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 themost 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) anda 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 whencompared to Borneo or Malaysia. The definition of Sumatrain 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 Natunaislands, please refer to Tan & Lim (2004). With regards tothe air breathing labyrinth fish fauna (Anabantoidei andChannoidei), recent collections in central Sumatra, Banka andPulau Bintan (Riau Archipelago) from 1993 to 1997 haveresulted in the discovery of eight new species of Betta anda new species of Parosphromenus (Osphronemidae) from thisarea, 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; andParosphromenus bintan Kottelat & Ng, 1998.
The intra-relationships of the suborder Anabantoidei wererecently revised (Britz, 1994a, 1995, 2001; Britz et al., 1995)
and several families have been shown to be monophyletic,
and Belontiidae and Luciocephalidae are synonymised withOsphronemidae (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 designatedand 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); the115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); andthe Zoological Reference Collection of the Raffles Museumof Biodiversity Research, National University of Singapore(ZRC). Material examined are listed by Province (from northto south) and followed by the collection abbreviation andcatalogue number with the number of specimens in brackets.
Full locality data is quoted only for new species andredescriptions 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 speciesof anabantoids and channoids of Sumatra and their localities
by province and island is provided in Table 1, based onliterature and collections. Material examined is listed by
locality, catalogue number and the number of specimens inbrackets; more details are provided for new taxa andredescriptions. Abbreviations used are SL: standard length,
TL: total length, HL: head length.

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.

TAXONOMY
SUB-ORDER ANABANTOIDEI

Material examined. – Sumatra Utara - BMNH 1889.12.26:30(lectotype of B. fasciata); ZRC 42468 (10); ZRC 42478 (4). Riau
Archipelago - ZRC 44057 (14).

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 & deBeaufort, 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).

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 (23.638.8% 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 lesssloping lateral head profile (vs. more sloping with a slightconcavity (see Tan & Ng, 1996).

FAMILY HELOSTOMATIDAE
Helostoma temminckii Cuvier *Betta simorum Tan & Ng †Helostoma temmincki - Bleeker, 1879: 15; Volz, 1905: 125; 1907:
125; Weber & de Beaufort, 1912: 540; 1922: 340; Hardenberg,
1933: 308.

Betta bellica (non Sauvage, 1884) - Kottelat et al., 1993: 161 (part),pl. 75.
Betta simorum Tan & Ng, 1996: 151, figs. 3C, 4C.

116

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. rajaB. renataB. rubraB. 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*reflects extensive collection done

+
+

+
+

+

+
+

117

+

+
17

+

+
+

12

+
+
+

+

+

+
+
+

+

+

+

+

+

+

+

+

Sumatra Utara

+

Aceh

3

+

+

+

Nias

+
7

+
+

+

+

+

+

Sumatra Barat

+
20

+
+
+
+

+
+

+
+

+

+
+

+

+
+

+

+

+

+

+

Riau

+
19*

+
+
+
+

+

+
+

+

+

+

+

+
+

+

+

+

+

+

Riau Archipel.

+
+
+
26*

+
+
+
+
+

+
+
+

+
+

+
+
+

+

+
+
+

+

+
+

+

+

+

Jambi

+
+
+
+
+
+
+
20

+

+
+
+

+

+
+

+
+

+

+

+

+

Sumatra Selatan

+
18*

+
+
+
+
+

+

+

+

+
+

+

+

+

+
+

+

+

Banka

Table 1. Distribution of Anabantoids (Anabantidae, Helostomatidae, Osphronemidae) and Channoids (Channidae) of Sumatra (based on literature and collections)

+
16

+
+
+
+
+

+

+

+
+

+

+

+

+

+

+

Biliton

+
6

+

+

+

+

Bengkulu

5

+
+

+
+

+

+

Lampung

THE RAFFLES BULLETIN OF ZOOLOGY 2005

Tan & Ng: Anabantoidei and Channoidei of Sumatra
Material examined. – Holotype - Jambi - MZB (holotype), 63.8mm SL.

Remarks. – Betta simorum is restricted to the southern halfof Sumatra and was recently recorded from the Kapuas basinin Kalimantan Barat (pers. observ.). It is closely allied to B.
bellica (see Tan & Ng, 1996).

dorsal fin rays (9-12, vs. 14-15); fewer subdorsal scales (78, 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 baselength (15.2-20.4% SL, vs. 26.8-28.6). Betta coccina differsfrom B. miniopinna in the following characters: presence oflateral iridescent blotch in males (vs. absence); red bodycolour (vs. black); black pelvic fin tip (vs. white); presenceof 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 coccina species group

Betta miniopinna Tan & Tan

Betta burdigala Kottelat & Ng

Betta miniopinna HH Tan & SH Tan, 1994: 42, fig. 1; SH Tan &HH Tan, 1994: 357.

