THE RAFFLES BULLETIN OF ZOOLOGY 2005
THE RAFFLES BULLETIN OF ZOOLOGY 2005 Supplement No. 13: 43-99

THE FIGHTING FISHES (TELEOSTEI: OSPHRONEMIDAE: GENUS BETTA) OF
SINGAPORE, MALAYSIA AND BRUNEITan 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. – The taxonomy of the fighting fishes of the genus Betta from Singapore, Malaysia and Brunei
are revised, and 23 species are recognised. The taxonomy of some allied species in other parts of Southeast
Asia are also considered. Species groups for the entire genus are redefined and two new groups are recognised:
the B. bellica and B. albimarginata groups. Betta rubra is tentatively referred to the B. foerschi group. The
B. pugnax and B. splendens groups are the most widely distributed; whilst those of the B. coccina and B.
waseri groups are the most restricted, being essentially stenotopic blackwater species. The problematic B.
pugnax group is revised and two new species, B. stigmosa and B. lehi, are described from Peninsular Malaysia
and Sarawak, respectively. Betta climacura is regarded as a junior subjective synonym of B. akarensis; anda new species from the group, B. obscura, is described from Kalimantan Tengah. Betta ocellata is redescribedfrom fresh material and regarded as distinct from B. unimaculata; and from this group, two new species: B.
gladiator, from the Maliau Basin, Sabah; and B. pallifina, from Kalimantan Tengah, are described. Betta
taeniata (B. picta group) is redescribed from fresh material and its range is extended to Kalimantan Barat.
The taxonomy of B. imbellis (B. splendens group) is discussed and the complicated taxonomic status of B.
splendens and B. marchei dealt with. Betta marchei is suppressed and B. stiktos, new species, from Cambodia
is described.
KEY WORDS. – Betta, new species, Osphronemidae, taxonomy.

INTRODUCTION

The genus as currently defined, contains 48 recognised species(Kottelat et al., 1993; Ng, 1993; Kottelat & Ng, 1994; Tan& Tan, 1994, 1996; Ng & Kottelat, 1994; Kottelat, 1994,
1995; Tan & Ng, 1996, 2004a; Tan, 1998; Tan & Kottelat,
1998a, 1998b; Tan & Lim, 2004; Tan & Ng, 2004. Six
additional species are described here making it a total of 54
species known thus far from the genus (Table 1).

Betta Bleeker, 1850, is the most speciose genus in the family
Osphronemidae. Britz (1994, 1995, 2001) and Britz et al.
(1995) recently included in the Osphronemidae species
previously placed in the families Luciocephalidae andBelontiidae. The large number of species of Betta is probably
due to several factors. The warm and humid climate ofSoutheast Asia provides a stable environment with numerous
niches and habitats (Whitmore, 1986: 15). The presence ofmany types of water-bodies, e.g. rivers, ox-bow lakes, forest
streams, open country streams, hill streams, freshwater andpeat swamps supports a great diversity (see Kottelat & Ng,1994). Many species are also adapted to survive in extreme
environmental conditions, like in blackwaters with highacidity (pH between 3 and 5) in peat swamp forests. Betta
species also tend to have ranges restricted to one or a few
drainage systems, delimited by various obstacles likemountain ranges or large rivers (see Kottelat & Ng, 1994).

Preserved specimens of Betta are generally difficult to
identify. Colour is helpful in distinguishing live specimens,
but this is a character often difficult to use by ichthyologists
unfamiliar with the infraspecific variation. Colour may be
dependent upon maturity, sex, reproductive condition andgeographic variation. Large and old specimens tend to be
darker, which may mask body and opercle patterns, making
species identifications even more difficult (Roberts, 1989;
Kottelat et al.,1993; Kottelat & Ng, 1994). Juveniles are also
taxonomically unusable in most instances. Their
physiognomy is often quite different from that of the adult43Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei(e.g. relatively larger eyes, proportionately larger heads,
undeveloped or indistinct colour patterns, etc.) and most lacksufficient diagnostic characters to be useful (Ng, 1993).
Meristic characters are generally conservative, especiallywithin species groups. Species distinction based only on thepresence, absence or number of spines in the dorsal and anal
fins are often unreliable (Tan & Tan, 1996). Morphometrics
are useful to distinguish certain species, but characters ofgiven species groups are difficult to define objectively andmay possess some geographic variation (see Tan & Tan,
1996). Morphometric characters are useful for distinguishing
between species groups (e.g. B. coccina group), but is rather
conservative for members within the same group (see Tan &Ng, 1996). Fin and head shapes are generally usefulcharacters, especially when enough fresh material is available
to ascertain intraspecific variation. For example, membersof the B. bellica and B. unimaculata groups can be effectively
separated by their head shapes (Tan & Ng, 1996; present
study).

Natuurhistorisch Museum, Leiden (ex Rijksmuseum van
Natuurlijke Historie, RMNH); Sabah Museum, Kota
Kinabalu, Sabah (SBM); Sabah Parks Collection, Sabah(SBP); Sarawak Museum, Kuching, Sarawak (SM);Senckenbergischen Naturforschenden Gesellschaft, Frankfurt
am Main (SMF); Universiti Malaysia Sabah, Sabah (UMS);
U.S. National Museum of Natural History, Smithsonian
Institution, Washington D.C. (USNM); Instituut voor
Systematiek and Populatiebiologie, Universiteit van
Amsterdam (ZMA); Zoologisches Museum der Universität,
Zürich (ZMZ); and the Zoological Reference Collection of
the Raffles Museum of Biodiversity Research, National
University of Singapore (ZRC).
Toponymy: Kampung = village; Batang or Sungai = river;
Bukit or Gunung = hill; Tasik = lake; Pulau = island.
Abbreviations used: SL = standard length; TL = total length;and HL = head length.
A diagnosis of each species group is provided and this isfollowed by the treatment of each species covered in the study
area. The term ‘species group’ is used here for an assemblageof species sharing a set of diagnostic characters and which
may or may not be a monophyletic lineage. For the two
recently revised species groups, B. coccina and B. waseri,
only a diagnosis of each species is provided. Instead, thereader is referred to Witte & Schmidt (1992), Ng & Kottelat(1992, 1994) and Tan (1998) for the detailed reports on the
B. coccina and B. waseri groups.

This present revision deals with the species native toSingapore, Malaysia (Peninsular Malaysia, Sarawak, Sabah)and Brunei; but also includes the description of three new
species from Kalimantan Tengah and Cambodia. The feral
B. splendens populations which are found in Peninsular
Malaysia and Singapore are excluded in this study (Tweedie,
1952a, b; Lim & Ng, 1990; Ng et al., 1993). Certain species
found just outside the present area of study have also been
included (e.g. B. unimaculata, B. patoti) to clarify theirtaxonomic status and aid in the general discussions.

All measurements were taken point to point on the left sideof the specimens with dial calipers allowing readings to thenearest 0.05 mm. Methods for obtaining meristic andmorphometric data follow Witte & Schmidt (1992), exceptthat all measurements are taken from point to point and thatthe predorsal scale counts are counted continuously (following
Ng & Kottelat, 1994). Chin-bar is used to refer to the darkthin stripe running from throat area below snout to eye, justunder the preorbital stripe (Fig. 1). The pattern of dark,narrow, often slightly curved bars perpendicular to the rayson the interradial membranes of the dorsal, caudal and anal
fins are called dorsal, caudal and anal transverse bars (Fig.
2) (Tan & Kottelat, 1998a).

MATERIAL AND METHODS
The species concept used is the phylogenetic species concept(Cracraft, 1989; Warren, 1992; see discussion in Kottelat,
1997, and methods in Tan & Kottelat, 1998a). For theliterature citation, only primary scientific literature is used.
However, in some cases, useful aquarium literature has been
cited, for instance the books by Linke (1991) and Kubota etal. (1996). These two books are covered, even though thecoverage of the genus Betta is incomplete and certain
information is inaccurate (e.g. locality data, scientific names)
primarily because of their excellent illustrations and synopses(where English versions are available). Other aquarium
literature are used only when useful information on breedingand habitats are provided.

Freshly caught specimens were fixed in 10% formalin solution
for one to two weeks, then placed into water for leaching
before permanent storage in 70-75% denatured ethanol.
Colour descriptions are based on live and acclimatised
specimens unless otherwise stated. The keys provided are foruse with mature specimens only.

Specimens examined are deposited in the Natural History
Museum, London (ex British Museum of Natural History,
BMNH); Muzium Brunei, Brunei Darussalam (BRM);
California Academy of Sciences, San Francisco (CAS); thecollection of Maurice Kottelat, Cornol (CMK); Danum Valley
Field Centre, Lahad Datu, Sabah (DVFC); Field Museum of
Natural History, Chicago (FMNH); Muséum d’Histoire
Naturelle, Genève (MHNG); Muséum National D’Histoire
Naturelle, Paris (MNHN); Museo Civico di Storia Naturale,
Genova (MSNG); Research and Development Centre for
Biology (ex Museum Zoologicum Bogoriense), The
Indonesian Institute of Sciences (MZB); Nationaal

TAXONOMIC KEYS
The keys to the species groups and members of the species
group from the study area are based upon morphologicalfeatures the authors feel are easy to use. The size ranges are
based on wild specimens. Captive specimens can generally
grow larger.

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
Table 1. Betta species list and localities
Betta bellica group
Betta bellica Sauvage, 1884: Malaysia (Selangor*, Perak, Pahang, Johor); Sumatra (North Sumatra)
Betta simorum Tan & Ng, 1996: Sumatra (Jambi*, Riau, South Sumatra)
Betta pugnax group
Betta pugnax (Cantor, 1850): Malaysia (Pulau Penang*, Kedah, Terengganu, Pahang, Selangor, Johor); Singapore; Sumatra (Riau, Jambi),
AnambasBetta fusca Regan, 1910: Sumatra (North Sumatra*)Betta schalleri Kottelat & Ng, 1994: Sumatra (Banka*)
Betta prima Kottelat, 1994: Thailand (East Thailand*); Cambodia (West Cambodia)
Betta enisae Kottelat, 1995: Kalimantan Barat (Kapuas*)
Betta pulchra Tan & Tan, 1996: Malaysia (Johor*)
Betta breviobesus Tan & Kottelat, 1998: Kalimantan Barat (Kapuas*)
Betta lehi, new species: Sarawak (south Sarawak*); Kalimantan Barat (Kapuas)
Betta stigmosus, new species: Malaysia (Terengganu*)
Betta akarensis group
Betta akarensis Regan, 1910: Sarawak (north to central* Sarawak); Brunei (Belait, Tutong and Bandar Seri Begawan Districts)
Betta balunga Herre, 1940: Sabah (Tawau*); Kalimantan Timur (Sebuku, Mahakam)
Betta chini Ng, 1993: Sabah (Beaufort*)
Betta pinguis Tan & Kottelat, 1998: Kalimantan Barat (Kapuas*)
Betta aurigans Tan & Lim, 2004: Natuna Besar*
Betta ibanorum Tan & Ng, 2004: Sarawak (south Sarawak*)Betta obscura, new species: Kalimantan Tengah (Barito*)
Betta unimaculata group
Betta unimaculata (Popta, 1905): Kalimantan Timur (Kayan*, Howong*)
Betta macrostoma Regan, 1910: Sarawak (north Sarawak*); Brunei (Belait District)
Betta patoti Weber & de Beaufort, 1922: Kalimantan Timur (Balikpapan*, Samarinda)
Betta ocellata de Beaufort, 1933: Sabah (Sandakan*, Kinabatangan, Lahad Datu, Tawau); Kalimantan Timur (Sebuku)
Betta gladiator, new species: Sabah (Maliau*)
Betta pallifina, new species: Kalimantan Tengah (upper Barito*)
Betta picta group
Betta picta (Valenciennes, in Cuvier & Valenciennes, 1846): Java (Bogor*, Bandung, Yogjarkata)
Betta taeniata Regan, 1910: Sarawak (south Sarawak*); Kalimantan Barat (Kapuas)
Betta simplex Kottelat, 1994: Thailand (south Thailand - Krabi*)
Betta falx Tan & Kottelat, 1998: Sumatra (Jambi*, Sumatra Utara)
Betta splendens group
Betta splendens Regan, 1910: Thailand (central Thailand*)Betta smaragdina Ladiges, 1972: Thailand (east and north* Thailand)
Betta imbellis Ladiges, 1975: Thailand (south Thailand); Malaysia (Selangor*, Pulau Penang, Perak, Kedah, Terengganu, Johor); Singapore;
Sumatra (North Sumatra)
Betta stiktos, new species: Cambodia*
Betta coccina group
Betta coccina Vierke, 1979: Sumatra (Jambi*, Riau); Malaysia (Johor)
Betta tussyae Schaller, 1985: Malaysia (Pahang*)
Betta persephone Schaller, 1986: Malaysia (Johor*)
Betta rutilans Witte & Kottelat, in Kottelat, 1991: Kalimantan Barat (Kapuas*)
Betta brownorum Witte & Schimdt, 1992: Sarawak (central* to south Sarawak; Kalimantan Barat)
Betta livida Ng & Kottelat, 1992: Malaysia (Selangor*)
Betta miniopinna Tan & Tan, 1994: Sumatra (Riau - Bintan*)
Betta burdigala Kottelat & Ng, 1994: Sumatra (Banka*)
Betta waseri group
Betta waseri Krummenacher, 1986: Malaysia (Pahang*, Terengganu, Kuantan)
Betta hipposideros Ng & Kottelat, 1994: Malaysia (Selangor*); Sumatra (Riau)
Betta spilotogena Ng & Kottelat, 1994: Sumatra (Riau - Bintan*, Singkep)
Betta chloropharynx Kottelat & Ng, 1994: Sumatra (Banka*)
Betta renata Tan, 1998: Sumatra (Jambi*, South Sumatra)
Betta pi Tan, 1998: Thailand (south Thailand - Sungai Kolok*); Malaysia (Kelantan)

45

Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei
Table 1. Betta species list and localities (Cont'd)
Betta edithae group
Betta edithae Vierke, 1984: Kalimantan Selatan*, Tengah and Barat; Sumatra (Riau - Bintan, Banka, Biliton)
Betta foerschi group
Betta foerschi Vierke, 1979: Kalimantan Tengah (Mentaya*)
Betta strohi Schaller & Kottelat, 1989: Kalimantan Tengah (Sukamara*)?Betta rubra Perugia, 1893: Sumatra (North Sumatra*, Aceh)
Betta anabatoides group
Betta anabatoides Bleeker, 1851: Kalimantan Selatan* and Tengah
Betta albimarginata group
Betta albimarginata Kottelat & Ng, 1994: Kalimantan Timur (Sebuku*)
Betta channoides Kottelat & Ng, 1994: Kalimantan Timur (Mahakam*)
Betta dimidiata group
Betta dimidiata Roberts, 1989: Kalimantan Barat (Kapuas*)* type locality
5. Body depth 18.5-24.7% SL; opercle scales with green
iridescence or red patch when alive; caudal transverse bars
present or absent, or caudal fin with black margin; pelvic
fins with distinct filamentous anterior rays; adult size large(SL more than 60 mm) ............. Betta unimaculata group– Body depth 24.2-32.2% SL; opercle scales with green
iridescence when live; caudal transverse bars absent orpresent, caudal fin with red distal crescent margin when
alive, yellowish when preserved (Fig. 5a); pelvic fins withall rays elongate; adult size small (SL less than 40 mm)...................................................... Betta splendens group
6. Body brown; iridescent green scales on body, lateral medianiridescent scales absent; pelvic fin hyaline, tip white when
alive; adult size medium to large (SL more than 40 mm)........................................................................................ 7– Body red or black; no iridescent scales on body, lateral medianiridescent green scales present in some species (Fig. 6a);pelvic fin red or black, pelvic fin tip black, white or green
when alive; adult size small (SL 25-40 mm) ....................
......................................................... Betta coccina group
7. Head large; caudal transverse bars distinct in male; lanceolatecaudal fin with 2-3 median rays elongated; anal fin withposterior 1-2 rays elongated into filaments in male; anal-fin
rays 27-30; dorsal-fin rays 8-11 .... Betta akarensis group– Head small; caudal transverse bars absent; rounded caudal fin
with 4-6 median rays elongated into filaments; anal fin withposterior 3-4 rays elongated into filaments in male; anal-fin
rays 30-33; dorsal-fin rays 11-13 ...... Betta bellica group

Key to the Betta species groups occurring in Malaysia,
Singapore and Brunei
1. Males with dark iridescent opercle scales when alive; with darkly
pigmented opercle and throat when preserved .............. 2– Males without dark iridescent opercle scales when alive; opercle
and throat not darkly pigmented when preserved ......... 6
2. Body brown; iridescent opercle scales blue or green when alive;chin-bar present or absent (Fig. 1) ................................. 3– Body clay-yellow; iridescent opercle scales gold (present inmature male of all species except B. tomi) when alive; throat
with black blotches which may join with black lower jaw(Fig. 3); chin-bar absent .................... Betta waseri group
3. Unpaired fins without broad, coloured margins when alive, thindark margin may be present when preserved; chin-bar
present or absent; adult size small (SL less than 40 mm) or
large (SL more than 60 mm) ......................................... 4– Unpaired fins with broad iridescent green or blue margin when
alive, dark when preserved (B. taeniata mature males) (Fig.
4); chin-bar present; adult size medium (SL between 40 and
60 mm) .................................................. Betta picta group
4. Caudal fin rounded (Fig. 5a); caudal transverse bars absent orpresent; chin-bar present or absent; adult size large or small,body depth variable ........................................................ 5– Caudal fin lanceolate (Fig. 5b-c); caudal transverse bars usuallypresent (mature male) (Fig. 2); chin-bar present; adult size
medium to large, body relatively stocky (body depth 2437% SL) ............................................ Betta pugnax group

v

iii

i

ii

iv

vii

vi

Fig. 1. Schematic diagram of an adult fighting fish showing: i)preorbital stripe, ii) chin-bar, iii) postorbital stripe, iv) second
postorbital stripe, v) central stripe, vi) second central stripe, vii)
black spot at base of caudal fin.

