. Mémoire sur les poissons à pharyngiens
labyrinthiformes de l’Inde archipélagique. Natuurkundig
Verhandenlingen Koninkl. Akademie, 19: 1-56.
Britz, R., 1994a. Ontogenic features of Luciocephalus (perciformes,
Anabantoidei) with a revised hypothesis of anabantoid
intrarelationships. Zoological Journal of the Linnean Society,
112: 491-508.
Britz, R., 1994b. Ablaichverhalten und Maulbrutpflege bei
Luciocephalus pulcher. Die Aquarien und Terrarien –
Zeitschrift, 47: 790-794 (with English summary).
Britz, R., 1995. Zur phylogenetischen Systematik der Anabantoidei
(Teleostei, Percomorpha) unter besonderer Berücksichtigung
der Stellung des Genus Luciocephalus. Morphologische und
ethologische Untersuchungen. PhD dissertation, Eberhard-
Karls-Universität Tübingen, Germany 125 pp.
Britz, R., 2001. The genus Betta – monophyly and
intrarelationships, with remarks on the subfamilies
Macropodinae and Luciocephalinae (Teleostei:
Osphronemidae). Ichthyological Exploration of Freshwaters, 12:
305-318.
Britz, R., M. Kokoscha & R. Riehl, 1995. The anabantoid genera
Ctenops, Luciocephalus, Parasphaerichthys, and Sphaerichthys
(Teleostei: Perciformes) as a monophyletic group: evidence
from egg surface structure and reproductive behaviour. Japan
Journal of Ichthyology, 42: 71-79.
Brown, A. & B. Brown, 1987. A survey of freshwater fishes of the
family Belontiidae in Sarawak. Sarawak Museum Journal, 37:
155-170.
Cantor, T. E., 1850. Catalogue of Malayan fishes. Journal of the
Royal Asiatic Society, Bengal, 18: i-xii + 981-1143, pls. 1-14.
Courtenay, W. R. Jr. & J. D. Williams, 2004. Snakeheads (Pisces,
Channidae) – A Biological Synopsis and Risk Assessment. U.S.
Department of the Interior, U.S. Geological Survey, Circular
1251, 143 pp.
Cuvier, G. & A. Valenciennes, 1846. Histoire naturelle des
poissons. Levrault, Paris, 18: xix + 505pp., pls. 520-553.
Dawson, W. R., 1946. On the history of Gray and Hardwicke’s
Illustrations of Indian Zoology, and some biographical notes
on General Hardwicke. Journal of the Society for Bibliophy of
Natural History, 2: 55-69.
Duncker, G., 1904. Die Fische de Malayischen Halbinsel.
Mitteilungen aus dem Naturhistorischen Museum, Hamburg,
21: 133-207.
Eschmeyer, W. N., 1998. Catalog of Fishes. California Academy
of Sciences, San Francisco, 2905 pp.
Fowler, H. W., 1904. A collection of fishes from Sumatra. Journal
of the Academy of Natural Sciences of Philadelphia, Ser. 2, 12:
497-559, Pls. 7-27.
Fowler, H. M., 1938. A list of the fishes known from Malaya.
Fisheries Bulletin of Singapore, 1: 1-268.
Fowler, H. W. & B. A. Bean, 1927. Notes on fishes obtained in
Sumatra, Java and Tahiti. Proceedings of the United States
National Museum, 71: 1-15.
Gray, J. E., 1830-1834. Illustrations of Indian zoology; chiefly
selected from the collection of Major-General Hardwicke.
Treuttel, Wurtz, Treuttel & Richter, London, 1: 100 pls.,
Richter, London & Parbury, Allen and Co., London, 2: 2 pp.,
102 pls.
Gray, J. E., 1831. Description of twelve new genera of fish,
discovered by General Hardwicke, in India, the greater part in
the British Museum. Zoological Miscellaneous 1831: 7-10.
Günther, A., 1861. Catalogue of the Acanthopterygian fishes in
the collection of the British Museum. British Museum, London,
3, 586 pp.
Hardenberg, J. D. F., 1931. The fishfauna of the Rokan mouth.
Treubia, 13: 81-168.
Hardenberg, J. D. F., 1933. Additional notes to my paper “The
fishfauna of the Rokan mouth”. Treubia, 14: 299-312.
Hardenberg, J. D. F., 1935-36. On a collection of fishes from the
estuary and the lower and middle course of the River Kapuas
(W. Borneo). Treubia, 15: 225-254.
Herre, A. W. C. T. & G. S. Myers, 1937. A contribution to the
ichthyology of the Malay Peninsula. Bulletin of the Raffles
Museum, Singapore, 13: 5-75.
ICZN (International Code for Zoological Nomenclature), 2000.