Paratypes - Jambi - ZRC 38549 (4 paratypes); CMK 11182 (5paratypes); 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).

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 (2paratypes).

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 (2426, 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 iridescentgreen 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 (26.828.6% SL, vs. 16.0-24.6); greater postdorsal length (16.717.4% SL, vs. 12.3-13.3); smaller predorsal length (52.854.9% SL, vs. 56.2-65.4); and a greater preanal length (42.545.5% SL, vs. 37.9-42.6).

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 coccina Vierke †Betta coccina Vierke, 1979: 288 (type locality: Jambi); Kottelat etal., 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 groupand is also found in Peninsular Malaysia. It is a stenotopic
blackwater species. It differs from B. burdigala in thefollowing characters: presence of lateral iridescent blotch inmales (vs. absence); black pelvic fin tip (vs. white); fewer

118

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 3515961 (3). Biliton - ZRC 46182 (3).

Remarks. – Betta edithae belongs to its own species group.
It has a widespread distribution and is found in Kalimantan

THE RAFFLES BULLETIN OF ZOOLOGY 2005
Timur, Kalimantan Selatan, Kalimantan Tengah, Kalimantan
Barat, Sumatra Selatan, Banka, Biliton and Pulau Bintan (ofthe 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: chinbar 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 (48.251.8% 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.

pelvic fin rounded with short filamentous ray (28.1-38.4%
SL); pectoral fin rounded.

Betta rubra Perugia(Fig. 1)

Preserved coloration. – See Figs. 1a, b for generalappearance. Dorsum of body dark brown, ventrum lighter
brown. Dorsum of head brown with black spots. Mouth withlower lip black. Distinct pre- and postorbital dark brown
stripes on head, postorbital stripe interrupted by verticalparallel reddish bars on opercle, opercle with distal darkbrown margin. Broad dark brown chin-bar present, whichmerges with anterior part of second postorbital stripe, forminga triangular dark brown mark below eye. Body withirregularly 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 finswithout transverse bars on interradial membrane. Pectoral fin
rounded, and hyaline. Pelvic fin rounded, with reddish
filamentous second ray.

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).

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.

Betta foerschi species group

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 othermembers 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 partof second postorbital stripe, forming a triangular dark markbelow 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 thanB. foerschi (28.1-38.4% SL, vs. 21.4-28.5); smaller dorsalfin 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 inTable 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); analfin base length about half of standard length (48.3-55.9% SL);

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 freshcollections (viz., Tan & Ng, 2005) (present volume). The
specimens from Genova (MSNG 13019) were in goodcondition, but the opercle and pelvic, anal, caudal and dorsal
fins were tinged pinkish-red. On closer examination under amicroscope, 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.

abFig. 1. Betta rubra - a. MSNG 13019 (lectotype), 29.4 mm SL,
Siboga; b. ZRC 42497, 33.4 mm SL, Aceh Barat (right side,
reversed).

119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 (6paratypes); 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 (5paratypes).
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 thefollowing 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 (46.550.3% SL, vs. 42.6-48.4); in life - male with distal margins
of anal and caudal fins reddish (vs. bluish); iridescentgreenish-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 orabsent); 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).

other members of the B. pugnax group in having the following
unique combination of characters: most slender body in thespecies group (body depth at dorsal-fin origin 21.2-24.2%
SL); anal fin rays 27-29 (mode 28); lateral scales 32-33 (mode33); 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 andblunt (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 65.067.1% SL); caudal fin with distal part of rays extended slightly
beyond interradial membrane, median rays elongated; analfin 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. Dorsumof body brown, ventrum lighter brown. Dorsum of headbrown, 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 andjuvenile. Male with some body scales on dorsal half withblue iridescence, rest of body scales with faint dark brown
posterior edge, more distinct on belly scales. Body with 57 faint irregularly spaced vertical brown bars. Female andjuvenile with indistinct central and second central stripes, andindistinct caudal peduncle spot. Fins brownish, dorsal andcaudal fins with dark brown dorsal transverse bars in male,
only dorsal fin with transverse bars in female. Both male andfemale with thin dark blue edge on anal fin. Pectoral finhyaline. Pelvic with bright whitish distal tip.
Preserved coloration. – See Fig. 2b for general appearance.
Dorsum of body dark brown, ventrum lighter brown towhitish. Dorsum of head dark brown, male with distal partof opercle after eye black. Distinct pre- and postorbital blacka

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 aftermain 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 the120bFig. 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).