Fig. 2. Transverse bars on caudal fin (similar bars on dorsal andanal fins).

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
Key to the Betta species
The key is for 47 out of 55 species of Betta recognised and is only
for the species groups covered in the study area. The species coveredin the study area are indicated by an asterisk (*). All characters listed
here are for mature specimens only. When colour notes are used,
they are for live and aquarium-acclimatised specimens.

3.
–

4.

Betta bellica group
1. Head slanted, dorsal surface distinctly concave just behind eye;pelvic fin filamentous tip extending up to 14th anal ray(pelvic fin length 31.3-48.3% SL); lateral scales 331/2-35(mode 34); adpressed pelvic fin reaching beyond anal-fin
origin (distance between pelvic and anal-fin origins 8.513.1% SL, mean 9.9). Central to southern Sumatra,
Kalimantan Barat (Kapuas) ...................... Betta simorum– Head rounded, dorsal surface not concave just behind eye; pelvic
fin filamentous tip extending up to 8th anal ray (pelvic fin
length 23.6-38.8% SL); lateral scales 33-34 (mode 33);
adpressed pelvic fin not reaching anal-fin origin (distance
between pelvic and anal-fin origins 10.5-12.6% SL, mean
11.2). Peninsular Malaysia, northern Sumatra ...................
.................................................................... Betta bellica*

–

5.
–
6.
–
7.

Betta pugnax group
1. Dark anal and caudal fin distal margins when preserved (blueish
when alive) ..................................................................... 2– No dark anal and caudal fin distal margins when preserved.......................................................................................... 5
2. No dorsal, caudal and anal transverse bars in male and female;no interrupted second postorbital stripe on opercle in maleand female (Fig. 1); interorbital width 64-84% postorbital
length .............................................................................. 3– Dorsal, caudal and anal transverse bars present in male andfemale (Fig. 2); interrupted second postorbital stripe on

–

8.

–

opercle present in male and female; interorbital width 6265% postorbital length. Peninsular Malaysia (Terengganu)................................................................ Betta stigmosa*No caudal transverse bars; lateral scales 28-30; predorsal length62.1-67.5% SL ................................................................ 4
Caudal transverse bars present; lateral scales 26-28; predorsal
length 67.3-70.3% SL. Kalimantan Barat (middle Kapuas)...................................................................... Betta enisae
Two black marks below postorbital stripe on opercle in female
and juvenile (Fig. 6b); chin-bar present; orbital diameter
27.2-32.2% HL; interorbital width 70-84% postorbital
length. Southern Sarawak, Kalimantan Barat (lower Kapuas)........................................................................ Betta lehi*
No black marks below postorbital stripe on opercle; no chinbar; orbital diameter 24.2-28.5% HL; interorbital width 4869% postorbital length. Kalimantan Barat (upper Kapuas)............................................................. Betta breviobesus
No chin-bar ........................................................................... 6
Chin-bar present (Fig. 1) ...................................................... 7
Anal-fin rays 27; dorsal-fin rays 10-11; lateral scales 31;
predorsal scales 17-19. Banka .................. Betta schalleri
Anal-fin rays 24-25; dorsal-fin rays 8-9; lateral scales 29;
predorsal scales 22-23. Northern Sumatra ..... Betta fusca
No complete second postorbital stripe on opercle; caudal
transverse bars present; predorsal scales 18-21 (mode 20);preanal length 40.6-51.2% SL ....................................... 8
Complete second postorbital stripe on opercle (Fig. 1); no caudal
transverse bars; predorsal scales 20-22 (mode 22); preanallength 50.6-54.8% SL. Cambodia, eastern Thailand .........
Betta prima
Caudal fin narrowly lanceolate in male (Fig. 5c); opercle scales
iridescent green when alive; body depth 26-37% SL; pelvicfin length 29.1-53.2% SL; adult size medium (SL up to55mm); stenotopic blackwater species. Peninsular Malaysia(Pontian) .................................................... Betta pulchra*Caudal fin broadly lanceolate in male (Fig. 5b); opercle scales
iridescent blue when alive; body depth 24-32% SL; pelvicfin length 26.6-43.8% SL; adult size large (SL up to 67 mm);clear to brown water species. Peninsular Malaysia,
Singapore, Sumatra (Riau), Anambas, Natuna ..................
................................................................... Betta pugnax*

Betta akarensis group
1. Opercle pigmentation absent to relatively dense, with 3 or fewer
dark marks below postorbital stripe (Fig. 7); body slender
to relatively stocky (body depth 23.3-32.7% SL); dorsal-fin
origin above lateral scales 14-18 ..................................... 2– Opercle pigmentation dense, with 5-7 dark marks below
postorbital stripe (Fig. 7d); body stocky (body depth 30.631.6% SL); dorsal-fin origin above lateral scales 18-19.
Kalimantan Barat (Upper Kapuas) .............. Betta pinguis

Fig. 3. Schematic diagrams of throat patterns of the Betta waseri
group and unique eye coloration pattern: a) B. waseri, b) B.
hipposideros, c) B. tomi, d) B. spilotogena, e) B. chloropharynx, f)
B. renata, g) B. pi, h) unique eye coloration pattern.

Fig. 4. Schematic diagram of an adult Betta taeniata having unpaired
fins with broad iridescent darkly pigmented margin.

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Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei
2. Predorsal scales 21-26; dorsal-fin rays modally 9; subdorsal
scales modally 5-6; vertebrae 29-32 (modes 30, 31) .... 3– Predorsal scales 19-21 (mode 20); dorsal-fin rays modally 10;
subdorsal scales modally 61/2; vertebrae 28-30 (mode 28).
Kalimantan Tengah (Barito) ....................... Betta obscura
3. Opercle with second postorbital stripe (complete or interrupted)(Fig. 1) ............................................................................ 4– Opercle without second postorbital stripe ............................ 6
4. Opercle with interrupted second postorbital stripe; lateral scales
29-31 (mode 31); vertebrae 29-31; adult size medium (SL40-60 mm) ...................................................................... 5– Opercle with complete second postorbital stripe; lateral scales
31-33 (mode 32); vertebrae 31-32; adult size large (SL 6080 mm). Central to northern Sarawak, Brunei ..................
............................................................... Betta akarensis*
5. Eye yellow when alive; lateral scales 30-31; preanal length 42.645.7% SL; anal-fin base length 54.8-58.5% SL. Western
Sabah ............................................................. Betta chini*– Eye orange to red when alive; lateral scales 29-30; preanal length48.5-52.6% SL; anal-fin base length 47.8-55.4% SL. Eastern
Sabah, Kalimantan Timur (Sebuku) ....... Betta balunga*
6. Iridescent green to blue on posterior half of scales when alive;
lateral scales 30-32 (mode 30); predorsal scales 21-23 (mode22); anal-fin rays 27-29 (mode 28). Southern Sarawak ....
................................................................ Betta ibanorum*– Gold scales on belly and body when alive; lateral scales 33-331/2(331/2); predorsal scales 24-26; anal-fin rays 29-30 (mode30). Natuna (Pulau Bunguran) ................. Betta aurigans

2.

–

3.
–

4.

–

5.

–

Betta unimaculata group
1. Male with yellowish to brown body; no red blotch on opercle,
opercle scales iridescent; no dorsal fin ocellus; dorsal-fin
rays 6-9; anal-fin rays 27-33; subdorsal scales 4-61/2 ... 2– Male with red body; a red blotch on opercle, opercle scales not
iridescent; dorsal fin ocellus present (Fig. 8a); dorsal-fin rays
9-11; anal-fin rays 25-28 (mode 27); subdorsal scales 6-8(mode 7). Brunei, northern Sarawak ..................................
............................................................ Betta macrostoma*
Dark opercular scales present when preserved; no dark verticalbars on body; caudal fin rounded or lanceolate; dorsal, caudal
and anal transverse bars present; anal-fin rays 27-31 ..... 3
No dark opercular scales absent when preserved; 10-14 darkvertical bars on lower half of body; caudal fin lanceolate;no transverse bars on dorsal, caudal and anal; anal-fin rays
30-33 (mode 31). Kalimantan Timur (Balikpapan) ...........
....................................................................... Betta patoti
Sheath scales on anal base; pectoral fin rays 11-14 (mode 12 or13); lateral scales 31-34 (mode 32 or 33) ..................... 4
No sheath scales on anal base; pectoral fin rays 16-17 (mode17); lateral scales 34-36 (mode 35). Sabah (Maliau) ........
................................................................ Betta gladiator*
Female with subdistal margin of anal fin uniformly coloured;
subdorsal scales modally 5 or 51/2; predorsal length 67.773.0% SL; head width 19.6-24.0% SL .......................... 5
Female with subdistal margin of anal fin orange when live,
hyaline when preserved; subdorsal scales modally 6;
predorsal length 64.6-69.4% SL; head width 17.7-20.8% SL
Kalimantan Tengah (Barito) ...................... Betta pallifina
Head blunt, slightly truncate (Fig. 8b); anal-fin rays 27-30 (mode28 or 29); lateral scales 31-33 (mode 32); length of anal-finbase 45.9-53.7% SL; length of dorsal-fin base 7.9-10.4%
SL; head width 19.6-21.8% SL. Kalimantan Timur (Kayan,
Howong) ............................................. Betta unimaculata
Head pointed, slightly conical (Fig. 8c); anal-fin rays 30-31(mode 30); lateral scales 32-34 (mode 33); length of analfin base 52.2-54.5% SL; length of dorsal-fin base 10.511.3% SL; head width 21.0-24.0% SL. Sabah, Kalimantan
Timur (Sebuku) ......................................... Betta ocellata*

Betta picta group
1. Male with blueish to greenish iridescent opercle scales and chinbar indistinct; body depth 25.3-32.2% SL; length of analfin base 48.3-52.7% SL .................................................. 2bacFig. 5. Schematic diagram of caudal fins: a) rounded caudal fin with darkly pigmented distal margin (B. imbellis), b) broadly lanceolatecaudal fin (B. pugnax), c) narrowly lanceolate caudal fin (B. pulchra).

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THE RAFFLES BULLETIN OF ZOOLOGY 2005–

Male with yellowish gold to greenish-blue iridescent opercle
scales, chin-bar distinct and part of second postorbital stripe
visible; body depth 21.5-28.0% SL; length of anal-fin base42.6-50.3% SL ................................................................ 3
2. Body and fins yellowish, head blackish, iridescent blue scales
present on dorsal half of body in male; anal-fin rays 23-26(mode 26); transverse scales 9-91/2 (mode 91/2); predorsal
length 62.4-67.7% SL; head length 30.3-34.5% SL; body
depth 25.3-30.1% SL. Southern Sarawak, Kalimantan Barat(Kapuas) ................................................... Betta taeniata*– Body and fins pale reddish, head dark reddish, no iridescent
blue scales on body in male; anal-fin rays 23-24 (mode 24);transverse scales 91/2-111/2 (mode 111/2); predorsal length66.7-69.8% SL; head length 34.6-37.2% SL; body depth
29.3-32.2% SL. Southern Thailand (Krabi) .......................
.................................................................... Betta simplex
3. Opercle scales iridescent yellowish gold; blueish anal and caudal
fin margins; faint dorsal transverse bars in male; faint caudal
transverse bars in female; dorsal-fin origin above 12-14th
lateral scale. Java (Bogor, Bandung, Ambarawa) .............
........................................................................ Betta picta– Opercle scales iridescent greenish blue; reddish anal and caudal
fin margins; distinct dorsal transverse bars in male; distinct
caudal transverse bars female; dorsal-fin origine above 1112th lateral scale. Sumatra (Sumatra Utara, Jambi) ..........
.......................................................................... Betta falx

postorbital stripes distinct when preserved. Cambodia .....
...................................................................... Betta stiktos
3. Body dark brown to black, head blackish; opercle with iridescent
blue scales; fins iridescent blue; no caudal transverse bars;
caudal fin iridescent blue with distal red crescent margin(Fig. 5a); distal half of posterior anal-fin rays red. Southern
Thailand, Peninsular Malaysia, northern Sumatra .............
.................................................................. Betta imbellis*– Body light brown, head brown; opercle with iridescent green
scales; fins reddish with iridescent green rays; caudal
transverse bars present; no distal red crescent margin on
caudal fin; distal half of posterior anal-fin rays not red.
Northern and eastern Thailand, Laos .. Betta smaragdina

Betta coccina group
1. Body blackish; pelvic fin blackish or red; adult size 25-30 mm
SL .................................................................................... 2– Body reddish; pelvic fin red; adult size 30-40 mm SL ....... 3
2. Dorsal surface of head behind eye sloping gently; pelvic fin often
blackish, sometimes red; predorsal scales 17-18; lateralscales 27-29. Peninsular Malaysia (Ayer Hitam, Muar) ...
............................................................ Betta persephone*– Dorsal surface of head behind eye sloping steeply; pelvic finalways red; predorsal scales 20; lateral scales 30-31. Sumatra(Bintan) ................................................. Betta miniopinna
3. A mid-lateral iridescent green blotch on body (Fig. 6a) ..... 4– No mid-lateral iridescent green blotch on body .................. 6
4. Male and female with a mid-lateral iridescent green blotch on
body; tip of pelvic fin green or white ........................... 5

Betta splendens group
1. Iridescent blue or green scales on opercle, fins iridescent blue
or green in male; head and body slender (body depth 24.230.1% SL) ....................................................................... 2– No iridescent scales on opercle, with parallel red vertical bars,
fins blue, green or red in male; head and body stocky (bodydepth 27.1-32.2% SL). Central Thailand ...........................
................................................................. Betta splendens
2. No transverse bars on caudal fin; body with more than 8 verticalrows of dark marks on posterior edge of scales; lateral scales
29-32 (mode 30-31); male with chin-bar, postorbital andsecond postorbital stripes distinct when preserved ....... 3– Transverse bars on caudal fin in male and female; body with 67 vertical rows of dark marks on posterior edge of scales;
lateral scales 31-34 (mode 33); male with only secondafbFig. 6. a) Schematic outline of Betta livida showing median lateralblotch, b) head of B. lehi showing interrupted second postorbital
stripe, usually present as a series of two black blotches.

Fig. 7. Schematic diagrams of opercle patterns of the Betta akarensis
group: a) B. akarensis, b) B. balunga, c) B. chini, d) B. pinguis, e)
B. ibanorum and B. aurigans, f) B. obscura.