Opinion 1946. Osphronemus deissneri Bleeker, 1859 (currently
Parosphromenus deissneri; Osteichthyes, Perciformes):
holotype replaced by a neotype. Bulletin of Zoological
Nomenclature, 57: 60, March 2000.
Imaki, A., A. Kawamoto & A. Suzuki, 1981. Results of an
ichthyological survey in the Kapuas River, West Kalimantan,
Indonesia. Research Institute of Evolutionary Biology Science
Report, 1: 33-54.
Károli, J., 1881. Prodomus Piscium Asiae Orientalis, a domine
Joanne Xantus annia 1868-70 collectorum. Természetrajzi
Füzetek, 5: 147-187.
Klausewitz, W., 1955. See- und Süsswasserfische von Sumatra und
Java. Senckenbergiana Biologica, 36: 30-323.
Kottelat, M., 1989. Zoogeography of the fishes from Indochinese
inland waters with an annotated check-list. Bulletin Zoologisch
Museum Universiteit van Amsterdam, 12: 1-56.
Kottelat, M., 2001. Fishes of Laos. WHT Publications (Pte) Ltd,
Colombo, Sri Lanka, 198 pp.
Kottelat, M. & K. K. P. Lim, 1995. Freshwater fishes of Sarawak
and Brunei Darussalam: a preliminary annotated check-list.
Sarawak Museum Journal, 48 (69): 227-256.
Kottelat, M. & P. K. L. Ng, 1994. Diagnoses of five new species
of fighting fishes from Banka and Borneo (Teleostei:
Belontiidae). Ichthyological Exploration of Freshwaters, 5: 65-
78.
Kottelat, M. & P. K. L. Ng, 1998. Parosphromenus bintan, a new
osphronemid fish from Bintan and Bangka islands, Indonesia,
with redescription of P. deissneri. Ichthyological Exploration
of Freshwaters, 8: 263-272.
Kottelat, M., P. K. L. Ng & K. K. P. Lim, 1992. Recent collections
of freshwater fish from Terengganu, Peninsular Malaysia.
Malayan Naturalist, 46: 7-12.
Kottelat, M. & A. J. Whitten, 1996. Freshwater fishes of western
Indonesia and Sulawesi: additions and corrections. Periplus,
Hong Kong, 8 pp.

137
THE RAFFLES BULLETIN OF ZOOLOGY 2005
Kottelat, M., A. J. Whitten, S. N. Kartikasari & S. Wirjoatmodjo,
1993. Freshwater Fishes of Western Indonesia and Sulawesi.
Periplus Editions, Hong Kong, 259 pp., 84 pls.
Krummenacher, R., 1986. Betta waseri spec. nov. Aquarien und
Terrarien – Zeitschrift, 33: 177-181.
Kubota, K., M. Majima, K. Miki & K. Yamazaki, 1996.
Anabantoids. Tropical fish collection 5 (in Japanese). Pisces,
Tokyo, 144 pp.
Lauder, G. V. & K. F. Liem, 1981. Prey capture by Luciocephalus
pulcher: implications for models of jaw protrusion in teleost
fishes. Environmental Biology of Fish, 6: 257-268.
Lee, P. G. & P. K. L. Ng, 1994. The systematics and ecology of
snakeheads (Pisces: Channidae) in Peninsular Malaysia and
Singapore. Hydrobiologia, 285: 59-74.
Lim, K. K. P. & P. K. L. Ng, 1990. A guide to the freshwater fishes
of Singapore. Singapore Science Centre, 160 pp.
Lim, K. K. P., P. K. L. Ng & M. Kottelat, 1990. On a collection of
freshwater fishes from Endau-Rompin, Pahang-Johore,
Peninsular Malaysia. The Raffles Bulletin of Zoology, 38: 31-
54.
Mizuno, T. & J. I. Furtado, 1982. Ecological notes on fishes. In:
Furtado J. I. & S. Mori (eds.), The ecology of a freshwater
swamp, Tasek Bera. Dr. W. Junk Publisher, The Hague. pp. 321-
349.
Mohsin, A. K. M. & M. A. Ambak, 1983. Freshwater fishes of
Peninsular Malaysia. Penerbit Universiti Pertanian Malaysia,
284 pp.
Morton, W., 1908. Récit de voyage Ceylan et Sumatra - Novembre
1906-Juin 1907. Bulletin de la Société Vaudose des Sciences
Naturelles, (5) 44 (163): 143-204.
Ng, H. H., P. K. L. Ng & R. Britz, 1999. Channa harcourtbutleri
(Annandale, 1918): a valid species of snakehead (Perciformes:
Channidae) from Myanmar. Journal of South Asian Natural
History, 4: 57-63.