THE RAFFLES BULLETIN OF ZOOLOGY 2005
Table 2. Meristics and morphometrics of Betta cracens, B. fusca, B. raja and B. rubra.

Sample sizeStandard length (mm)
MERISTICS
Vertebrae

Anal fin rays
Dorsal fin rays
Caudal fin rays
Pelvic fin rays
Pectoral fin rays
Anal fin rays below dorsal-fin origin
Anal fin rays at pectoral fin end
Anal fin rays at pelvic fin end
Subdorsal scales
Transverse scales at dorsal-fin origin
Lateral scales
Lateral scales below dorsal-fin origin
Lateral scales above anal-fin origin
Predorsal scales
Postdorsal scales
MORPHOMETRICS - % SL
Total length
Predorsal length
Postdorsal length
Caudal peduncle depth
Preanal length
Head length
Body depth at dorsal-fin origin
Pelvic fin length
Anal-fin base length
Dorsal-fin base length
Body length
Head width% HL
Orbit diameter
Postorbital lengthInterorbital width
Snout length
Head width% Postorbital length
Orbit diameter
Interorbital width% Predorsal length
Head length

Betta cracens

Betta fusca

Betta raja

Betta rubra

638.2-57.4

2
53.1-59.7

20
45.9-63.9

4
33.1-35.0

2-3 + 8 + 20-21(total 31-32, 31)(n = 11)
I-II, 26-27(total 27-29, 28)
0-I, 8-9(total 8-9, 8 or 9)ii,5+6,ii
I, 1, 413-

2-3 + 8 + 19
(total 29-30, 29 or 30)(n = 2)0-I, 23-25(total 24-25, 24 or 25)
0-II, 7-8
(total 8-9, 8 or 9)ii,6+6,ii
I, 1, 4
12
9
5-6
11-12

2 + 8 + 18-19(total 28-29, 29)(n = 4)
II-III, 24-26(total 27-29, 27)0-II, 7-8
(total 8-9, 9)ii-iii,5+6,i-iii
I, 1, 413-

51/2-6 (51/2 or 6)
91/2-10 (91/2)
32-33 (33)
16-19 (17 or 18)
6-7 (7)23-24 (23 or 24)
11-12 (11 or 12)

51/2-6
10
29
16
6
22-23
11

2-3 + 8 + 18-19(total 28-30, 29)(n = 20)
II, 23-26(total 25-28, 26)
I-II, 7-10
(total 8-11, 9)
ii,5+6-7,i (ii,5+7,i)
I, 1, 412-13 (13)10-13 (11)
6-9 (8)
11-17, females
15 and above, males
6-7 (61/2)
10-101/2 (10)30-32 (31)14-16 (15)
5-7 (6)20-22 (22)10-11 (11)138.9-145.8
65.0-67.1
20.4-24.6
15.1-16.7
42.0-46.1
30.8-31.7
21.2-24.2
36.1-43.4
53.4-55.7
10.5-11.6
68.5-71.3
17.1-19.9

138.6
68.5-70.2
22.2-23.5
19.2-20.4
49.0-50.3
33.2-33.8
30.9-31.7
38.6-39.2
47.8-51.4
13.7-13.9
70.4-70.8
19.8-23.6

138.5-159.4
63.2-69.9
20.0-24.6
18.3-23.2
46.1-52.4
33.9-37.4
28.1-33.5
37.4-69.8
48.9-54.8
11.0-15.3
62.8-70.7
18.5-22.3

131.6-142.3
62.9-67.6
20.3-23.3
13.1-15.3
41.7-47.4
27.3-33.6
23.2-25.8
28.1-38.4
49.2-52.8
10.0-12.2-

24.7-28.2
46.3-50.7
32.7-37.1
24.0-25.8
53.8-64.0

23.8-26.7
48.3-50.5
33.5-34.2
25.0-26.7
59.7-69.8

24.0-28.7
45.1-53.9
32.2-39.6
26.3-29.8
51.8-62.0

24.5-25.8
49.5-57.1
22.7-29.5
17.9-24.7
-

50-58
67-75

47-55
68-70

46-64
67-78

-

46.3-48.4

47.2-49.4

50.4-55.7-

stripes on head, chin-bar present but may be obscured byblackened opercle scales. Body scales with black posterioredge, 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 withtransverse black bars in female, anal fin with thin distal blackedge. Pectoral fin hyaline. Pelvic fin dark brownish, withhyaline filamentous second ray.
Distribution. – Betta cracens is currently known only from

5-6 (6)
81/2-91/2 (91/2)
281/2-30 (29)
15-16
6-8 (6)22-24 (24)10-11the 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 inlarge 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, Sphaerichthysosphromenoides,
Trichogastertrichopterus(Osphronemidae), Channa gachua and C. lucius (Channidae).