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Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei–

Only male with a mid-lateral iridescent green blotch on body;tip of pelvic fin black ............................... Betta coccina*
5. Discrete iridescent green spots on median fins; parallel vertical
iridescent gold bars on opercle; tip of pelvic fin wide,
iridescent green; body dark red. Peninsular Malaysia(Selangor) ..................................................... Betta livida*– No discrete iridescent green spots on median fins; no parallel
vertical iridescent gold bars on opercle; tip of pelvic fin
filamentous, white; body magenta red. Southern to central
Sarawak ............................................... Betta brownorum*
6. No discrete iridescent green spots on median fins; subdorsal
scales 8-10; dorsal-fin rays 11-12; anal-fin rays 27-28 .... 7– Discrete iridescent green spots on median fins; subdorsal scales
11-111/2; dorsal-fin rays 14-15; anal-fin rays 24-26. Sumatra(Banka) ..................................................... Betta burdigala
7. Faint parallel vertical iridescent gold bars on opercle; body
reddish-orange with darker central and second central
stripes; subdorsal scales modally 8. Peninsular Malaysia(Pahang) ..................................................... Betta tussyae*– No faint parallel vertical iridescent gold bars on opercle; bodyuniform deep red; subdorsal scales modally 9. Kalimantan
Barat (Kapuas) ............................................ Betta rutilans

Betta waseri group
1. Dark blotch on throat not joined to black lower lip (Fig. 3) .. 2
– Dark blotch on throat joined to black lower lip (Fig. 3) .... 5
2. Anal fin never with dark distal margin; throat pattern with nonelongated median dark blotch; subdorsal scales 51/2-7 ..... 3– Anal fin usually with broad and distinct dark green distal margin;
throat pattern with elongated semi-horseshoe dark blotch(Fig. 3c); subdorsal scales 5-51/2 (mode 5). Peninsular
Malaysia (Johor) ............................................ Betta tomi*
3. Throat pattern with two discrete or semi-fused median dark
blotches; no dorsal and caudal transverse bars; subdorsal
scales 51/2-61/2 (51/2 or 6); lateral scales 31-321/2 (mode 32);
anal-fin rays 28-31 (mode 30) ....................................... 4– Throat pattern with a single median kidney-shaped dark blotch(Fig. 3f); faint dorsal and caudal transverse bars; subdorsal
scales 6-7 (61/2); lateral scales 31-32 (mode 31); anal-fin
rays 28-31 (mode 29). Central Sumatra ...... Betta renata
4. Throat with tear-drop shaped black blotches (Fig. 3a); opercle
not spotted, with dark blotch on posterior edge; subdorsal
scales 6-61/2 (mode 6); postdorsal scales 101/2-12 (mode 11);body depth 23.1-27.6% SL. Peninsular Malaysia (Pahang,
Terengganu) ................................................ Betta waseri*– Throat with a median pear-shaped black blotch (Fig. 3d); opercle
with 7 or more spots; subdorsal scales 51/2-6 (mode 51/2);
postdorsal scales 10-101/2 (mode 10); body depth 25.6-28.7%
SL. Sumatra (Bintan) ............................ Betta spilotogena
5. Throat with a pair of elongated black bars; no iridescent greenishgold opercle scales ......................................................... 6– Throat with ∞-shaped black pattern (Fig. 3e); iridescentgreenish-gold opercle scales. Sumatra (Banka) ... Betta
chloropharynx
6. Throat with horseshoe shaped black mark (Fig. 3b); dorsal andcaudal transverse bars present; anal fin without dark distal
margin; lower margin of opercle brown; subdorsal scales
61/2-7 (mode 61/2); postdorsal scales 9-10 (mode 91/2); lateralscales 31-32 (mode 32). Peninsular Malaysia (Selangor),central Sumatra ................................. Betta hipposideros*– Throat with pi (π)-shaped black mark (Fig. 3g); no dorsal andcaudal transverse bars; anal fin with broad and distinct dark
blue distal margin; lower margin of opercle black; subdorsal
scales 51/2-6 (mode 6); postdorsal scales 11-12 (mode 11);
lateral scales 32-33 (2mode 33). Southern Thailand (Sungai
Kolok), Peninsular Malaysia (Kelantan) ........... Betta pi*

aTAXONOMY
Betta Bleeker, 1850
Betta Bleeker, 1850: 12 (type species Betta trifasciata Bleeker, 1850,
by monotypy).
Micracanthus Sauvage, 1879: 95 (type species Micracanthus
marchei Sauvage, 1879, by monotypy).
Parophiocephalus Popta, 1905: 184 (type species Parophiocephalus
unimaculatus Popta, 1905, by monotypy).
Oshimia Jordan, 1919: 342 (unnecessary replacement name forMicracanthus Sauvage, 1879).
Pseudobetta Richter, 1981: 273 (type species Macropodus pugnax
Cantor, 1850, by original designation).

bRemarks. – The first fighting fish species described wasPanchax pictum by Valenciennes (in Cuvier & Valenciennes,
1846) from Java. The genus name Betta, however, was only
established by Bleeker (1850) for his new species, B.
trifasciata, also from Java. Cantor (1850) describedcFig. 8. a) Dorsal fin ocellus of B. macrostoma, b) dorsal, lateral andventral head profiles of B. unimaculata, c) dorsal, lateral and ventral
head profiles of B. ocellata.

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
Macropodus pugnax from the Malay Peninsula and Bleeker(1851) described B. anabatoides, from Borneo. Bleeker(1879), however, decided that all described species of fighting
fishes up to that time were conspecific with B. picta.

Schaller (1985) and Schaller & Kottelat (1989) separated thegenus into two categories: mouthbrooders and bubblenest
brooders. Schaller (1985) further divided the bubblenest
brooders into the B. splendens and B. bellica groups. These
two groups were not described with apomorphies, but withsuperficial similarities, behavioural and physiological
adaptations to different environments, viz. B. splendens group(B. imbellis, B. smaragdina, B. splendens, B. persephone, B.
strohi): characterised by a compressed and relatively slenderbody, presence of two vertical iridescent patches on opercle
and members inhabiting stagnant to slow flowing waters; andB. bellica group (B. bellica, B. coccina and B. tussyae):characterised by a cylindrical and slender body, dark stripe
running through snout and eye to opercle edge (preorbital
and postorbital stripes) with iridescent scales and members
inhabiting flowing waters.

Sauvage (1879) described Micracanthus marchei on the basisof a single specimen with the locality data “bassin del’Ogooué“ in Africa. The erroneous locality data is discussed
by Roberts (1981) who argued that M. marchei is a synonymof B. splendens. Roberts (1981) also commented that the
specimen of M. marchei was most probably from Thailand
or Peninsular Malaysia. There are, however, complications
as there are several species in that same region which could
fit Sauvage’s (1879) description. For example, Roberts (1981)
did not take into consideration B. smaragdina and B. imbellis.
In any case, there is no doubt that Micracanthus, is asubjective junior synonym of Betta. Oshimia was proposed
by Jordan (1919) to replace Micracanthus because he thoughtthat the genus name had been used earlier by Swainson (1839)
for Microcanthus (family Chaetodontidae). Jordan’s action
was unnecessary because the spellings for the names are not
identical. The names Micracanthus and M. marchei have been
proposed for suppression by Tan & Ng (2000) and had been
suppressed by the International Commission on Zoological
Nomenclature in 2003.

Vierke (1991), however, suggested the recognition of three
species groups based almost entirely on ethological charactersand overall body shape, viz. the B. splendens group (B.
persephone, B. tussyae, B. coccina, B. bellica, B. smaragdina,
B. imbellis, B. splendens, B. foerschi), the B. pugnax group(B. taeniata, B. picta, B. edithae, B. climacura, B. pugnax,
B. anabatoides) and the B. macrostoma group (B.
unimaculata, B. patoti, B. macrostoma). The first groupconsists of supposed bubblenest brooders, except for the oralbrooding B. foerschi. The second and third groups consist oforal-brooders only.

Parophiocephalus was established by Popta (1905) for P.
unimaculatus. Regan (1910) treated Parophiocephalus as asynonym of Betta but did not provide reasons. There is no
doubt, after our examination of a large series of preserved
and fresh B. unimaculata that Parophiocephalus is asubjective junior synonym of Betta. Richter’s (1981) proposalof restricting Betta to bubblenest building species and torecognise a new genus, Pseudobetta, for oral-brooding species
is nomenclaturally unsound. The type species for Betta is B.
picta, which is an oral brooder; as is B. pugnax, the designatedtype species of Pseudobetta. Moreover, oral-brooding andbubblenest brooding traits are probably not evolutionary
stable characters in Betta, and may have been selected for
repeatedly in different species (Roberts, 1989) and within
Betta (Rüber et al., 2004). In any case, Richter’s proposal
has not been adopted by most authors.

Witte & Schmidt (1992) tentatively recognised 15 species
groups based on easily recognisable behavioural andmorphological characters, and to a large extent on details of
the head and body patterns which are usually diagnostic.
These are the B. unimaculata, B. patoti, B. edithae, B. picta,
B. akarensis, B. anabatoides, B. pugnax, B. species E, B.
splendens, B. coccina, B. foerschi, B. macrostoma, B.
dimidiata, B. bellica and B. rubra groups. The last four groups
consisted of a single taxon each. In particular, Witte &Schmidt (1992: 318-319) highlighted the B. coccina groupas having a monophyletic origin, using a partly resolved
cladogram of 28 characters and with other Betta groups as
outgroups.
Ng & Kottelat (1994) subsequently separated out the B. waseri
group from the B. anabatoides group of Witte & Schmidt(1992) on the basis of morphological, meristic and colour
characters.

SPECIES GROUPS
The term ‘species group’ is used here for an assemblage of
species sharing a set of diagnostic characters and which may
or may not be a monophyletic lineage. In some cases,
available data may support the monophyly of a given group(e.g. the B. coccina group, fide Witte & Schmidt, 1992), while
for other groups such data are (still) missing or have to be
re-evaluated. Our current inability to demonstrate monophyly
does not automatically imply that a given group is notmonophyletic.

In this paper, the species groups are a modification of Witte& Schmidt’s (1992) version as well as Ng & Kottelat’s (1994)
additional group, together with some alterations. The 13
recognised species groups are discussed below. The species
found in the present study area are indicated with an asterisk(*).
Betta bellica group

Several authors have recognised species groups in Betta based
mainly on superficial similarities, behavioural (breeding andbrood care mainly) and morphological characters. These are,however, not necessarily apomorphic character states.

We recognise a new species group, the B. bellica group,characterised by: long and slender body which has the dorsal
and ventral margins of the body almost parallel, body depth

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Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei
Table 2. Meristic and morphometric data of Betta bellica

Vertebrae
Standard length (mm)Total length (mm)Head length (mm)
MERISTICS
Anal-fin rays
Dorsal-fin rays
Caudal-fin rays
Pelvic-fin rays
Pectoral-fin rays
Subdorsal scales
Transverse scales
Lateral scales
Predorsal scales
Postdorsal scales
MORPHOMETRICS
In % SL
Total length
Predorsal length
Preanal length
Head length
Body depth at dorsal-fin origin
Pelvic fin length
Anal-fin base length
Dorsal-fin base length
Height of anal fin
Distance between
pelvic- and anal-fin origins
In % HL
Orbit diameter
Postorbital lengthInterorbital distance

Betta bellica

Betta fasciata

NEOTYPEZRC 39196
2+8+22 (32)
51.4
68.8
13.6

LECTOTYPEBMNH 1889.12.26:30
2+8+23 (33)
69.1
16.1

I, 29
I, 12
ii, 5+6, ii
I, 1, 4
13
9
9
33
26
8

II, 300, 11
ii, 5+6, ii
I, 1, 4
13
7
9
35
27
9

133.9
63.0
38.9
26.5
23.9
32.7
58.9
18.7
14.4

65.3
37.0
23.3
23.9
19.8
60.5
16.2
-

10.5

13.2

30.1
48.5
33.1

26.1
50.9
32.9

elongated, caudal fin lanceolate, male may exhibit caudal-fin
transverse bars. All members are male oralbrooders.

23-28% SL; short blunt head, head length 23-28% SL; analfin rays 30-33, dorsal-fin rays 11-13; total vertebrae 32-34;
body dark brown with iridescent green patch on every scale.
The members of the B. bellica group are bubblenest builders.

Witte & Schmidt (1992: 325) did not define this group and merely
stated the reported localities in which the member species can befound. They also established a “Betta sp. E” group which they
characterised by having no chin-bar; body depth at dorsal-fin origin
2.8-3.1 times in SL; head length 2.9-3.1 times in SL, head without
light spots; body length up to 72-92 mm SL; pelvic fin with very
long, bright blueish-white filamentous ray; adult male with brightiridescent opercle and throat and light blue body; and occurring in
Sumatra (Jambi), Kalimantan Barat (Kapuas basin) and Sarawak.
These characters are too general and we can discern no clear
evidence to recognise their Betta sp. E group and we consider their
Betta sp. E group as part of the B. pugnax group, along with B.
fusca. All species share the common feature of having a green or
blue iridescent opercle in life, a lanceolate caudal fin, and pointed
anal and dorsal fins.

Witte & Schmidt (1992: 325) defined B. bellica as having a total
anal-fin ray count of 30-34; some posterior rays of dorsal and anal
fins branched; caudal fin somewhat rhomboid with a few long,
filamentous principal rays and TL up to 110 mm. Tan & Ng (1996)
described an allied species (B. simorum) from Jambi, Sumatra.
Subsequently, B. simorum had been found in the lower Kapuas basinin Kalimantan Barat (pers. observ.).
Included species: B. bellica*, B. simorum.

Betta pugnax group
Members of the B. pugnax group are distinguished by the following
set of characters: a relatively large head, 28-40% SL; usually brownwith greenish to blueish iridescent spots on body scales when alive;
male with strongly coloured (blue, green or greenish blue) iridescent
opercle scales, occasionally even covering anterior portion of belly
in life; juvenile and female usually with a light brown backgrounddisplaying a central and a second central stripe, with a caudal
peduncle spot; head rhombic when viewed from above, with orwithout chin-bar; and fins of mature specimens pointed, often

Included species: B. pugnax*, B. fusca, B. schalleri, B. prima, B.
enisae, B. pulchra*, B. breviobesus, B. stigmosa*, B. lehi*.

Betta akarensis group
Members of the B. akarensis group are distinguished by thefollowing character suite (see Tan & Kottelat, 1998a): presence ofpreorbital and postorbital stripes (postorbital stripe may be

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
Table 3. Meristic and morphometric data of Betta stigmosus and B. lehi.

Standard length (mm)n
MERISTICS (mode)
Vertebrae
Anal fin rays
Dorsal fin rays
Caudal fin rays
Pelvic fin rays
Pectoral fin rays
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 stigmosus

Betta lehi

31.2-41.1
5

35.9-60.8
19

10 + 18-19 (total 28-29, 29)
I-II, 21-23 (total 23-24, 24)
0-I, 8-9 (total 9-10, 9)
ii, 5 + 6, ii
I, 1, 412-13 (13)61/2
91/2-10 (10)29-30 (29)
14-15 (14)
6-7 (7)20-21 (20)
10-11 (10)10 + 18-20 (total 28-30, 29)
II, 23-27 (total 25-29, 28)
I, 8-10 (total 9-10, 10)ii, 5 + 6, ii
I, 1, 412
5-61/2 (6)91/2
28-30 (29)20-23 (22)10-11 (11)141.5-148.0
62.8-67.5
21.2-23.6
14.4-17.3
47.2-52.8
33.6-36.5
25.6-28.2
33.2-51.8
48.2-51.3
12.2-13.9
64.4-69.1
19.0-21.5

136.2-152.7
62.6-67.5
47.4-52.5
33.4-36.8
27.3-32.5
34.8-50.6
48.1-54.2
12.3-15.6-

26.7-28.4
47.6-51.1
29.8-32.1
22.0-25.2
56.0-58.9

27.2-32.2
42.2-48.5
31.7-37.4
22.8-27.9
-

52-60
62-65

59-74
70-84

51.7-55.4

51.2-56.1

interrupted or faint), and a chin-bar; caudal fin lanceolate, medianrays distinctly elongated in mature male, with branching of caudalfin rays starting from ca. 40 mm SL, usually with caudal-fin
transverse bars (always distinct in mature male); greenish to blueish
iridescent body scales in some species; opercle without iridescentscales (fide Witte & Schmidt, 1992; Ng, 1993). Adult specimens
may possess greenish-gold opercle scales which tend to disappear
in captivity. Juvenile may possess gold opercle scales, but thesedisappear with age. The faintly coloured greenish-gold opercle scalesof the adults could be due to stress from handling. All members are
male oralbrooders.