Ng, P. K. L. & M. Kottelat, 1994. Revision of the Betta waseri
group (Teleostei: Belontiidae). The Raffles Bulletin of Zoology,
Singapore, 42: 593-611.
Ng, P. K. L. & M. Kottelat, 1998. Case 3071. Osphronemus deissneri
Bleeker, 1859 (currently Parosphromenus deissneri;
Osteichthyes, Perciformes): proposed replacement of holotype
by a neotype. Bulletin of Zoological Nomenclature, 55 (3): 155-
158.
Ng, P. K. L. & K. K. P. Lim, 1990. Snakeheads (Pisces: Channidae):
natural history, biology and economic importance. In: Essays
in Zoology. Papers commemorating the 40th anniversary of the
Department of Zoology, National University of Singapore: 127-
152.
Ng, P. K. L. & K. K. P. Lim, 1991. The identity of Channa
cyanospilos Bleeker from Sumatra, and a new record of Channa
bankanensis (Bleeker) from Peninsular Malaysia (Pisces:
Channidae). The Raffles Bulletin of Zoology, Singapore, 39: 119-
130.
Ng, P. K. L. & H. H. Tan, 1997. Freshwater fishes of Southeast
Asia: potential for the aquarium trade and conservation issues.
Aquarium Science Conservation, 1: 79-90.
Ng, P. K. L., J. B. Tay & K. K. P. Lim, 1994. Diversity and
conservation of blackwater fishes in Peninsular Malaysia,
particularly in the North Selangor peat swamp forest.
Hydrobiologia, 285: 203-218.
Ng, P. K. L., J. B. Tay, K. K. P. Lim & C. M. Yang, 1992. The
conservation of fish and other aquatic fauna of the north
Selangor peat swamp forest and adjacent areas. Asian Wetland
Bureau, Natn. Univ. Singapore, World Wide Fund for Nature-
Malaysia & Univ. Malaya; 90 pp., 110 figs.
Perugia, A., 1893. Di alcuni pesci raccolti in Sumatra dal Dott. Elio
Modigliani. Annali del Museo Civico di Storia Naturale
Giacoma Doria, Genova, 13: 241-247.
Regan, C. T., 1910. The Asiatic fishes of the family Anabantidae.
Proceedings of the Zoological Society London, 1909[1910]:767-
787, pls. 77-79.
Riehl, R. & M. Kokoscha, 1993. A unique surface pattern and
micropylar apparatus in the eggs of Luciocephalus sp.
(Perciformes, Luciocephalidae). Journal of Fish Biology, 43:
617-620.
Roberts, T. R., 1989. The freshwater fishes of western Borneo
(Kalimantan Barat, Indonesia). Memoirs of the California
Academy of Sciences, San Francisco, 14, 210 pp.
Roberts, T. R., 1992. Systematic revision of the Southeast Asian
anabantoid fish genus Osphronemus, with descriptions of two
new species. Ichthyological Exploration of Freshwaters, 2: 351-
360.
Sauvage, H.-E., 1884. Note sur une collection de poissons recueillie
à Pérak, presquîle de Malacca. Bulletin de la Société Zoologique
de France, 9: 216-220.
Schaller, D. & M. Kottelat, 1990. Betta strohi sp. n., ein neuer
Kampffisch aus Südborneo (Osteichthyes: Belontiidae). Die
Aquarien- und Terrarien-Zeitschrift, 43: 31-37.
Sim, C. H., 2002. A field guide to the fish of Tasek Bera Ramsar
site, Pahang, Malaysia. Wetlands International-Malaysia
Programme, Malaysia, 99 p.
Steindachner, F., 1901. Fische. [In: W. Kükenthal, Ergebnisse einer
zoologischen Forschungsreise in den Molukken and Borneo].
Abhandlungen der Senckenbergischen Naturforschenden
Gesellschaft, 25(2): 408-464, pls. 17-18.
Tan, H. H., 1998. Two new species of the Betta waseri group
(Teleostei: Osphronemidae) from central Sumatra and southern
Thailand. Ichthyological Exploration of Freshwaters, 8: 281-
287.
Tan, H. H. & M. Kottelat, 1998a. Two new species of Betta
(Teleostei: Osphronemidae) from the Kapuas basin, Kalimantan
Barat, Borneo. The Raffles Bulletin of Zoology, Singapore, 46:
41-51.
Tan, H. H. & M. Kottelat, 1998b. Redescription of Betta picta
(Teleostei: Osphronemidae) and description of B. falx n. sp. from
central Sumatra. Revue Suisse de Zoologie, 105: 557-568,
September 1998.
Tan, H. H. & K. K. P. Lim, 2004. Inland fishes from the Anambas
and Natuna Islands, South China Sea, with description of a new
species of Betta (Teleostei: Osphronemidae). The Raffles
Bulletin of Zoology, Singapore, Supplement no. 11: 107-115.