121

Tan & Ng: Anabantoidei and Channoidei of Sumatra
Etymology. – From the Latin cracens meaning slender,
graceful; alluding to the slender body, as compared with theother members of the B. pugnax group.
Remarks. – Betta cracens can be differentiated from the othermembers 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 countthan 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 (mode23-24, vs. 17-22); more postdorsal scales than B. prima (1112, vs. 9-10); smaller total length than B. enisae (138.9145.8% SL, vs. 143.0-155.2); smaller predorsal length thanB. fusca, B. prima and B. enisae (65.0-67.1% SL, vs. 67.370.3); smaller caudal peduncle depth than B. fusca and B.
raja (15.1-16.7% SL, vs. 18.3-23.2); smaller preanal lengththan all members except B. pugnax and B. pulchra (42.046.1% 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 widththan 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 (5058% postorbital length, vs. 57-76); and a smaller head lengththan 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 (2324, 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 baselength(53.4-55.7% SL, vs. 46.8-46.9); smaller dorsal-fin baselength (10.5-11.6% SL, vs. 14.4-14.7); and a greater bodylength (68.5-71.3% SL, vs. 65.3-67.2).

Paralectotype - Sumatra Utara - BMNH 1908.17.13:19 (1paralectotype), 53.1 mm SL; sama data as lectotype.

Diagnosis. – Betta fusca can be differentiated from the othermembers of the B. pugnax group by the following set ofcharacters: 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), headlong 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- andpostorbital 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 secondray.
Distribution. – Betta fusca is currently known only from thetype 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: 165174), 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 Muséed’Histoire Naturelle, Lausanne, Switzerland; from Morton’s
text, it seems that some material was also deposited in
Naturhistorisches Museum, Zürich, Switzerland, but noMorton fish material could be located in these two collections

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.

122

Fig. 3. Betta fusca - BMNH 1908.17.13:18 (lectotype), 59.7 mm
SL, Medan (caudal fin missing).

THE RAFFLES BULLETIN OF ZOOLOGY 2005
(M. Kottelat, pers. comm.). Attempts had been made to collectfresh material from Medan area, but we were not successful.
Remarks. – Betta fusca can be differentiated from the othermembers 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 thanB. schalleri (51/2-6, vs. 61/2-7); fewer subdorsal scales than B.
pulchra and B. breviobesus (mode 51/2-6, vs. 7); fewer lateralscales than B. schalleri and B. cracens (29, vs. 31-33); morelateral 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 thanB. 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 thanB. cracens (49.0-50.3% SL, vs. 42.0-46.1); smaller headlength 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 headlength than B. cracens (33.2-33.8% SL, vs. 30.8-31.7);
smaller body depth at dorsal-fin origin than B. schalleri andB. 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 lengththan 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 (23.826.7% HL, vs. 26.0-38.5) and smaller orbital diameter thanB. 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 widththan 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.

56.3 mm SL male; Sungai Ayer Merah, feeder stream to Danau
Souak Padang ca. 15 mins. by boat upstream (1°36’45.7”S103°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), 27.857.6 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), 35.1-63.9mm SL, ZMA 121.823 (5 paratypes), 36.3-50.7 mm SL, BMNH (5paratypes), 37.5-50.7 mm SL; Sungai Sentang, ca. 20 mins. walkthrough 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), 29.3-54.2mm 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), 36.063.8 mm SL, CMK 11159 (7); Danau Rasau, a blackwater lakedraining to Batang Hari, opposite Kampung Rantau Panjang; M.

aThe larger specimen (BMNH 1908.17.13:18, 59.7 mm SL)
has been chosen as the lectotype although its caudal fin isbroken 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 andmay lack several anal fin rays.

b

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),

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).