Betta unimaculata group
Members of the B. unimaculata group are characterised by thefollowing unique combination of characters: long, relatively slenderbody (body depth at dorsal-fin origin 18-25% SL); large blunt head
(head width 19-24% SL); long maxilla and lower lip (distance fromtip of lower jaw to posterior end of maxilla 27-54% HL); caudal fin
rounded, occasionally with elongated median rays; short filamentous
pelvic fin; relatively pointed dorsal and anal fins.

The B. akarensis and B. pugnax groups can be readily differentiatedby two characters: the presence of faintly coloured greenish-gold
opercle scales in members of the B. akarensis group (vs. the intense
blueish-green opercle scales in members of the B. pugnax group)
and usually more anal-fin rays (28-30, vs. 25-28).

Members of the B. unimaculata group differ from the other groups
in breeding behaviour. The breeding display consists of an open
gape (unique to this group) and flaring of dorsal, pelvic, anal andcaudal fins (vs. expansion of branchiostegal membranes with fins
flaring). The female of some members do not participate in the egg
gathering or egg transfer to the male (van den Nieuwenhuizen, 1995;
Tan, 1997).

Included species: B. akarensis*, B. balunga*, B. chini*, B. pinguis,
B. ibanorum*, B. aurigans, B. obscura.

Included species: B. unimaculata, B. macrostoma*, B. patoti, B.
ocellata*, B. gladiator*, B. pallifina.

53

Dorsal-fin rays
Pectoral-fin rays
Sub-dorsal scales
Lateral scales
Lateral scales below dorsal-fin origin
Lateral scales above anal-fin origin
Predorsal scales
Postdorsal scales
MORPHOMETRICS
In % SL
Total length
Head length
Body depth at dorsal-fin origin
Predorsal length
Preanal length
Postdorsal length
Anal-fin base length
Dorsal-fin base length
Pelvic-fin length
Lower jaw length
Height at pectoral-fin origin
In % HL
Orbit diameter
Postorbital lengthInterorbital width
Lower jaw length
Height at pectoral-fin origin

Number of specimens
MERISTICS (mode)
Anal-fin rays
0-III, 27

I-II, 26-28(total 28-30, 28)
I, 7-8 (total 7-9, 8)
12-13 (13)
5-6 (5)31-33 (32)
16-18 (17)6-8 (6)22-25 (23)10-12 (11)54132.8-141.3
32.6-35.5
28.0-31..5
66.7-71.7
49.2-52.2
22.6
47.8-55.4
13.7-14.1
35.0-40.6
10.2
16.1
29.5-33.3
46.7-49.2
28.8-35.3
28.8
45.5

134.4-154.2
29.8-34.9
25.1-30.9
64.3-70.0
43.4-48.5
19.9-25.1
49.6-56.8
9.9-14.1
30.0-50.0
8.8-10.7
15.7-19.6

24.5-32.3
45.7-51.6
29.4-37.4
25.4-34.4
47.3-61.9

0-II, 8-9
12
6
29-30
15-16
6
21-23
9-10

2

Betta balunga

18

Betta akarensis
8

Betta chini

26.2-30.8
46.7-52.5
29.3-37.7
25.8-31.5
47.6-59.3

137.2-144.6
31.4-33.3
26.3-29.0
64.5-69.8
42.6-45.7
19.9-23.3
54.8-58.5
11.0-14.0
30.3-45.8
8.9-11.3
16.9-21.5

II, 24-28(total 26-30, 28)
I, 7-9 (total 8-10, 8)
12-13 (12)51/2-7 (6)30-31 (31)16-17 (16)
5-7 (6)22-24 (23)9-11 (10)Table 4. Meristic and morphometric data of Betta akarensis, B. balung, B. chini, B. ibanorum and B. obscura.

22.2-30.5
45.7-50.2
29.6-37.7
28.0-36.1
44.7-59.9

138.2-165.6
31.7-36.4
23.3-28.1
64.5-68.5
45.8-49.4
18.9-24.2
50.0-55.8
10.6-14.8
28.5-50.6
9.3-12.0
15.0-21.9

I-II, 25-27(total 27-29, 28)
I, 8-10 (total 9-10, 9)
13-14 (13)
5-6 (51/2)
30-32 (30)
15-17 (16)
6-7 (7)
21-23 (22)10-11 (11)16

Betta ibanorum

21.1-25.3
45.0-50.0
29.9-37.1
28.9-34.1
49.5-61.6

135.5-160.2
30.9-35.4
28.7-32.7
63.8-68.3
46.1-50.7
19.8-23.9
51.3-56.4
12.1-15.4
33.1-52.6
9.2-11.3
16.5-20.4

II-III, 24-27(total 26-29, 28)0-II, 8-9, (total 8-10, 10)
10-13 (11, 12)
51/2-7 (61/2)
28-30 (29)
14-15 (15)6-71/2 (7)19-21 (20)
10-11 (11)14

Betta obscura

Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei

Pectoral-fin rays
Subdorsal scales
Body depth at dorsal-fin origin
scalesLateral scales
Lateral scales below dorsal-fin
originLateral scales above anal-fin
originPredorsal scales
Postdorsal scales
MORPHOMETRICS
in % 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
Height at pectoral-fin origin
Head width
in % HL
Orbit diameter
Postorbital lengthInterorbital width
Lower jaw length
Head width
Height at pectoral-fin origin

Anal-fin rays

No. of specimens
MERISTICS (mode)
Dorsal-fin rays

55

24
10-11 (11)21-24 (23)10-11 (11)21.6-24.8
51.3-56.0
28.7-33.0
27.3-33.1
63.3-74.2
50.0-57.7

6-8 (7 or 8)

7-11 (8)19.8-26.4
49.3-56.7
29.2-35.8
38.0-54.0
57.4-70.2
34.1-49.6

32-34 (33)
18-20 (18 or 19)

32-33 (33)
15-17 (17)131.9-135.6
68.8-71.9
20.3-22.5
14.8-17.0
44.8-46.8
31.4-33.3
21.6-24.5
24.0-32.1
52.2-54.5
10.5-11.3
16.7-18.7
21.0-24.0

0-I, 7-8 (total 8,
mode 8)
I-II, 28-30 (total 30-31,
mode 30)
12-13 (13)5-51/2 (51/2)
91/2

0-I, 9-11 (total 9-11,
mode 10)0-I, 24-27 (total 25-28,
mode 27)
13 or 14
6-8 (7)
91/2-10 (91/2)

131.0-137.0
62.3-66.7
18.2-22.0
12.9-17.3
44.7-50.3
30.4-33.3
18.5-21.1
21.6-27.6
43.2-49.2
12.2-15.7
10.6-16.0
17.7-22.4

13

Betta ocellata

14

Betta macrostoma

21.4-27.4
49.7-56.3
30.4-35.4
31.6-37.9
61.3-69.6
42.7-56.5

130.5-139.7
67.7-73.0
20.2-25.5
13.3-16.4
43.9-51.3
29.0-33.3
18.9-24.7
20.4-32.4
45.9-53.7
7.9-10.4
12.4-18.6
19.6-21.8

24-26 (24)10-11 (11)7-9 (7)31-33 (32)
17-20 (18)0-I, 6-8 (total 6-8,
mode 8)
I-II, 26-30 (total 27-30,
mode 28 or 29)
12-13 (12)4-5 (5)9-91/2 (91/2)

13

Betta unimaculata

20.5-24.8
48.6-53.3
30.8-40.1
32.9-38.5
57.9-67.2
40.6-53.7

132.3-143.0
67.5-69.9
19.3-21.8
14.3-16.5
41.1-46.1
29.7-31.4
20.0-22.6
25.9-34.7
51.0-56.3
9.4-11.4
12.7-16.6
18.0-20.7

24-26 (25)11-12

22.6-28.3
46.3-53.5
29.5-38.3
28.3-38.3
64.5-77.3
35.2-57.3

135.4-139.2
68.0-72.8
17.2-22.2
13.5-15.3
43.4-47.2
28.5-31.3
18.4-20.4
29.0-35.7
48.1-52.8
7.3-11.4
10.6-16.5
19.2-23.4

24-27 (26)10-12 (11)

6-8 (8)34-36 (35)
19-21 (19, 20)

33-341/2 (34)
18-19
5-8 (7)0, 7-9 (total 7-9,
mode 8)
I-II, 27-30 (total 28-31,
mode 30)
16-17 (17)5-61/2 (6)1
9 /2 -10 (91/2)

16

Betta gladiator

0-I, 7-9 (total 8-9,mode 8)I, 29-32 (total 30-33,
mode 31)12-14 (13)6-61/2 (6)91/2-108

Betta patoti

Table 5. Meristic and morphometric data of Betta macrostoma, B. ocellata, B. unimaculata, B. patoti, B. gladiator and B. pallifina.

19.0-24.8
50.3-55.5
30.4-36.9
30.6-39.6
56.4-70.2
37.6-59.5

129.7-140.9
64.6-69.4
19.1-23.3
13.2-16.2
39.6-50.5
26.8-33.4
20.5-23.0
22.5-34.7
47.4-57.7
9.3-12.5
12.1-16.0
17.7-20.8

21-26 (23)10-13 (11)

6-8 (6)32-34 (33)
16-19 (18)0-I, 6-9 (total 7-9,
mode 8)
I-II, 26-31 (total 27-32,
mode 30)
11-14 (13)5-61/2 (6)9-10 (91/2)

25

Betta pallifina

THE RAFFLES BULLETIN OF ZOOLOGY 2005

Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei
Table 6. Meristic and morphometric data of Betta taeniata

Number of specimens
MERISTICS (mode)
Vertebrae
Anal-fin rays
Dorsal-fin rays
Pelvic-fin rays
Pectoral-fin rays
Caudal-fin rays
Pelvic-fin length to anal-fin ray
Subdorsal scales
Body depth at dorsal-fin origin
Lateral scales
Lateral scales below dorsal-fin origin
Lateral scales above anal-fin origin
Predorsal scales
Postdorsal scales
MORPHOMETRICS
In % SL (mean)
Total length
Predorsal length
Postdorsal length
Preanal length
Head length
Body depth at dorsal-fin origin
Pelvic-fin length
Anal-fin base length
Dorsal-fin base length
Caudal peduncle depth
In % HL (mean)
Orbit diameter
Postorbital lengthInterorbital distance

Betta taeniata
type material

Betta taeniata
fresh material

48

2 + 8 + 18 (total 28)(n=3)
total 23-24
total 9I,1,4
11-12 (12)ii,5+7,i
5-7 (6)5-6 (6)ca. 9
ca. 25-26 (26)
ca. 12-14 (12)ca. 6-8 (7)19
9-10 (9)2 + 8 + 17-19 (total 27-29,
mode 28) (n=8)
total 23-26 (26)
total 8-10 (10)I,1,4
12ii,5+6-7,i (ii,5+7,i)
5-6 (6)5-6 (6)
9-91/227-29 (28)13-14 (13)7
19-20
9-10 (10)133.7-140.3 (137.3)
65.9-70.4 (67.5)
21.4-23.4 (22.4)
49.1-53.3 (51.8)
33.5-37.6 (35.6)
24.4-30.4 (27.4)
26.5-32.5 (29.4)
43.8-51.8 (48.9)
11.2-15.4 (14.0)
15.3-17.1 (16.4)

132.9-139.9 (135.7)
62.4-66.5 (64.6)
20.3-25.5 (22.5)
49.5-54.1 (51.8)
30.3-36.1 (33.0)
24.8-28.9 (26.7)
23.5-28.4 (25.9)
49.4-53.2 (51.0)
10.8-15.8 (13.0)
15.2-16.7 (15.9)

27.2-31.0 (29.1)
45.5-46.5 (46.0)
24.6-28.6 (26.4)

23.8-30.4 (27.1)
41.1-48.1 (45.9)
26.9-36.2 (31.2)

body scales (other colours for domesticated stocks of B. splendens);
dorsal-fin rays 0-II, 7-9; anal-fin rays II-V, 21-26.

Betta picta group
Members of the B. picta group are characterised by: roundedunpaired fins (anal fin occasionally bluntly pointed); anal-fin rays
I-III,18-24; dark pigmented distal margin on anal and caudal fins,especially in mature male (Fig. 4); preorbital and postorbital stripes
present, throat with chin-bar; opercle with iridescent gold, green or
blue scales (after Witte & Schmidt, 1992). All members are male
oralbrooders.

Included species: B. splendens, B. smaragdina, B. imbellis*, Betta
stiktos.

Betta coccina group
The B. coccina group is unique in having a small adult size (lessthan 40 mm SL), either a uniform red or black body colour, midlateral body blotch present in some species (see Key) (Fig. 6a) andnine abdominal vertebrae (vs. 10-12 in all other species groups)(Witte & Schmidt, 1992: 325). All members of the group are
stenotopic inhabitants of peat swamp forests. They typically inhabit
the shallow pools of submerged leaf litter. All are bubblenest
brooders.

Included species: Betta picta, B. taeniata*, B. simplex, B. falx.

Betta splendens group
Members of this group were defined by Witte & Schmidt (1992:
325-327) as having a small head (22-31% SL); often brightly
coloured body; iris of eye with iridescent green or blue patches;
bubblenest brooders; body elongate or slender; head with parallel
opercles when viewed dorsally; red or brown anal and caudal fin
rays contrasting with iridescent interradial membranes (especiallyin male); unpaired fins without fine iridescent margin; in male,
opercle with red or iridescent blue bars; TL up to 70 mm, SL up to
57 mm; iridescent green opercle scales (B. imbellis, B. smaragdina)or red parallel bars on opercle (B. splendens); distinct red distal
crescent on the caudal fin (in B. imbellis) (Fig. 5a); iridescent green

Included species: B. coccina*, B. tussyae*, B. persephone*, B.
rutilans, B. brownorum*, B. livida*, B. miniopinna, B. burdigala.

Betta waseri group
The B. waseri group is diagnosed by: black mark(s) (various speciesspecific shapes) on throat and black lower lip (Fig. 3); distinctive
markings on opercle; large adult size (more than 60 mm SL); clayyellow body colour; and golden iridescent belly and opercle scales

56

THE RAFFLES BULLETIN OF ZOOLOGY 2005
Table 7. Meristic and morphometric data of the Betta splendens, B. imbellis, B. smaragdina and B. stiktos..

Number of specimens
MERSITICS (mode)
Anal fin rays
Dorsal fin rays
Pectoral fin rays
Subdorsal scales
Transverse scales
Lateral scales
Predorsal scales
Postdorsal scales
Lateral scales below dorsal-fin origin
Lateral scales below anal-fin origin
MORPHOMETRICS (% SL)
Total length
Predorsal length
Postdorsal length
Preanal length
Head length
Body depth
Caudal peduncle depth
Pelvic fin length
Anal fin base length
Dorsal fin base length(% HL)
Orbit diameter
Postorbital lengthInterorbital width
Snout length

B. splendens

B. imbellis

B. smaragdina

B. stiktos

7

24

5

5

28-30 (30)
10-14 (11)11-12 (11)7-10 (7, 8)
9-91/2 (91/2)
29-31 (29, 31)
22-26 (25)
8-11 (9)14-16 (14)5-7 (6)25-31 (27)9-12 (10)11-12 (12)51/2 -8 (7)9-91/2 (91/2)
30-32 (31)21-24 (23)9-12 (11)15-17 (16)
6-8 (7)26-29 (29)
9-11 (10)12
6-7 (6)91/2 -10 (91/2)
29-32 (30)22-24 (23, 24)
10-11 (10)15-16 (15)
5-7 (6, 7)

27-29 (29)
10-11 (10)1261/2-7 (7)91/2-10 (91/2)
33-34 (33)
24-25 (25)
10-11 (10)16
6-7 (6)128.4-136.6
60.1-65.6
15.3-22.8
40.5-45.3
28.2-30.9
27.1-32.2
18.6-20.3
26.1-63.9
56.8-58.8
16.7-23.8

131.9-141.7
61.3-66.7
17.3-22.1
38.7-45.8
25.8-30.9
24.2-30.1
15.9-19.2
31.6-51.0
54.2-61.9
12.9-18.9

132.3-140.4
63.9-66.4
21.4-23.2
42.3-45.1
27.8-30.1
24.7-28.4
18.1-19.3
36.8-51.0
54.2-60.0
14.9-16.7

136.0-139.7
63.5-66.4
17.5-20.1
40.5-43.5
27.7-30.3
26.2-27.7
15.4-17.4
33.2-39.0
53.7-60.3
15.2-15.6

22.7-26.6
49.6-52.7
30.2-34.3
21.1-26.9

24.7-31.1
43.6-55.2
25.6-36.0
15.6-28.2

25.0-29.0
48.8-51.2
31.0-32.5
17.5-23.0

27.7-29.7
45.9-50.0
26.4-31.6
16.9-21.6

in some species (Ng & Kottelat, 1994). An additional diagnostic
character is the colour zonation of the eye (Fig. 3h). The pupil issurrounded by an iridescent yellow rim, followed with a reddish
iris rimmed with another thicker iridescent yellow ring on thecircumference of the orbit. This is most obvious in juveniles andsub-adults. Eyes of old or large specimens are darker and the colour
zonation may not be so obvious. The throat markings of each of thespecies of the B. waseri group are diagnostic (Fig. 3). They are only
displayed when specimens are disturbed, well acclimatised or inbreeding condition. Well-acclimatised specimens invariably display
a brown body colour, with a black or brown postorbital and central
stripe which runs from just behind the eye to the middle of the caudal
peduncle. A distinct broken black stripe runs just above the anal fin
sheath scales.