Tan, H. H. & P. K. L. Ng, 1996. Redescription of Betta bellica
Sauvage, 1884 (Teleostei: Belontiidae), with description of a
new allied species from Sumatra. The Raffles Bulletin of Zoology,
Singapore, 44: 143-155.
Tan, H. H. & P. K. L. Ng, 2005. The fighting fishes (Teleostei:
Osphronemidae: genus Betta) of Singapore, Malaysia and
Brunei. In: Kottelat, M. & D. C. J. Yeo (eds.), Southeast Asian
Freshwater Fish Diversity. The Raffles Bulletin of Zoology,
Singapore, Supplement no. 13: 43-99.
Tan, H. H. & S. H. Tan, 1994. Betta miniopinna, a new species of
fighting fish from Pulau Bintan, Riau Archipelago, Indonesia
(Teleostei: Belontiidae). Ichthyological Exploration of
Freshwaters, 5: 41-44.

138
Tan & Ng: Anabantoidei and Channoidei of Sumatra
Tan H. H. & S. H. Tan, 1996. The identity of Betta pugnax
(Teleostei: Belontiidae), with the description of a new species
of Betta from the Malay Peninsula. The Raffles Bulletin of
Zoology, Singapore, 44: 419-434.
Tan, S. H. & H. H. Tan, 1994. The freshwater fishes of Pulau Bintan,
Riau Archipelago, Sumatera, Indonesia. Tropical Biodiversity,
2: 351-367.
Tweedie, M. W. F., 1952. Notes on Malayan fresh-water fishes.
Bulletin of the Raffles Museum, Singapore, 24: 63-95.
Tweedie, M. W. F., 1953. Malayan aquarium fishes. Malayan Nature
Journal, 8: 47-51.
Vaillant, L., 1902. Résultats zoologiques de l’expédition scientifique
Néerlandaise au Bornéo central. Poissons. Notes Leyden
Museum, 24: 1-166.
Vidthayanon, C., 2002. Peat swamp fishes of Thailand. Office of
Environmental Policy and Planning, Bangkok, Thailand, 136
pp.
Vidthayanon, C., J. Karnasuta & J. Nabhitabhata, 1997. Diversity
of Freshwater Fishes in Thailand. Office of Environmental
Policy and Planning, Bangkok, Thailand, 102 p.
Vinciguerra, D., 1879, Appunti ittiologici sulle collezioni del Museo
Civico de Genova. I. Enumerazione di alcune specie di pesci
raccolti in Sumatra dal Dr. O. Beccari nell’ anno 1878, Annali
del Museo Civico di Storia Naturale Giacoma Doria, Genova,
14: 384-397.
Vinciguerra, D., 1926. Catalogo dei pesci raccolti a Borneo dai sigg.
Marchese G. Doria e Dott. O. Beccari negli anni 1865-67. Annali
del Museo Civico di Storia Naturale Giacoma Doria, Genova,
serie 3, 10: 532-628, pl. 1.
Vierke, J., 1979. Betta coccina nov. spec., ein neuer Kampffisch
von Sumatra. Das Aquarium, 13 (121): 288-289.
Volz, W., 1904. Fische von Sumatra gesammelt von Herrn G.
Schneider. Revue Suisse de Zoologie, 12: 451-493.
Volz, W., 1905. Catalogue of the fishes of Sumatra. Natuurkundig
Tijdschrift voor Nederlandsch Indié, 9: 35-250.
Volz, W., 1907. The fishes of Sumatra. Natuurkundig Tijdschrift
voor Nederlandsch Indié, 10: 35-250.
Weber, M. & L. F. de Beaufort, 1912. Die Fische. In: Mass, A.
(ed.), Durch Zentral-Sumatra, Vol. 2. Behr, Berlin & Leipzig.
Pp. 522-541, pls. 11-12.
Weber, M. & L. F. de Beaufort, 1915. Fische aus den Süsswasser
von Nias, in: Zoologische Resultate aus J. P. Kleiweg de Zwaan,
Die Insel Nias bei Sumatra. Nijhoff, Haag, 1915.
Weber, M & L. F. de Beaufort, 1922. The fishes of the Indo-
Australian archipelago. 4. Brill, Leiden, xiii + 410 pp.
Whitten, A. J., S. J. Damanik, J. Anwar & N. Hisyam, 1987. The
Ecology of Sumatra. Gadjah Mada University Press, i-xvii, 583
pp.
Witte, K.-E. & J. Schmidt, 1992. Betta brownorum, a new species
of anabantoids (Teleostei: Belontiidae) from northwestern
Borneo, with a key to the genus. Ichthyological Exploration of
Freshwaters, 2: 305-330.