123

Tan & Ng: Anabantoidei and Channoidei of Sumatra
Kottelat & H. H. Tan, 1-2 Jun.1994. — ZRC 42504 (16), 54.3-65.6mm 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 othermembers of the B. pugnax group by the following set ofcharacters: mature males with very long pelvic fin (15-23
anal fin rays at pelvic fin end, even exceeding anal-fin basein some specimens); dark margins on anal fin and lower halfof 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; bodylength 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), headlong 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. Dorsumof body brown, ventrum lighter brown to yellow. Dorsum ofhead brown, male with iridescent green opercle, female withgold opercle. Distinct pre- and postorbital black stripes onhead, 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 posterioredge. Body with faint irregularly spaced 3-5 vertical brownbars. Female and juvenile with more distinct central andsecond 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 to124whitish. Dorsum of head dark brown, male with distal partof 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 withirregularly 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 withtransverse bars, caudal transverse bars absent. Pectoral finhyaline. Pelvic fin brownish, with whitish filamentous secondray.
Distribution. – Betta raja is found in the lowland swamp
forests of the Batang Hari basin in Jambi, the northern partof Sumatra Selatan, in the Indragiri basin in Riau and easternpart of Sumatra Barat in Sijunjung.
Field notes. – In Danau Rasau, a blackwater lake draining
to Batang Hari, fisherfolk catch them with hook-and-line anduse 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,
Sphaerichthysosphromenoides, 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 kingor prince, also from the Indonesian vernacular for this species- Chupang Raja. A noun in apposition.
Remarks. – Betta raja can be differentiated from the othermembers 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 somespecimens); more anal fin rays than B. fusca (25-28, vs. 2425); 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 lengththan 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. 15.116.7); greater preanal length than B. cracens (42.0-46.1% SL,
vs. 46.1-52.4); greater pelvic fin length than B. prima (37.469.8% SL, vs. 30.1-37.2); greater dorsal-fin base length thanTHE RAFFLES BULLETIN OF ZOOLOGY 2005
B. enisae (37.4-69.8% SL, vs. 8.7-12.1); slightly smaller headwidth 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, 6176).

(26.7-27.6% SL, vs. 21.2-24.2); greater dorsal fin base lengththan 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. 53.455.7); greater orbital diameter than B. cracens and B. fusca(61-76% postorbital length, vs. 47-58).

BETTA SPELENDENS SPECIES GROUP
Betta pugnax (Cantor)
Betta imbellis Ladiges
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

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 thehighlands of Kerinci is most probably introduced. This
population has adapted to the colder waters and did not takewell to the warmer temperatures of captivity in the lowlands.

Material examined. – Nias - ZMA 121.592 (11).

BETTA WASERI SPECIES GROUP
Remarks. – The specimens from Nias (ZMA 121.592) aretentatively 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 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 (1paratype).

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 (3paratypes); 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 thanB. 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 thanB. pugnax, B. cracens and B. fusca (35.5-36.5% SL, vs. 27.535.2); smaller predorsal length than B. fusca (62.7-66.3% SL,
vs. 68.5-70.2); greater preanal length than B. cracens (47.850.9% SL, vs. 42.0-46.1); greater body depth than B. cracens

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 thefollowing characters: µ-shaped black throat mark (vs. horseshoe 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).

125

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); presenceof both dorsal and caudal transverse bars (vs. absence); more
subdorsal scales (mode 61/2, vs. 5-6). Betta hipposideros canbe differentiated from B. renata in the following characters:
horseshoe-shaped black throat mark (vs. kidney-shaped); nonspotted opercle (vs. spotted opercle); opercle with yellow orbrown 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 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 beenpreviously 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 markon 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; andsubdorsal scales 6-61/2. Fresh material would be necessary toascertain 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.

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 (3paratypes); 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 thefollowing characters: kidney-shaped black throat mark (vs.
µ-shaped black throat mark); presence of both dorsal andcaudal transverse bars (vs. absence); spotted opercle (vs. nonspotted); opercle with black lower margin (vs. yellow margin);more subdorsal scales (mode 61/2, vs. 5-6). Betta renata canbe 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 (mode12, 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).

126

Paratypes - Riau Archipelago - ZRC 35418-20 (3 paratypes); CMK
10021 (2 paratypes); ZRC 35421-31 (11 paratypes); MZB (3paratypes); 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 canbe 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 beseparated 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. kidneyshaped 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:

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), 35.8-81.3mm SL. Selangor - ZRC 28530-31 (2), 84.8-106.4 mm SL; ZRC
15054 (1), 128.3 mm SL. Pahang - ZRC 25117-19 (3), 59.0-67.8mm 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), 73.7mm 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), 106.1mm SL. SUMATRA: Sumatra Utara - ZMA 121.701 (2), 43.454.4 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;abFig. 5. a. Diplopterus pulcher – reproduction of Gray, 1830: pl. 87;
b. Luciocephalus pulcher - ZRC 42520 (neotype), 106.2 mm SL,
Malaysia: Johor.