Included species: B. waseri*, B. chloropharynx, B. hipposideros*,
B. tomi*, B. spilotogena, B. renata, B. pi*.

Betta edithae group
The B. edithae group is characterised by: branchiostegal membraneand posterior part of opercle translucent or opaque; and size up to
62 mm SL. Witte & Schmidt (1992: 326) commented that they had
seen specimens resembling B. edithae from Malay Peninsula.
Despite extensive collections in Peninsular Malaysia, we have notcome across any specimens resembling B. edithae yet. Also, there
are no preserved museum specimens from this area which can bereferred to this species group.
The group currently has a single member, B. edithae, known fromseveral widely separated areas: Kalimantan Barat, Kalimantan
Tengah, Kalimantan Selatan, Kalimantan Timur (Roberts, 1989;
Witte & Schmidt, 1992), Palembang (unpublished data), Banka(unpublished data), Biliton (unpublished data) and Pulau Bintan (Tan
& Tan, 1994). Whether all these populations are conspecific has
yet to be determined. Witte & Schmidt (1992: 324) listed Jambi,
Sumatra as another locality (B. (edithae) sp. B), however, these arelikely to be B. falx (Tan & Kottelat, 1998b) instead.

All the species in the B. waseri group are paternal oral-brooders.
They exhibit a ‘S’ mating embrace. The male curves its body aroundthe female, positioning its genital opening next to the female’s inorder to fertilize the eggs as they are extruded. The embrace of themale and female resembles a ‘S’ shape. The female recovers fromthe embrace first and collects the fertilized eggs from the cup formedby the curved anal fin of the male. The female spits the eggs out
one at a time in front of the male. The male then collects the egg
quickly, otherwise the female gulps the egg back. This egg gulping
ritual is repeated until the male has collected all the eggs in its buccal
sac. At this stage, the male will have a grossly enlarged throat. The
above mating ritual had been observed for B. spilotogena and there
can be as many as 180 fry in the mouth (Tan & Tan, 1994; pers.
observ.). An apparently undescribed species of the B. waseri groupfrom Pekan Nanas (misidentified as B. macrophthalma by Schmidt,
1988) also possesses this form of behaviour. Unfortunately, we do
not have sufficient material to resolve the identity of this species.

Betta foerschi group
The B. foerschi group was defined by Witte & Schmidt (1992: 325)
as having iridescent blue or green unpaired fins, with rays andinterradial membranes not contrasting; dorsal fin with thin iridescent
white margin; opercle with two distinct reddish-yellow bars; andsize up to 51 mm SL.

57

Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei
The position of B. rubra requires comment. Witte & Schmidt (1992)
defined B. rubra as having a triangular mark below the eye; foundin the lowlands of northwestern Sumatra; and size up to 38 mm SL.
They accomodated this taxon into its own species group based on
the above mentioned characters. We tentatively place B. rubra in
the B. foerschi group on the basis of the following features: presenceof two vertical lighter coloured bars on opercle; vertical bars on
body when preserved (also present, but faint in B. strohi); and generalbody shape. Until fresh material becomes available, the relationshipsof B. rubra can only be speculated. However, considering the known
characters, there is little reason to treat B. rubra as a distinct species
group. The prime character used by Witte & Schmidt (1992: 325)
to delineate the ‘B. rubra’ group was the triangular patch beneaththe eye, which is a weak character as this mark is part of the chinbar extending to the opercle. In head and body form, B. rubra isclearly either a member of the B. foerschi or B. splendens group.
These two species groups are very close morphologically. From thedistinctiveness of the two vertical bars on the opercle and stockinessof the head of B. rubra, it fits better in the B. foerschi group as
currently understood.

fin origin almost above anal-fin origin; and small size (all known
specimens less than 30 mm SL); and a unique colour pattern (fide
Kottelat & Ng, 1994). Paternal oral-brooder (Grams & Dickmann,
1997).
Included species: B. albimarginata, B. channoides.

Betta dimidiata group
The B. dimidiata group includes a single species. Witte & Schmidt(1992: 323) defined it as having: elongate body, body depth 2125% SL; pale interspace between upper and central stripe thin, about
half a scale apart, appearing like a thin stripe; largest known size
35 mm SL; 22-26 articulated anal-fin rays. This species certainly
deserves its placement in its own group, which can be further
distinguished by the following characters: extreme sexual
dimorphism (observed from captive specimens), male with greatly
elongated unpaired and pelvic fins, bright blueish-white pelvic finreaching up to two-thirds of anal fin, anal fin with iridescent light
blue distal margin reaching up to lower margin of caudal fin; female
with somewhat rounded unpaired fins; throat with chin-bar andopercle with iridescent green scales (in both sexes, more intense inmale), mature male with anterior portion of throat purple, up to
anterior portion of opercle; several vertical black bars on body justbehind opercle edge in both sexes, more prominent in male. Oralbrooder.

Included species: B. foerschi, B. strohi, B. rubra.

Betta anabatoides group
The B. anabatoides group was defined by Witte & Schmidt (1992)
by the following characters: no iridescent opercle scales; filamentous
pelvic ray usually not extended; head with two light spots behind
eye; adult body colour uniform reddish brown or black. The groupis further characterised by: body relatively short and stout (bodydepth 30-33% SL); no chin bar; terminal portion of caudal rays
protruding behind caudal membrane; caudal transverse bars present
or absent; convex head dorsal profile and stout body reminescentof Anabas.

SPECIES ACCOUNTSBETTA BELLICA GROUP
Betta bellica Sauvage, 1884
(Figs. 9a-b, 46a)

The group includes a single species, B. anabatoides, which Witte& Schmidt diagnosed as having no markings on lower jaw or no
throat pattern; central stripe between lower lip and eye with “variable
modifications” [sic, no further elaboration]; size up to 90 mm SL;
found in Kalimantan Selatan, Kalimantan Tengah, Kalimantan Baratand possibly in Sarawak. Bleeker (1851: 269) described B.
anabatoides from Banjermassing (=Banjarmasin), Borneo, on thebasis of two specimens 101 and 115 mm TL. He provided the analfin ray count II, 29; dorsal-fin ray count II, 7; and lateral scale count
33. There was no accompanying figure, but the high anal-fin ray
count, extended fin ray tips and convex head profile allies B.
anabatoides to the B. waseri and B. akarensis groups. It differs,
however, mainly in the shape of the head and anterior part of body.
Vierke (1979b) commented on B. anabatoides, on the basis of fresh
specimens collected from the type locality of B. foerschi - Mentayabasin, 250 km northwest from Banjarmasin. His figure of B.
anabatoides and the figure in plate 75 of Kottelat et al. (1993) depict
specimens that come closest to Bleeker’s description and to its
namesake - Anabas (Anabantidae). To confirm the status of thisspecies group, the types should be examined, together with fresh
topotypic material. The B. anabatoides group is tentatively regarded
as valid and confined to Borneo.

Betta bellica Sauvage, 1884: 217, fig.; Regan, 1910: 779; Weber &de Beaufort, 1922: 363; Herre, 1940: 43 (key only); Kottelat,
1989: 19 (list only); Witte & Schmidt, 1992: 326 (part); Ng etal., 1992: 28; 1994: 210; Kottelat et al., 1993: 161 (part), pl. 75;
Tan & Ng, 1996: 144, fig. 3a, 4a.
Betta fasciata Regan, 1910: 782, pl. 75, Fig. 4; Weber & de Beaufort,
1922: 362; Herre, 1940: 42 (key only); Witte & Schmidt, 1992:
326; Tan & Ng, 1996: 149, fig. 3b.
Material examined. – Neotype - Malaysia: ZRC 39196, 51.4 mm
SL male; Malaysia: Selangor, north Selangor peat swamp forest,
43 km towards Sungai Besar (3°39’12.9"N 101°18’00.4"E); D. S.
L. Chung et al., Sep.1993.
Others - Malaysia: ZRC 39253, 2 ex., 43.2-44.1 mm SL; Aug.1992,- ZRC 39254, 1 ex., 54.4 mm SL; Sep.1992, - ZRC 39255, 3 ex.,43.9-48.3 mm SL; Jul.1993, - ZRC 39256, 1 ex., 49.8 mm SL;
Sep.1993, - ZRC 39262, 3 ex., 39.0-40.7 mm SL; Jul.1993, same
locality data as neotype. — ZRC 39257, 1 ex., 49.4 mm SL; Jul.1993,- ZRC 39258, 7 ex., 39.1-57.6 mm SL; Sep.1993, - ZRC 39263, 1
ex., 48.6 mm SL; Mar.1993; Malaysia: Selangor, north Selangor
peat swamp forest, streams running parallel to road at 46.5 km
towards Sungai Besar (3°39’51.4"N 101°19’43.0"E); D. S. L. Chunget al. — ZRC 39259, 1 ex., 49.4 mm SL; Sep.1992, - ZRC 39260,
1 ex., 54.3 mm SL; Jul.1993, - ZRC 39261, 2 ex., 46.0-49.7 mm
SL; Sep.1993; Malaysia: Selangor, north Selangor peat swamp
forest, stream parallel to road at 47.6 km towards Sungai Besar(3°40’12.7"N 101°20’10.2"E); D. S. L. Chung et al. — ZRC 15059,
1 ex., 56.1 mm SL; Malaysia: Selangor, north Selangor peat swamp
forest, stream at 43 km mark on road to Sungai Besar; 1991-92

Betta albimarginata group
The B. albimarginata group is characterised by: broad head andbody; flat head top; pelvic fin broad and falcate (seldom
filamentous); great number of anal-fin spines (IX-XII vs. 0-IV inother species); few articulated anal-fin rays (11-13 vs. 18-32); dorsal-

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
Honours class, 19 Jun.1991. — ZRC 39577, 2 ex., 39.8-47.7 mm
SL; Malaysia: Selangor, north Selangor peat swamp forest, stream
at 43 km mark on road to Sungai Besar, coll. P. K. L. Ng et al.,Sep.1994. — ZRC 39578, 2 juv.; Malaysia: Selangor, north Selangor
peat swamp forest, road from Sungai Besar to Tanjung Malim, 0.65
km from 35 km stone, coll. P. K. L. Ng et al., Sep.1994. — ZRC
38274, 1 juv.; Malaysia: Selangor, north Selangor peat swamp forest,
coll. P. K. L. Ng et al., Sep.1994. — ZRC 15053-54, 2 juv.;
Peninsular Malaysia, north Selangor peat swamp forest, stream at
34 km mark on road to Tanjung Malim; NUS 1991-92 Honours
class, 17 Jun.1991. — ZRC 16700, 1 juv.; - ZRC 15055-58, 4 juv.;
Malaysia: Selangor, north Selangor peat swamp forest, stream at
50 km mark to Tanjung Malim (United Plantations Berhad); NUS
1991-92 Honours class, 18 Jun.1991. — ZRC 17374-76, 3 juv.;
Malaysia: Selangor, north Selangor peat swamp forest, coll. P. K.
L. Ng, Aug.1991. — ZRC 39195, 1 ex., 47.7 mm SL; Malaysia:
Selangor, north Selangor peat swamp forest, stream running throughedge of United Plantations; D. S. L. Chung et al., Sep1993. — ZRC
28636-37, 2 ex., 48.2-58.0 mm SL; Malaysia: Johor, Segamat;
aquarium collectors, Oct.1992. — ZRC 29126, 1 ex., 39.5 mm SL;- ZRC 29052, 1 juv.; Malaysia: Johor, Pontian, Sri Bunian, Kampung
Pt. Tekong, blackwater stream, coll. P. K. L. Ng et al., 8 May.1992.
— ZRC 28799-800, 2 juv.; Malaysia: Johor, near Pontian,Tenggoyon (between Pekan Nanas and Kukup), coll. P. K. L. Ng
et al., 4 Mar.1992. — ZRC 21949-50, 2 juv.; Malaysia: Johor, 56
km Pontian-Johor Bahru road, blackwater stream, coll. K. K. P. Lim& D. S. L. Chung, 14 May.1992. — ZRC 17377-79, 3 juv.; Malaysia:
Johor, about 2 km north of Ayer Hitam, blackwater stream, coll. P.
K. L. Ng & K. K. P. Lim, 23 May.1991. — ZRC 17373, 1 juv.;
Malaysia: Johor, about 2 km north of Ayer Hitam, in swamp forest,
coll. P. K. L. Ng & K. K. P. Lim, 23 May.1991. — ZRC 2867677, 2 juv.; Malaysia: Pahang, pool along Mersing-Pekan road (73km to Kuantan, 400 m to Sungai Beto), coll. M. Kottelat & P. K.
L. Ng, 9 Mar.1992. — ZRC 25674-76, 3 juv.; Malaysia: Pahang,
69 km, Mersing-Kuantan road, blackwater stream, coll. P. K. L. Ng
et al., 10 Mar.1992. — ZRC 29408, 1 ex., 42.8 mm SL; Malaysia:
Pahang, Pekan road, 66 km from Mersing to Kuantan road,
blackwater stream, coll. P. K. L. Ng et al., 24 Jul.1992. — ZRC
38049, 1 ex., 77.3 mm SL; Malaysia: Selangor, Serbak Bernam,
coll. P. K. L. Ng, Jun.1991. Sumatra: BMNH 1889.12.26:30,
lectotype of B. fasciata, 69.1 mm SL; Indonesia: Sumatra, North
Sumatra, Deli (Medan); Iversen. — ZRC 42468, 10 ex., 15.3-40.1mm SL; Indonesia: Sumatra, North Sumatra, ca. 80 km south ofMedan, Sei Rampah, Pantai Berdagai, pond at Kampung Nagur(3°31’14.0”N 99°10’32.2”E), coll. H. H. Tan et al., 15 Jun.1996.

— ZRC 42478, 4 ex., 18.8-24.3 mm SL; Indonesia: Sumatra, North
Sumatra, ca. 250 km south of Medan, Rantau Berapat, tributary ofSungai Semang Alam (2°27’06.4”N 99°43’03.0”E), coll. H. H. Tan
et al., 16 Jun.1996.

Diagnosis. – Betta bellica differs from B. simorum (the only
other member of the species group) in the following
characters: lower mode of anal-fin rays (31, vs. 32); lower
mode of dorsal-fin rays (12, vs. 11); lower mode of lateralscales (33, vs. 34); lower mode of postdorsal scales (8, vs.
9); shorter pelvic fin (length 23.6-38.8% SL, vs. 31.3-48.3;
reaching base of 8th anal ray, vs. 14th); shorter pelvic finflange (ends halfway down pelvic fin filament, vs. two-thirdsway); adpressed pelvic-fin not reaching anal-fin origin (vs.
overlapping); and a less sloping lateral head profile (vs. more
sloping and slightly concave).
Description. – General body shape and appearance as shownin Figs. 9a-b, 46a. Body long, slender, body depth at dorsalfin origin 23.7-26.2% SL; head length 26.0-28.0% SL; headblunt and small; opercle without iridescent scales; body brownwith green iridescence on scales; caudal fin rounded withmedian rays extended, dorsal and anal fins pointed, pelvic
fin filamentous, flange (a narrow membrane) on first simple
pelvic fin ray ending abruptly before one-third of ray.
Meristics: lateral scales 33-35; transverse scales at dorsal-fin
origin 81/2-9; predorsal scales 24-27; subdorsal scales 7-10;
postdorsal scales 7-9; dorsal-fin rays 0-I, 11-12 (total 12-13,
mode 12); caudal fin rays ii, 5+6, ii; anal-fin rays I-II, 29-31(total 30-32, mode 31); pelvic fin rays I, 1, 4; pectoral fin
rays 12-13; vertebrae 2 + 8-9 + 22-23 = 32-33 (mode 33,n=10). Meristic and morphometric data of neotype in Table
2. Maximum known size 77.3 mm SL (ZRC 38049).
Coloration. – For live coloration, see Fig. 46a (see also Linke,
1991: 28; Kubota et al., 1996: 28). Background body colour
dark brown dorsally, lighter brown ventrally. Fins withwhitish to golden distal margins, brownish with some greenish
iridescence in mature specimens when alive. Dorsal fin withtransverse bars. Pelvic fin with white tip. Male with more
iridescent green scales than female. Eye with bright blue
iridescence. Juvenile with distinct central stripe.
Distribution. – Widely distributed in Peninsular Malaysia:
Terengganu, Perak, Selangor, Pahang and Johor. Also foundnear Medan, North Sumatra (Fig. 10).

ab

Field notes. – Betta bellica is usually found in peat swamp
habitats, although it can also be found in swamp forest habitats(Ng et al., 1992). Other syntopic Betta species are: B. tussyae,
B. waseri (Pahang), B. hipposideros, B. livida (Selangor), B.
imbellis, B. persephone, B. pulchra (Johor). Gut contents ofwild caught specimens include a relatively high proportionof odonate nymphs (Chung et al., 1994).