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), 58.4-110.5mm SL; ZRC 39505 (3), 17.5-96.7 mm SL; ZRC 39501 (1), 69.7mm SL; ZRC 42538 (6), 68.3-126.3 mm SL; ZRC 422 (1), 141.9mm 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 inlacking 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, whichappear circular when body is viewed ventrally (vs. absenceof such markings); yellowish caudal fin with 3-5 broad
vertical black bars (vs. brownish caudal fin with 3-7 narrowiridescent vertical gold bars); presence of markings on analand 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 longprotrusible 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 45.752.2% 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 halfof 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 (29.143.8% anal-fin base length); pelvic fin with filamentous firstray, 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 withinterrupted 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 withvery 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 proximallypigmented dark brown, rest hyaline; caudal fin yellowish with
3-5 broad vertical black bars, either in a regular or irregularpattern.
Preserved coloration. – Preserved coloration illustrated in127Tan & Ng: Anabantoidei and Channoidei of Sumatra
Table 3. Meristics and morphometrics of Luciocephalus pulcher and L. aura.

sample size, n
MERISTICS
vertebraedorsal-fin rayanal-fin ray
pelvic-fin ray
pectoral-fin ray
subdorsal scales
transverse scales
lateral scales
predorsal scales
postdorsal scales
caudal peduncle scales
lateral scales below dorsal-fin origin
lateral scales above anal-fin origin
MORPHOMETRICS - % SL
total length
body length
predorsal length
postdorsal lengthpreanal lengthcaudal peduncle length
head length
body depth
caudal peduncle depth
anal-fin base length
dorsal-fin base length
distance between pelvic- and anal-fin origins
pelvic-fin length
head width
snout length% HL
orbit diameter
postorbital length
head widthinterorbital width
snout length
upper jaw length

Luciocephalus pulcher

Luciocephalus aura

23

618-19 + 20-22(total 38-40, 39) (n = 28)
10-12 (11)0-I, 19-21 (total 19-21, 19)I, 5
16-18 (17)71/2-8 (81/2)
141/2-161/2 (151/2)
42-45 (43)60-65 (61)11-14 (13)91/2-12 (11)25-27 (26)
15-17 (15)17-19 + 20-22(total 38-40, 39) (n = 13)
11-12 (11)0, 18-21 (total 18-21, 20)
I, 5
16-17 (17)
81/2
151/2
43
60-63 (60)11-12 (12)10-11 (11)25-28 (26)15-16 (15)120.0-125.6
50.8-56.8
77.5-81.4
10.4-12.4
62.9-68.2
11.1-15.2
44.1-48.6
13.6-17.6
8.9-10.6
19.3-23.4
8.3-10.4
18.1-22.0
38.7-48.2
9.5-13.4
18.8-21.9

123.7-127.1
54.9-56.9
77.1-79.1
12.0-13.2
62.5-63.8
11.4-14.0
43.3-45.9
15.4-17.7
10.5-11.2
23.3-24.9
10.1-11.0
18.1-20.3
39.9-46.9
9.8-10.7
18.7-20.2

13.7-17.0
38.2-43.2
21.5-28.9
16.4-19.6
41.6-45.1
35.8-39.5

13.3-14.1
40.0-42.9
22.3-24.7
15.8-18.6
42.6-45.3
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 justabove anal-fin base, which appear as circular markings whenbody is viewed ventrally; broad dark brown preorbital andpostorbital 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 beneathpreorbital to central stripe covered with faint or strong darkbrown to black irregular markings; lower jaw whitish withblack 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.

128

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 foundonly in freshwater swamp forest and peat swamp habitats. It
is usually found among submerged vegetation along thebanks. It is an ambush predator, suddenly extending its funnellike 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 foundin the gular pouch (Kottelat et al., 1993: 159). Details on its
spawning as documented by Britz (1994b).

THE RAFFLES BULLETIN OF ZOOLOGY 2005
Remarks. – Further to the characters mentioned in the
diagnosis, L. pulcher can be differentiated from L. aura inhaving 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 analfin base length (35.4-42.3% body length, vs. 41.2-44.4); andgreater 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 expectedfrom L. pulcher.
The name Diplopterus pulcher was first used by Gray in thecaption 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 thenorthernmost 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 thatno 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 localityof 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 toascertain the precise locality from which the specimen in thedrawing is based (see Dawson (1946) for a history behind
Gray and Hardwicke’s illustrations). This uncertainty has notbeen a problem as it has long been accepted that
Luciocephalus is a monospecific genus. There has also neverbeen 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, insteadof 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 andpatterning, 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 asecond 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 thetype species of Luciocephalus Bleeker, 1851, and that thecurrent understanding of the name refers to the morewidespread species that occurs in Borneo, Malaysia andSumatra, 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 thepresent 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 (1paratype), 86.8 mm SL, ZRC 41715 (3 paratypes), 74.2-98.4 mm
SL; Jambi: Danau Semangkat, a lake connected to Batang Hari by

a

b

c

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 and102.2 mm SL, Jambi (Sungai Sentang).