Fig. 9. a) Betta bellica – ZRC 39196, neotype, 51.4 mm SL, b) B.
fasciata - BMNH 1889.12.26:30. lectotype, 69.1 mm SL.

Betta bellica is the largest known bubblenest brooding Betta.
Details of its spawning are given in Vierke (1987, 1991) andvan den Nieuwenhuizen (1993). Mohsin & Ambak (1983:
249) listed B. bellica as rare or extinct, but this is incorrect,
as recent collections show. Their statement probably reflects
the poor knowledge and lack of sampling in peat swamp

59Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei
101; Weber & de Beaufort, 1922: 357; Herre & Myers, 1937:
72; Fowler, 1938: 121; Herre, 1940: 46.
Betta pugnax - Günther, 1861: 389; Hanitsch, 1901: 4; 1912: 27;
Duncker, 1904: 164; Breder, 1934: 128; Tweedie, 1952b: 73;
1953: 50, pl. 9, fig. 5; Alfred, 1958: 161; 1961a: 162; 1961b:
18; 1963: 147, fig. 2; 1966: 49, pl. 8, fig. 1; Moshin & Ambak,
1983: 173; Kottelat, 1984: 275; Zakaria-Ismail, 1987: 409;
Kottelat, 1989: 19; Lim & Ng, 1990: 94; Lim et al., 1990a: 317;
1990b: 49; Witte & Schmidt, 1992: 327; Kottelat, 1994: 297;
Tan & Tan, 1994: 357; Lim, 1995: 161; Tan & Tan, 1996: 420;
Vidthayanon et al., 1997: 54; Sim, 2002: 74; Tan & Lim, 2004:
111.
Betta picta (non Valenciennes, in Cuvier & Valenciennes, 1846) Bleeker, 1879: 26; Weber & de Beaufort, 1922: 360; Fowler,
1938: 121; Herre, 1940: 44.
Betta trifasciata (non Bleeker, 1851) - Károli, 1881: 172 (part).
Betta macrophthalma Regan, 1910: 781.
Betta bleekeri Regan, 1910: 780 (part).
Betta brederi Myers, 1935: 25.
Betta fusca (non Regan, 1910) - Tweedie, 1936: 22; Fowler, 1938:
259; Herre, 1940: 46.
Betta taeniata (non Regan, 1910) - Tweedie, 1936: 22; Herre &Myers, 1937: 72; Herre, 1940: 45.
Betta rubra (non Perugia, 1893) - Herre, 1940:45.

habitats until recent findings have found otherwise (see Ng
et al., 1994).
Remarks. – See Tan & Ng (1996) for details of the taxonomic
history of B. bellica. The record of B. fasciata from Jambi
by Weber & de Beaufort (1922) is likely to be based on B.
simorum (fide Tan & Ng, 1996).
Betta fasciata was originally described from Sumatra andfrom meristic and morphometric counts, it is no different from
B. bellica. Betta fasciata was demonstrated to be a juniorsynonym of B. bellica with the designation of the larger
syntype as the lectotype by Tan & Ng (1996) (BMNH
1889.12.26:30, 69.1 mm SL), and a species (B. simorum)closely allied to B. bellica was described from Sumatra (where
both B. bellica and B. simorum occur). In the same paper,
Tan & Ng (1996) chose a neotype for B. bellica to stabilise
the taxonomy. The neotype is a male specimen, 51.4 mm SL(ZRC 39196) collected from the north Selangor peat swamp
forest in the state of Selangor, Peninsular Malaysia. This
locality is adjacent to the state of Perak, the original type
locality of B. bellica.
BETTA PUGNAX GROUP

Material examined. – Lectotype - Malaysia: Pulau Penang:
BMNH 1860.3.19:930, 60.6 mm SL; Pinang, coll. Cantor.

Betta pugnax (Cantor, 1850)(Figs. 11a-b, 46b)

Paralectotypes – Malaysia: Pulau Penang: BMNH 1860.3.19:317318, 2 ex. 51.8-66.2 mm SL; same data as lectotype.

Macropodus pugnax Cantor, 1850: 84.
Betta anabatoides (non Bleeker, 1851) - Bleeker, 1860a: 334; 1860b:

Others – Malaysia: Pulau Penang: ZRC 1610, 8 ex., 40.3-59.4mm SL; Batu Ferringi Catchment Area; Alfred et al., 1961. — ZRC

Fig. 10. Distribution map of B. bellica (circle) and B. simorum (triangle).

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
1611, 2 ex., 36.4-37.1 mm SL; Sungai Bayan Lepas; Alfred et al.,1961. — ZRC 1612, 6 ex., 42.4-58.2 mm SL; Sungai Telok Bahang,
141/2 miles Telok Bahang Road; Alfred et al., 1961. — ZRC 1613,
1 ex., 27.4 mm SL; Pulau Betong, Kampung Trang; Alfred et al.,1961. — ZRC 1614, 1 ex., 36.3 mm SL; Sungai Balik Pulau, 1 mile
north-east of Balik Pulau; Alfred et al., 1961. — ZRC 14753, 2 ex.,48.4-48.9 mm SL; Kolam Ayer; Alfred et al., 1964. — ZRC 31990,
4 ex., 44.3-57.0 mm SL; Kampung Tengah, coll. H. H. Tan & S.
H. Tan, 8 Jun.1993. — ZRC 32047, 2 ex., 44.4-59.5 mm SL; Pantai
Acheh, coll. H. H. Tan & S. H. Tan, 8 Jun.1993. — ZRC 32065,
7 ex., 40.3-50.9 mm SL; Sungai Relau (upper reaches), Kampung
Darat, coll. H. H. Tan & S. H. Tan, 9 Jun.1993. — ZRC 32302, 1
ex., 48.1 mm SL; Balik Pulau, Kampung Titi Serong (upstream),coll. H. H. Tan & S. H. Tan, 10 Jun.1993. — ZRC 39264, 1 ex.,60.1 mm SL; Penang, coll. K. K. P. Lim, 1995. Kedah: ZRC 11631,
1 ex., 47.3 mm SL; Pulau Langkawi, Dayang Bunting; not known,
Mar.1960. — ZRC 29633, 3 ex., 35.9-51.5 mm SL; Pulau Langkawi,
Telaga Tujuh waterfalls, coll. K. K. P. Lim, 30 Jan.1990.
Terengganu: ZRC 21720, 1 ex., 41.2 mm SL; Sekayu waterfalls,
coll. P. K. L. Ng et al., 18 Mar.1992. Pahang: ZRC 9003, 3 ex.,46.9-52.4 mm SL; Tasik Chini; 205 SQN RAF, Changi, 19-25
Aug.1967. — ZRC 8306-9, 3 ex., 46.7-67.3 mm SL; Rompin, easterntributaries of Sungai Kinchin; Rompin-Endau Expedition ZRC team,
13 Jun.1989. — ZRC 3476, 5 ex., 45.4-56.0 mm SL; Tasik Bera;
C. C. Lindsey, 29-31 Mar.1965. Perak: ZRC uncat., 4 ex., 41.352.9 mm SL; Taiping, base of Bukit Larut, coll. K. K. P. Lim et al.,21 Dec.1994. — ZRC 39572, 2 ex., Taiping, base of Bukit Larut,coll. H. H. Tan et al., 18 Nov.1995. Selangor: ZRC 435, 2 ex.,48.7-49.1 mm SL; West of Ginting Sempak (143/4miles from Kuala
Lumpur); A. W. Herre & Tweedie, 1937. — ZRC 19138, 1 ex.,54.5 mm SL; University of Malaya campus, in peaty stream, coll.
P. K. L. Ng, 20 Nov.1989. — ZRC uncat., 3 ex., 41.7-48.6 mm SL;
University of Malaya campus, in remnant swamp forest, coll. H. H.
Tan et al., 22 Dec.1994. — ZRC 39579, 1 ex., north Selangor
peatswamp forest, coll. P. K. L. Ng et al., Sep.1994. Johor: ZRC
23096, 2 ex., 47.5-48.6 mm SL; Ulu Endau, Sungai Anak Jasin; T.
H. T. Tan & J. B. Tay, 4 Apr.1992. — ZRC 28404, 3 ex., 38.9-39.8mm SL; Kota Tinggi, Sungai Tementang, coll. P. K. L. Ng et al.,15 Oct.1992. — ZRC 38220, 2 ex., 55.2-55.8 mm SL; Kota Tinggi,
Sungai Tementang, coll. H. H. Tan et al., 21 Aug.1994. — ZRC
11912, 2 ex., 49.0-54.3 mm SL; Kota Tinggi; C. F. Lim. — ZRC
25765, 1 ex., 47.6 mm SL; Sungai Mupor along Kota TinggiMersing road, coll. P. K. L. Ng et al., 24 Jul.1992. — ZRC 18130,
2 ex., 42.4-45.2 mm SL; 100 m north of road marker 175 km on
Johor Bahru-Kuantan road (north of Mersing), coll. P. K. L. Ng etal., 19 Oct.1991. — ZRC 19490, 1 ex., 43.1 mm SL; swamp nearabFig. 11. a) Betta pugnax - BMNH 1860.3.19:930, lectotype, 60.6mm SL, b) B. macrophthalma - BMNH 1868.7.10:28, holotype, 36.3mm SL.

Mersing, coll. P. K. L. Ng, Sep.1991. — ZRC 8442, 2 ex., 53.155.0 mm SL; Muar, 14th mile on Yong Peng Road; anon., 7
May.1962. — ZRC 19072, 1 ex., 47.3 mm SL; Mawai district; M.
W. F. Tweedie, 1935. — ZRC 8501, 1 ex., 48.3 mm SL; 14th mileon Muar-Yong Peng road; E. R. Alfred, 28 Sep.1967. — ZRC 28720,
1 ex., 42.2 mm SL; near Pontian, coll. P. K. L. Ng et al., 4 Mar.1992.
— ZRC 28995, 1 ex., 53.1 mm SL; Gunung Pulai Reservoir, coll.
P. K. L. Ng et al., 4 Mar.1992. — ZRC 19135, 1 ex., 50.6 mm SL;
stream near Gunung Pulai Reservoir (south), coll. P. K. L. Ng, 14
Mar.1990. — ZRC 1576.1-1576.11, 11 ex., 58.4-67.2 mm SL;
Gunung Pulai; L. K. Charles, Apr.1934. — ZRC uncat., 2 ex., 41.742.1 mm SL; Gunung Pulai, stream running under access roadtowards Pekan Nanas, coll. H. H. Tan et al., 20 May.1993. — ZRC
38240, 5 ex., 38.3-46.8 mm SL; Pontian, stream draining from
Gunung Pulai Reservoir, coll. P. K. L. Ng et al., 29 Sep.1994. —ZRC 13887, 3 ex., 42.9-52.5 mm SL; Gunung Panti foothills, coll.
P. K. L. Ng, 31 Aug.1990. — ZRC 19492, 2 ex., 59.0-59.6 mm SL;
Gunung Panti, forest, coll. P. K. L. Ng, Oct.1991. — ZRC 32485,
1 ex., 49.9 mm SL; Air Hitam, Parit Botak, blackwater stream, km
8; Yong, 23 I 1993. — USNM 094400, holotype of B. brederi, 64.2
mm SL female, Johor; A. Ramsperger, 22 Oct.1934. Singapore:
ZRC 38297, 9 ex., 44.7-52.6 mm SL; Nee Soon swamp forest, coll.
P. K. L. Ng et al., 23 Aug.1993. — ZRC 38298, 9 ex., 41.6-56.3mm SL; Nee Soon swamp forest, coll. P. K. L. Ng et al., 29 Jul.1993.
— ZRC 12196, 2 ex., 42.4-49.2 mm SL; forest stream draining into
northern arm of Lower Pierce Reservoir, coll. P. K. L. Ng, 13
Jul.1990. — ZRC 34621, 1 ex., 54.3 mm SL; Rifle Range road; C.
Y. Chang et al., 27 May.1993. — ZRC 30376, 2 ex., 40.6-44.2 mm
SL; Nee Soon swamp forest, edge of abandoned kampung near Upper
Thomson road, coll. P. K. L. Ng, 15 Jan.1993. — BMNH
1868.7.10:28, holotype of B. macrophthalma, 36.3 mm SL juvenile;
Singapore.

Diagnosis. – Betta pugnax is distinguished from othermembers of the B. pugnax species group by the following
combination of characters: chin-bar present; no dark marks
below postorbital stripe on opercle; dorsal transverse bars
present; caudal transverse bars present only in male; greenish
to blueish iridescent scales on opercle and body; anal-fin with
red subdistal band and black edge; no dark edge on caudalfin; anal-fin rays 25-28 (mode 26); lateral scales 28-31 (mode29); postdorsal scales 11-12 (mode 11); pelvic fin length 26.643.8% SL; head length 27.5-35.2% SL; inter-orbital width
32.4-53.0% HL.
Description. – General body shape and appearence as in Figs.
11a-b, 46b. Body slender; dorsal-fin origin above 13th lateralscale; posterior extremity of dorsal fin pointed, reaching aboutanterior third of caudal fin; anal-fin origin below 7th lateralscale; posterior extremity of anal fin pointed and reaching
beyond midlength of caudal fin; caudal fin broadly lanceolate(Fig. 5b); pelvic fin filamentous. Meristics: lateral scales 2930; predorsal scales 19-22 (mode 21); subdorsal scales 5-8(mode 6); postdorsal scales 11-12 (mode 11); dorsal-fin rays
0-II, 7-10 (total 8-10, mode 9); anal-fin rays I-II, 24-26 (total
25-28, mode 26); caudal fin rays ii, 4+5, ii; pelvic fin rays
I, 1, 4; pectoral fin rays 13; vertebrae 2 + 8 + 18-21 = 28-31.
Morphometrics: total length 130.6-148.1% SL; predorsal
length 60.9-73.4% SL; preanal length 40.6-51.2% SL; pelvicfin length 26.6-43.8% SL; head length 27.5-35.2% SL; body
depth at dorsal-fin origin 24.8-32.1% SL; length of anal-finbase 44.2-54.0% SL; length of dorsal-fin base 9.9-15.7% SL;
orbital diameter 27.9-38.5% HL; postorbital length 45.0-

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Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei
and ranged from fast flowing to sluggish. The substrate wasusually sandy with rocks. There was little or no submerged
vegetation. Overhanging bank vegetation, submerged leaf
litter and exposed roots of bank vegetation constituted themain habitats for B. pugnax. It was mostly found in and at
the periphery of stagnant areas . The pH was around 7.4 (range
7.1-7.6). The water depth ranged from 10 to 80 cm. There
were no other syntopic osphronemids in these habitats. The
only other Betta found on Pulau Penang is B. imbellis.