129

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 (8paratypes), 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 inhaving numerous iridescent green spots on central stripe whenlive (vs. absence); absence of black irregular pattern on area
below preorbital to central stripe (vs. presence); area justabove 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 onanal 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 dorsalfin 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 longprotrusible 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 46.151.2% 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 thanhalf of anal-fin base (43-44% anal-fin base length); caudalfin 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 baselength); 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 andpostorbital dark brown broad stripe joining with central darkbrown 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 inFig. 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; broad130

dark brown preorbital and postorbital stripe joining withcentral stripe, ending at caudal peduncle with black spot;
middle of central stripe uniformly pigmented, green iridescentspots 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 proximallypigmented 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: Chelamaassi, 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 inlimited 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 fromL. pulcher in having slightly smaller preanal length (62.563.8% 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. 21.928.6); 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 inJambi. 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.

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 anddescribed 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 & Ngof body pattern. Ng & Kottelat (1998) had applied to theCommission to designate a neotype and had been successful(ICZN, 2000). The neotype (ZRC 31377) is a freshly collectedmale 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 theposterior extremity of the dorsal-fin base.

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 (15paratypes); CMK 11925 (6 paratypes); ZRC 30815-33 (19paratypes).
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 describedby Kottelat & Ng (1998), and can be found in Pulau Bintan,
Pulau Lingga and the island of Banka (pers. observ.). It canbe distinguished principally from P. deissneri in having arounded 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 canbe differentiated from P. sumatranus in missing theconspicuous black spot at the posterior extremity of the dorsalfin base; and a white pelvic fin filament (vs. black) (Kottelat& Ng, 1998).

Parosphromenus sumatranus Klausewitz †(Fig. 7)
Parosphromenus deissneri - Regan, 1910: 775 (part); Kottelat etal., 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).

aParosphromenus deissneri (Bleeker)
Osphronemus deissneri Bleeker, 1859: 376 (type locality:
Baturussak, Banka).
Polyacanthus deissneri - Günther, 1861: 381.
Parosphromenus deissneri - Bleeker, 1879: 20; Weber & deBeaufort, 1922: 348; Kottelat et al., 1993: 164 (part); Kottelat& Ng, 1998: 265, figs. 1-3.

bMaterial 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 islandof Banka, but recently also found in Biliton. Males havefilamentous extension to the median caudal ray. All previous
records of this taxon from Malay Peninsula and Borneo referto other taxa. The holotype (RMNH 6363) is in very poor
condition, with missing caudal fin, translucent body and loss

Fig. 7. Parosphromenus sumatranus - a. not preserved male, ca. 20mm SL, Jambi (Pijoan); b. SMF 3711 (paratype), 22.8 mm SL,
Jambi.

131

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. – 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. – 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. – 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)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.

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.

Trichogaster leerii (Bleeker) †Material examined. – Sumatra Utara - ZRC 42471 (1). Riau
Archipelago - ZRC 32600 (15). Jambi - ZRC 41739 (1); ZRC

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 & deBeaufort, 1922: 367.
Trichogaster leerii - Kottelat et al., 1993: 164.

a46211 (4); ZRC 43258 (2).
Remarks. – Several captive strains of T. trichopterus havebeen 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 asfollows (see Fig. 9): greyish body colour, darker at the dorsal,
almost white at ventral; head with dark grey dorsal andsilverish opercle scales with 2-3 blackish bars, eyes bright
red or orange when life; body with up to 20 blackish thinbars, 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,b

Fig. 8. Sphaerichthys selatanensis – a. not preserved; b. ZRC 46183,
32.4 mm SL, Biliton.

132

Fig. 9. Trichogaster trichopterus - ZRC 43258, 57.8 mm SL, Jambi(Bertam).

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, mostoften found in disturbed habitats (e.g. padi fields, ditches).
Specimens from Jambi have an iridescent green body colour.

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 notencountered 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 themarket shows more variation. The spots on the throat can beelongated and more blotchy. The spots on the lateral side ofbody is present in some, but absent in others.

SUB-ORDER CHANNOIDEIFAMILY CHANNIDAE

Channa gachua (Hamilton)

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 havebrownish transverse bars. For C. melasoma and C.
marulioides, the young is plain with orangish dorsal, withocellus on posterior part of dorsal fin and upper proximal
part of caudal fin respectively. For C. pleurophthalma, theyoung have a reticulated pattern of four to six longitudinalinterrupted black stripes on body, with faint black patcheson 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 theirclose affinity. Ng & Lim (1990) and Courtenay & Williams(2004) provide overviews of Channidae.