55.7% HL; interorbital width 32.4-53.0% HL; head length41.6-54.3 in% predorsal length. Maximum known size 67.3mm SL (ZRC 8306).
Coloration. – For live coloration, see Fig. 46b (see also Linke,
1991: 43; Kubota et al., 1996: 38). Live topotypes from
Penang have blue iridescent opercle with a light brownbackground. each scale of body and opercle, a median areawith blue to greenish-blue iridescence. Specimens from
Taiping, Perak, also exhibit the same character (these two
localities are hill streams with clear waters, pH 7 and above).
Specimens from more acidic waters (e.g., Singapore), usuallywith less blue and more green iridescence. Caudal transversebars in males (alive: strongly barred depending on state offish; in preserved specimens: faintly barred or not apparent)(Fig. 2). Preserved specimens dark brown dorsally and lightbrown ventrally, fins slight brownish to hyaline; male withblackened opercle region, female and juvenile with chin-bar,central and second central stripes fairly distinct, with caudal
peduncle spot.

Comparative notes. – Betta pugnax is distinguished from theother members of the B. pugnax species group by thefollowing characters: more anal-fin rays than B. fusca (2528, mode 26, vs. 24-25); more predorsal scales than B.
schalleri (19-21, mode 20, vs. 17-19, mode 19); fewer
predorsal scales than B. fusca (19-21, mode 20, vs. 22-23);
shorter preanal length than B. prima and B. enisae (40.651.2% SL, vs. 50.1-54.8); relatively shorter head length thanB. fusca, B. enisae, B. schalleri, B. pulchra, B. prima and B.
lehi (27.5-35.2% SL, vs. 32.7-37.7); larger orbital diameter
than B. fusca and B. enisae (27.9-38.5% HL, vs. 23.8-28.5).

Distribution. – Betta pugnax is widely distributed in
Peninsular Malaysia. It is found in Pulau Penang, Kedah,
Terengganu, Pahang, Perak, Selangor, Johor and Singapore(Fig. 12). It is also found in the Anambas islands (Tan &Lim, 2004), Riau Archipelago, Lingga Archipelago, Riau
province and Jambi province (Tan & Ng, 2005) (present
volume).

Remarks. – Bleeker (1879) identified 130 specimens fromvarious localities as B. picta. In the volume 9 of his Atlas(1877: pl. 395, fig. 3), he figured the species, but he did notindicate the origin of the figured specimen. Neither did he in
his 1879 paper. The text due to accompany this plate has neverbeen published (Bleeker died in 1878).

Field notes. – Specimens from Pulau Penang were mainlycollected in hill streams and foothills. These waters were clear

Regan (1910: 780) described B. bleekeri based on Bleeker’s
1877 figure and 1879 text. He listed B. picta (sensu Bleeker)

Fig. 12. Distribution map of Betta pugnax (circle), B. pulchra (triangle), B. stigmosa (upside-down triangle) and B. lehi (square).

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
Johor, Pontian, ca. 4 km towards Pontian Kechil from Sri Bunian(1°27’13.1"N 103°24’52.1"E), coll. P. K. L. Ng et al., 15 Aug.1995.
Others - ZRC. 39251, 15 ex., 21.5-36.5 mm SL; Malaysia: Johor,
Pontian, Sri Bunian, Kampung Pt. Tekong (1°27’59.2"N
103°26’07.7"E), coll. P. K. L. Ng et al., 15 Aug.1995. — ZRC.
39250, 13 ex., 21.7-43.1 mm SL; Malaysia: Johor, Pontian, Sri
Bunian, Kampung Jasa Sapakat (1°31’30.5"N 103°27’47.7"E), 5
km into side road leading into oil-palm estate, coll. P. K. L. Ng etal., 15 Aug.1995.

as a synonym of B. bleekeri. He also commented that thedescription of B. picta (sensu Bleeker) is based on the figure
in Bleeker’s Atlas (1877: pl. 395, fig. 3). Regan (1910) clearlyconsiders the figure in Bleeker’s Atlas to be a species distinct
from B. picta. This means that all of the specimens mentioned
or used by Bleeker (1877, 1879) for B. picta are syntypes ofB. bleekeri (except the one used by Regan for description of
B. bleekeri, based on Bleeker’s 1877 figure). Since Bleeker(1879) included B. trifasciata, B. pugnax and B. anabatoides,in the synonymy of B. picta, all his specimens of B. trifasciata
and B. anabatoides and all his and Cantor’s specimens of B.
pugnax , also are syntypes of B. bleekeri. This is problematicwhen selecting a lectotype or clarifying the identity of B.
bleekeri.

Diagnosis. – Betta pulchra is distinguished from othermembers of the B. pugnax species group by the following
combination of characters: chin-bar present; no dark marks
below postorbital stripe on opercle; dorsal transverse bars
present; caudal transverse bars present only in male; greenish
iridescent scales on opercle and body; anal fin with red
subdistal border and black edge; caudal fin without distal darkedge; anal-fin rays 24-28 (mode 26); lateral scales 28-30(mode 29); postdorsal scales 10-11 (mode 10); pelvic fin
length 29.1-53.2% SL; head length 32.7-37.6% SL; interorbital width 34.3-41.8% HL.

Witte & Schmidt (1992: 326) synonymised B. bleekeri with
B. bellica. Bleeker’s (1877) figure of B. picta is very unlikely
to be based on a specimen of B. bellica. General morphologyof the figured specimen, meristics and morphometrics, it
seems to belong to the B. waseri group, as defined by Ng &Kottelat (1994). Interestingly, Witte & Schmidt (1992: 326)
regarded Bleeker’s figure as coming close to B. patoti, butthey still provisionally synonymised B. bleekeri under B.
bellica, arguing the single B. picta (RMNH 10741) they had
examined from Bleeker’s 130 specimens (!), presumably from
Sumatra, matched B. bellica. They did not state how they
identified the locality of the specimen.

Description. – General body shape and appearance as in Figs.
13, 46c. Body short and stocky, body depth at dorsal-fin origin
25.9-36.9% SL; head length 32.7-37.6% SL; head pointed
and stout; caudal fin narrowly lanceolate (Fig. 5c); pelvic fin
filamentous; dorsal and anal fins pointed, anal fin reaching
to half or more of caudal fin. Meristics: lateral scales 28-30(mode 29); transverse scales at dorsal-fin origin 91/2; subdorsal
scales 6-71/2 (mode 7); predorsal scales 18-21 (mode 20);
postdorsal scales 10-11 (mode 10); dorsal-fin rays I-II, 8-9(total 9-10, mode 9); anal-fin rays I-II, 22-26 (total 24-28,
mode 26); caudal fin rays ii, 4+5, ii; pectoral fin rays 12;
pelvic fin rays I, 1, 4; vertebrae 2 + 8 + 19 = 29.
Morphometrics: total length 136.6-156.6% SL; predorsal
length 60.8-66.0% SL; preanal length 44.7-50.0% SL; pelvicfin length 29.1-53.2% SL; head length 32.7-37.6% SL; body
depth at dorsal-fin origin 25.9-36.9% SL; length of anal-finbase 49.8-52.9% SL; length of dorsal-fin base 13.9-15.9%
SL; orbital diameter 26.0-31.3% HL; postorbital length 44.349.7% HL; interorbital width 34.3-41.8% HL; head length49.8-58.5% predorsal length. Maximum known size 55.9 mm
SL (ZRC 17368).

In order to definitively fix the nomenclatural status of B.
bleekeri, we have decided to synonymise B. bleekeri with B.
pugnax. Betta pugnax is listed by Bleeker (1879) as asynonym of B. picta, and all the material which Bleeker
considered as belonging to B. picta constitutes the type seriesof B. bleekeri, therefore the type series of B. pugnax is partof the type series of B. bleekeri. We have chosen as thelectotype of B. bleekeri the same specimens (BMNH
1860.3.19.930) which had been designated as lectotype of B.
pugnax by Alfred (1964). This makes B. bleekeri a juniorobjective synonym of B. pugnax. This action will not affectthe taxonomy of the other Betta species, as the name B.
bleekeri had been only used thrice (Regan, 1910: 780; Weber& de Beaufort, 1922: 357; Witte & Schmidt, 1992: 326).
Other details of the taxonomic history of B. pugnax are
discussed in Tan & Tan (1996).

Coloration. – For live coloration, see Fig. 46c (see also
Kubota et al., 1996: 39, bottom figure identified as B. pugnax).
Pattern similar to that of B. pugnax sensu stricto, except that
more distinctly greenish iridescent scales (cf. blue in B.
pugnax). Guanophores on body generally more intense than

Betta pulchra Tan & Tan, 1996
(Figs. 13, 46c)
Betta pulchra Tan & Tan, 1996: 428.
Material examined. – Holotype - Malaysia: - ZRC 28860, 49.9mm SL; Malaysia: Johor, Pontian, Kampung Jasa Sepakat, coll. P.
K. L. Ng et al., 8 May.1992.
Paratypes – ZRC 28861-28899, 39 ex., 19.8-46.5 mm SL; same
locality data as holotype. — ZRC 17368-17370, 3 ex., 34.4-55.9mm SL; Malaysia: Johor, Pontian, coll. P. K. L. Ng et al., 1989. —ZRC 29123-29125, 3 ex., 44.8-50.2 mm SL; Malaysia: Johor,
Pontian, Sri Bunian, Kampung Pt. Tekong, coll. P. K. L. Ng et al.,
8 May.1992. — ZRC. 39252, 5 ex., 39.0-45.3 mm SL; Malaysia:

Fig. 13. Betta pulchra – ZRC 28860, holotype, 49.9 mm SL (rightside, reversed).

63

Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei
Others – ZRC 21800, 1 ex., 20.0 mm SL; Malaysia: Terengganu,
Pasir Puteh Air Terjun, 4 km from turnoff, coll. P. K. L. Ng et al.,
20 Mar.1992.

those of B. pugnax in life. Opercle with iridescent greenishblue scales; body brown with iridescent green scales. Headwith preorbital and postorbital stripes, continuous with central
stripe, second central stripe and caudal peduncle spot in both
sexes and juvenile. Distal part of pelvic ray iridescent green;
dorsal transverse bars present; caudal fin brownish, caudal
transverse bars present in mature male; anal fin with red
subdistal margin and black edge.

Diagnosis. – Betta stigmosa is distinguished from the othermembers of the B. pugnax group by the following unique
combination of characters: 5-10 regularly spaced blacktransverse bars on caudal fin of male and female; in life, bothmale and female with alternating blueish and black transversebars on dorsal and caudal fins; both male and female withblack transverse bars on posterior few anal-fin rays; anal fin
with dark distal edge. Betta stigmosa shares with B. prima
the presence of a second postorbital stripe on opercle; small
adult size (31-41 mm SL); total anal-fin rays 23-24; lateralscales 29-30 (mode 29); postdorsal scales 10-11 (mode 10);pelvic fin length 33.2-51.8% SL; head length 33.6-36.5% SL;interorbital width 29.8-32.1% HL.

Field notes. – Betta pulchra is found in blackwater habitats;other syntopic osphronemids present are Belontia hasseltii,
Betta bellica, B. imbellis, Sphaerichthys osphromenoides,
Parosphromenus sp. and Trichopsis vittata. This species iscommon only in the overgrown bank vegetation of theirrigation canals. Much of the district of Pontian was peatswamp forest in the past, but most of it has been logged andplanted with oil-palm, pineapple and rubber trees. The pH of
the waters range from 3.9 to 4.2. Betta pulchra is an oral
brooder (Tan, 1997).

Description. – General appearance as illustrated in Figs. 14ab, 46d. Meristic and morphometric data in Table 3. Body
relatively slender (body depth at dorsal-fin origin 25.6-28.2%
SL), head relatively long and blunt (head length 33.6-36.5%
SL; head width 19.0-21.5% SL and 56.0-58.9% HL). Dorsal,
caudal and anal fins pointed; dorsal fin placed relatively far
back (predorsal length 62.8-67.5% SL); caudal fin with distal
part of rays extended slightly beyond interradial membrane,
median rays elongated in male; length of anal-fin base about
half of standard length (48.2-51.3% SL); pelvic fin roundedwith relatively long filamentous ray (33.2-51.8% SL); pectoral
fin rounded. Vertebrae 2 + 8 + 18-19 = 28-29 (mode 29, n= 7). Maximum known size 41.1 mm SL (ZRC 21720).

Distribution. – Betta pulchra appears to be restricted to theblackwaters in Pontian, southwest Johor, Malaysia (Fig. 12).
Comparative notes. – Betta pulchra is distinguished fromthe other members of the B. pugnax species group in having:
more subdorsal scales than B. fusca, B. prima and B. enisae(6-71/2, vs. 5-6); more lateral scales than B. enisae (28-30,
mode 29, vs. 26-28, mode 27); fewer lateral scales than B.
schalleri (28-30, vs. 31); fewer predorsal scales than B. fusca(18-21, vs. 22-23); shorter predorsal length than B. fusca, B.
prima and B. enisae (60.8-66.0% SL, vs. 67.3-70.3); shorter
preanal length than B. prima and B. enisae (44.7-50.0% SL,
vs. 50.1-54.8); relatively greater head length than B. pugnax(32.7-37.6% SL, vs. 27.5-35.2); greater length of dorsal-finbase than B. fusca and B. schalleri (13.9-15.9% SL, vs. 12.413.9); greater orbital diameter than B. fusca (26.0-31.3% HL,
vs. 23.8-26.7); greater interorbital width than B. fusca (34.341.8% HL, vs. 33.5-34.2); greater head length than B. fusca(49.8-58.5% predorsal length, vs. 47.2-49.4).

Coloration. – The following is based on freshly preserved
specimens. Male: reddish brown body, darker brown head,
opercle and throat area greenish-blue iridescence (darkly
pigmented when preserved longer). Lower lip black, chinbar present but may be masked by dark pigmentation. Preand postorbital stripes present, second postorbital stripe
masked by dark pigmentation. Belly whitish, ventral part ofbody lighter reddish brown. Dorsal and caudal fins brown.
Dorsal fin with distinct or faint 3-5 black transverse oninterradial membrane. Caudal fin with thin black edge on

Betta stigmosa, new species(Figs. 14a-b, 46d)a

Betta pugnax (non Cantor, 1850) - Tweedie, 1952a: 73 (part);
Mohsin & Ambak, 1983: 173; Tan & Tan, 1996: 420 (part).
Betta cf. pugnax (non Cantor, 1850) - Kottelat et al., 1992: 11 (listonly).
Material examined. – Holotype - ZRC 43392, male, 36.9 mm SL;
Malaysia: Terengganu, Sekayu waterfalls (04°57’49.7”N
102°57’14.0”E), near swampy stream, coll. P. K. L. Ng et al., 10
Oct.1997.

bParatypes – ZRC 41710, 5 ex., 22.3-34.0 mm SL; same locality
data as holotype. — ZRC 43393, 7 ex., 29.2-37.5 mm SL; Malaysia:
Terengganu, Sekayu, swampy stream draining into concrete pondnear chalets (04°57.80’N 102°57.21’E), coll. H. H. Tan & K. K. P.
Lim, 21 & 28 Oct.1998. — ZRC 21720, 1 male, 41.1 mm SL;
Malaysia: Terengganu, Sekayu waterfalls (4°57’49.7”N
102°57’14.0”E), coll. P. K. L. Ng et al., 18 Mar.1992.

Fig. 14. a) Betta stigmosa – ZRC 43392, male holotype, 36.9 mm
SL, b) ZRC 41710, female paratype, 32.2 mm SL.

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
lower half, 5-10 regularly spaced dark brown or blacktransverse bars on interradial membrane. Anal fin reddish withthin black edge, 2-6 regularly spaced dark brown transversebars on posterior 3-4 rays. Pelvic fin brown, filament darkly
pigmented. Pectoral fin hyaline.
Female: yellowish brown body, darker brown head, opercle
and throat area yellowish (not darkly pigmented after longterm preservation). Lower lip black, chin-bar present withanterior portion elongated towards mouth (Fig. 15). Pre- andpostorbital stripes present, second postorbital stripe
interrupted into 4-5 staggered parts (Fig. 15). Belly whitish,
ventral part of body light brown. Dorsal and caudal finsbrown. Caudal fin without thin black edge on lower half, 38 regularly spaced black transverse bars on interradial
membrane. Anal fin reddish brown with thin black edge, 25 regularly spaced dark brown transverse bars on posterior
3-4 rays. Pelvic and pectoral fins hyaline, pelvic filamenthyaline. Juvenile with faint central and second central stripes,
dorsal and caudal fins hyaline, anal fin with 2 longitudinalrows of dark brown spots.
Life coloration. The color pattern is similar to above described(see Fig. 46d). The transverse bar pattern on dorsal and caudal
fins consist of alternating slightly iridescent blue and blacktransverse bars. Pelvic fin bright white.