Ophiocephalus gachna – Volz, 1905: 131; 1907: 131.
Ophiocephalus gachua - Bleeker, 1879: 38; Perugia, 1893: 242;
Weber & de Beaufort, 1912: 539; 1915: 270; 1922: 321; deBeaufort, 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 bankanensis (Bleeker) *†Ophiocephalus bankanensis Bleeker, 1852b: 726 (type locality:
Toboali, Banka); 1879: 51; Günther, 1861: 475; Weber & deBeaufort, 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 3053032 (3). Banka - ZRC 30848 (1); ZRC 30934-38 (5).

Remarks. – This is a stenotopic blackwater species. Channabankanensis can be differentiated from C. lucius, even atjuvenile stage, in having a rounder head vs. pointed; numerous
spots on unpaired fins vs. bars and stripes; reddish-brownlife coloration vs. brownish-black. For more details on
taxonomy and morphological changes during growth, see Ng& Lim (1991) and Lee & Ng (1994) respectively.

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) andbrownish black life colour (vs. reddish brown).

133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.

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 differsin having a more pointed snout in lateral view (vs. blunt)(see Ng & Lim, 1991, for more details).

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, 187778, Pl. 399, fig. 2) with an ocellus on the upper proximal partof 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 withstriped pattern on dorsal, caudal and anal fins, with auniformly coloured body. Weber & de Beaufort (1922: 315,
fig. 84) depicted B. melanoptera as having irregular patchesof black on its lateral. We have come across and examined
specimens which have regularly spaced black patches oflateral scales with white posterior edges (ZRC 41508). The
identity of C. marulioides is characterised by the presence ofthe ocellus on the caudal fin, which is only distinct in young
individuals (ca. 50 mm SL). This ocellus later fades off andthe 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, fromrecent 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 theearlier conclusion.
The present series obtained from central Sumatra had four orfive patches of white edged black scales on posterior threequarters of body (Fig. 10). The ocellus on the caudal fin is
obscured by dark pigmentation. It is not commonly
encountered in the market.

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 patternconsisting of a metallic iridescent green base colour, with 34 lateral yellow to orange ocelli with black centre on the bodyaChanna melasoma (Bleeker, 1851) *Ophiocephalus melanosoma - Günther, 1861: 473; Bleeker, 1879:bFig. 10. Channa marulioides - ZRC 41508, ca. 420 mm SL, Jambi.

Fig. 11. Channa pleurophthalma - a. ZRC 41672, 46.9 mm SL,
Jambi (Ayer Hitam); b. aquarium stock, ca. 250 mm SL, Jambi(Ayer Hitam).

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
and an additional ocellus on both the opercle and caudal finbase, and orange patches on the base of the anal and caudal
fins (Fig. 11b). The young fish has a reticulated pattern of 46 longitudinal interrupted black stripes on body, with faintblack 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 (consistingof the Riau, Jambi and West Sumatra Provinces) includes
nearly all of the species previously recorded from the wholeof Sumatra (30 of 43 species = 70%) or from mainland
Sumatra (30 of 34 species = 88%); 21% (9 of 43 species) arefound exclusively in the smaller islands surrounding Sumatra.
Anabantoids and channids are a staple source of protein forthe local human population. They are found abundantly inlakes, rivers and streams. The presence of the labyrinth andaccessory organs enables them to survive out of the water inmoist conditions for extended periods of time. This featurefacilitates their transport to far-off markets for live sale.
Channids have firm and odourless flesh which is muchfavoured by locals (Ng & Lim, 1990). Trichogastertrichopterus 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 wellappreciated 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., Sphaerichthysosphromenoides, Trichogaster leerii, Channa gachua, C.
marulioides and C. pleurophthalma. This has made them
popular in the aquarium trade, and some species are evenbred commercially (e.g. Sphaerichthys osphromenoides andTrichogaster trichopterus; see Ng & Tan, 1997).
All species of Osphronemidae exhibit paternal care for theeggs and to a certain extent, to the fry. The genus Betta is
well noted for paternal care in oralbrooding and bubble nestbuilding species. There is also sibling brood care, as exhibitedby 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 anddonation 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, forproviding fresh material; M. Kottelat, Tommy H. T. Tan, S.
H. Tan, Darren C. J. Yeo and H. H. Ng, for their help andcompanionship in the field. This paper has been partially
supported by research grants RP 950326 and RP960314 toPN from the National University of Singapore.

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