Field notes. – The type locality is a swampy stream drainingout from a small hill into the Sekayu waterfalls. Syntopic
species present: Poropuntius smedleyi, Puntius binotatus,
Rasbora cf. bunguranensis (Cyprinidae), Clarias sp.
(Clariidae), Amblyceps mangois (Amblycepitidae), Channagachua, C. melasoma (Channidae) and tilapia (Cichlidae).
Etymology. – From the Latin stigmosus, meaning full ofmarks, pertaining to the patterned opercle and barred pattern
on dorsal, caudal and anal fins of both male and female
specimens.
Comparative notes. – Betta stigmosa is distinguished fromthe other members of the B. pugnax group in the following
characters: presence of caudal transverse bars in male andfemale (vs. in male only or absent in both sexes); transverse
bar pattern on posterior anal-fin rays (vs. absent); fewer analfin rays than other members (23-24, vs. 25-29); more
subdorsal scales than B. fusca, B. prima and B. enisae (61/2,
vs. 5-6); more lateral scales than B. enisae (29-30, vs. 2628); fewer lateral scales than B. schalleri (29-30, vs. 31-33);
more predorsal scales than B. schalleri (20-21, vs. 17-19);
fewer predorsal scales than B. fusca (20-21, vs. 22-24);
smaller predorsal length than B. fusca, B. prima and B. enisae(62.8-67.5% SL, vs. 67.3-70.3); smaller postdorsal length thanB. prima (21.2-23.6% SL, vs. 24.1-26.3); greater head lengththan B. fusca (51.7-55.4% predorsal length, vs. 47.2-49.4);
smaller body depth at dorsal-fin origin than B. fusca, B. prima
and B. breviobesus (25.6-28.2% SL, vs. 28.1-33.5); greater
length of dorsal-fin base than B. enisae (12.2-13.9% SL, vs.
8.7-12.1); smaller length of anal-fin base than B. breviobesus(48.2-51.3% SL, vs. 51.1-55.7); greater orbit diameter thanB. fusca (26.7-28.4% HL, vs. 23.8-26.7); smaller head widththan B. fusca (56.0-58.9% HL, vs. 59.7-69.8); smaller orbit
diameter than B. schalleri (52-60% postorbital length, vs. 6176); smaller interorbital width than B. fusca, B. schalleri, B.
prima, B. enisae, and B. lehi (62-65, vs. 65-84) (data for B.
prima; B. schalleri; B. enisae; B. pugnax and B. pulchra; andB. breviobesus from Kottelat, 1994, 1995; Kottelat & Ng,
1994; Tan & Tan, 1996; and Tan & Kottelat, 1998,
respectively).

Distribution. – Betta stigmosa is currently known only fromthe hill streams in Sekayu area, Terengganu, Peninsular
Malaysia (Fig. 12).

Remarks. – Tan & Tan (1996) revised the taxonomy of B.
pugnax from Peninsular Malaysia. They overlooked thedistinctiveness of B. stigmosa for which they had only two
specimens (ZRC 21720, ZRC 21800) at the time. The larger
series now available shows the unique character of the blacktransverse bars on the caudal fin of both male and female.
This is the second species of the B. pugnax group in which
this character is observed, the other being B. enisae.

Betta lehi, new species(Figs. 16, 46e)
Betta pugnax (non Cantor, 1850) - ?Károli, 1881: 172.
Betta (sp. E) sp. E Witte & Schmidt, 1992: 324 (key), 327 (part).
Betta spec. Brown & Brown, 1987: 166.
Betta sp. Kottelat & Lim, 1995: 248.

Fig. 15. Schematic drawings of lateral and ventral head views of B.
stigmosa (ZRC 41710, female, 31.6 mm SL).

Material examined. – Holotype - ZRC 39267, 50.7 mm SL male,65

Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei
Borneo: Sarawak, Sungai Stuum Muda (1°28’51.3"N
109°58’18.1"E), ca. 58 km to Sematan on road from Bau to Lundu,
21.1 km before Lundu ferry over Batang Kayan, coll. H. H. Tan &D. C. J. Yeo, 7 Sep.1995.

pugnax). Mature male with an iridescent light blue margin
on anal fin and faintly on caudal fin when live, not visible
when preserved. Body iridescence faint when alive, absent
when preserved. Opercle iridescent green when alive, darkbrown when preserved. Dorsal fin pointed, may reachmidlength of caudal fin; penultimate anal ray sometimes
extending beyond tip of caudal fin; tip of pelvic fin white;caudal fin interradial membrane faint blue when alive, brownwhen preserved. Females with less colour and shorter fins.
Throat with chin-bar, female and juvenile with two black
marks on opercle below postorbital stripe (Fig. 6b). Pattern
on male opercle obscured by iridescent scales, darkly
pigmented when preserved. Upper half of eye shiny yellow
when alive. Preserved specimens dark brown dorsally andlighter brown ventrally. Fins light brown, anal and caudal
fins with faint darkly pigmented margin.

Paratypes – ZRC 41582, 15 ex., 18.5-47.8 mm SL, same locality
data as holotype. — ZRC 41583, 11 ex., 14.0-49.1 mm SL; CMK
13021, 5 ex., 21.1-48.7 mm SL; ZMA 121.674, 5 ex., 22.5-47.9mm SL; SM uncat., 5 ex., 25.5-41.6 mm SL; same locality data as
holotype, coll. H. H. Tan et al., 6 Sep.1995.
Others – ZRC 37875, 4 ex., 23.1-60.8 mm SL; Borneo: Sarawak,
Lundu-Bau road, 27 km before Bau, coll. M. Kottelat et al., 8
May.1994. — ZRC 26055-57, 3 ex., 22.7-47.3 mm SL; - CMK 8414,
3 ex., 16.1-45.9 mm SL; Borneo: Sarawak, 42 km before Lundu
from Kuching, after Sungai Stinggang, coll. M. Kottelat, P. K. L.
Ng & K. K. P. Lim, 3 Jul.1992. — ZRC 40925, 38 ex., 17.4-41.6mm SL; USNM 344462, 10 ex., 26.0-36.3 mm SL; MZB uncat., 5
ex., 26.8-33.1 mm SL; Borneo: Sarawak, Sungai Sebiris(1°41’32.0”N 109°47’00.8”E), 13.8 km after Kampung Pueh turnoff,
towards Lundu on Lundu-Sematan road, coll. P. K. L. Ng et al., 1
Sep.1996. — ZRC 40926, 2 ex., 37.3-38.0 mm SL; Borneo: Sarawak,
Sungai Stunggang (1°37’28.5”N 109°53’10.9”E), 4.8 km before
Lundu ferry over Batang Kayan, coll. P. K. L. Ng et al., 2 Sep.1996.
— ZRC 40927, 39 ex., 14.9-49.6 mm SL; Borneo: Sarawak, Sungai
Stuum Muda (1°28’51.3"N 109°58’18.1"E), 21.1 km before Lundu
ferry point over Batang Kayan, coll. P. K. L. Ng et al., 2 Sep.1996.
— ZRC 46268, 9 ex., 43.8-54.0 mm SL; Kalimantan Barat,
Pontianak, aquarium trade; donor F. Yuwono, Oct.1999.

Distribution. – Betta lehi is apparently restricted to the area
southwest of Kuching, between the towns of Bau and Lundu
in south Sarawak; and in the lower Kapuas basin in
Kalimantan Barat (Fig. 12).
Field notes. – Betta lehi inhabits peat swamp forests and acidwater swamp forests within the area mentioned, and appears
to replace B. ibanorum in these habitats. At the type locality,
Sungai Stuum Muda is a stream about 3 m wide and at someparts up to 1.5 m deep (water level can rise much more after
heavy rains; pers. observ.). The water was fast flowing andclear with a pH ca. 6.8. All specimens of B. lehi were collectedfrom quiet parts of the stream among the overgrown
submerged bank vegetation. The aquatic plant Barclaya
motleyi (Nymphaeaceae) was found upstream of the bridge,
indicating the presence of freshwater swamp forest nearby.
Fish species syntopic with B. lehi include: Rasbora
sarawakensis, R. sumatrana, Puntius banksi, P. kuchingensis,
Hampala macrolepidota (Cyprinidae), Homaloptera tweediei,
Nemacheilus saravacensis (Balitoridae), Acanthopsoides
robertsi, Pangio semicincta, P. shelfordii (Cobitidae),Pseudomystus fuscus (Bagridae), Glyptothorax cf. major(Sisoridae), Hemirhamphodon kuekenthali (Hemiramphidae),
Doryichthys martensii (Syngnathidae), Pseudogobiopsis
siamensis (Gobiidae), Nandus nebulosus (Nandidae),
Luciocephalus pulcher (Osphronemidae), Channa lucius(Channidae), Phenacostethus sp. (Phallostethidae) andCarinotetraodon salivator (Tetraodontidae).

Diagnosis. – Betta lehi is distinguished from the othermembers of the B. pugnax species group by the following
combination of characters: no chin-bar; dorsal fin transversebars present; no caudal fin transverse bars; green iridescentscales on opercle and body; anal fin with iridescent light blue
distal border; and two black marks below postorbital stripe
on opercle of female and juvenile (Fig. 11); anal-fin rays 2529 (mode 28); lateral scales 28-30 (mode 29); postdorsal
scales 10-11 (mode 11); pelvic fin length 34.8-50.6% SL;head length 33.4-36.8% SL; interorbital width 31.7-37.4%
HL.
Description. – General body shape and appearance as in Figs.
16, 46e. Meristic and morphometric data in Table 3. Bodystocky, relatively long and pointed head; caudal fin lanceolate,dorsal and anal fins pointed, anal fin reaching end of caudal
fin, pelvic fin filamentous. Maximum known size 60.8 mm
SL (ZRC 37875).

Etymology. – Named after Charles M. U. Leh, curator of
Zoology at the Sarawak Museum, in recognition of his kind
help.

Coloration. – For live coloration, see Fig. 46e (see also
Kubota et al., 1996: 42, figures identified as B. sp. aff.

Comparative notes. – Betta lehi is distinguished from theother members of the B. pugnax species group in having:
more anal-fin rays than B. fusca (25-29, mode 28, vs. 2425); more lateral scales than B. enisae (28-30, mode 29, vs.
26-28, mode 27); fewer lateral scales than B. schalleri (2830, vs. 31); more predorsal scales than B. schalleri (20-23,
mode 22, vs. 17-19, mode 19); shorter predorsal length thanB. fusca, B. enisae and B. prima (62.6-67.5% SL, vs. 67.370.3); greater head length than B. fusca (33.4-36.8% SL, vs.
33.2-33.8); greater body depth at dorsal-fin origin than B.

Fig. 16. Betta lehi – ZRC 39267, male holotype, 50.7 mm SL.

66

THE RAFFLES BULLETIN OF ZOOLOGY 2005of airport runway end on Jalan Teku, coll. M. Kottelat et al., 15
May.1994. — ZRC 37845, 8 juv.; outskirts of Sibu, 4.2 km northof airport runway end on Jalan Teku, coll. M. Kottelat et al., 15
May.1994. — ZRC 37925, 6 ex., 19.7-60.2 mm SL; ditch 34 km
from Mukah, on Mukah-Sibu road, coll. M. Kottelat et al., 14
May.1994. — ZRC 37914, 1 juv.; Sungai Mantala, about 28 km
from Mukah on Mukah-Sibu road, coll. M. Kottelat et al., 14
May.1994. — ZRC 37940, 2 juv.; Sungai Gayao, about 40 km from
Mukah on Mukah-Sibu road, coll. M. Kottelat et al., 14 May.1994.
— ZRC 37902, 1 juv.; Sungai Ngeli, about km 6 Simunjan-Balairoad, coll. M. Kottelat et al., 11 May.1994. — ZRC37952, 5 ex.,17.5-50.8 mm SL; Sibu-Sarikei road, 2-3 km south of Durin ferry,coll. M. Kottelat et al., 15 May.1994. — ZRC 40928, 7 ex., 10.851.1 mm SL; Miri, stream below cascades flowing out of Lambir
Hill park (3°45’20.1”N 113°45’25.5”E), coll. H. H. Tan & Y. Y.
Goh, 26 Sep.1996. — ZRC 40929, 20 ex., 12.6-32.7 mm SL; Miri,lower Tukao, remnant swampforest ca. 200m from sea (4°18’53.7”N
113°58’07.5”E), coll. H. H. Tan & Y. Y. Goh, 26 Sep.1996. —MNHN 1891-515 to 518, 4 ex., 68.4-75.6 mm SL; Borneo, coll.
Chaper. — MNHN 1891-521 to 528, 8 ex., 49.7-71.3 mm SL;
Borneo, coll. Chaper. Brunei: ZRC 31812-17, 6 ex., 9.4-56.6 mm
SL; Belait District, near Seria, coll. K. K. P. Lim, 11 May.1993. —ZRC 35437-441, 5 ex., 34.1-71.6 mm SL; Belait District, Sungai
Liang area, coll. Helen & S. C. Choy, 4 Oct.1992. — ZRC 3186364, 2 juv.; Tutong District, Lamunin, coll. K. K. P. Lim & S. C.
Choy, 14 May.1993. — ZRC 40930, 25 ex., 15.3-71.1 mm SL; Belait
District, Sungai Bang Balat, Sungai Kebun, Sungai Mengiris, Sungai
Temusu - all draining into Sungai Melilas, tributary of Sungai Belait(jetty point 4°15’24.3”N 114°39’40.2”E), coll. H. H. Tan et al., 8
and 12 May.1996. — ZRC 40931, 3 ex., 39.3-75.1 mm SL; Belait
District, Sungai Ingei (4°9’42.6”N 114°42’59.5”E), coll. H. H. Tan
et al., 9-11 May.1996. — ZRC 40932, 3 ex., 22.7-61.4 mm SL;
Belait District, Sungai Jaung, stream below hot spring, coll. H. H.
Tan et al., 10 May.1996. — ZRC 40933, 40 ex., 16.8-72.7 mm SL;
Belait District, Sungai Sepan, tributary of Sungai Ingei running
through Kerengas forest, coll. H. H. Tan et al., 11 May.1996. —ZRC 40986, 42 ex., 20.7-59.1 mm SL; Belait District, peat swamp
forest next to Sungai Sepan, coll. H. H. Tan et al., 11 May.1996.
— ZRC 40934, 47 ex., 12.8-72.5 mm SL; BRM uncat., 10 ex., 24.851.1 mm SL; CMK 13121, 10 ex., 23.6-62.8 mm SL; ZMA 121.676,
5 ex., 34.4-60.4 mm SL; CAS 91716, 5 ex., 33.7-57.4 mm SL; Belait
District, Sungai Pelok, tributary of Sungai Ingei, coll. H. H. Tan etal., 11 May.1996. — ZRC 40935, 50 ex., 21.1-60.5 mm SL; BRM
uncat., 10 ex., 29.0-50.0 mm SL; CMK 13122, 10 ex., 27.4-58.1mm SL; BMNH 1997.4.3.1-5, 5 ex., 28.9-52.0 mm SL; RMNH

schalleri (27.3-32.5% SL, vs. 26.7-27.6); greater length of
dorsal-fin base than B. enisae (12.3-15.6% SL, vs. 8.7-12.1);
greater orbital diameter than B. breviobesus (27.2-36.8% HL,
vs. 24.2-28.5); shorter postorbital length than B. fusca (42.248.5% HL, vs. 48.3-50.5); greater interorbital width than B.
breviobesus (70-84% postorbital length, vs. 64-74); greater
head length than B. fusca (51.2-56.1% predorsal length, vs.
47.2-49.4).
Remarks. – Károli (1881) listed B. pugnax from Matang and
Palandok in Borneo. Specimens obtained from recent
collections from Matang area in Sarawak consist only of B.
ibanorum (see later). Thus, Károli may have misidentified
the specimens from Matang, Sarawak. The locality Palandokcannot be found in current maps and cannot match any site
in Sarawak (assuming it is in Sarawak). Witte & Schmidt(1992: 327) treated material from Sumatra and Sarawak as
a Betta (sp. E) sp. E. Based on their locality data, their
Sarawak specimens are most probably B. lehi. Kottelat & Lim(1995) had only collected very few specimens of B. lehi up
to 1995 and had listed it tentatively as only Betta sp.

BETTA AKARENSIS GROUP
Betta akarensis Regan, 1910
(Figs. 17 a-b, 46f)
Betta akarensis Regan, 1910: 779, pl. 77, Fig. 3; Weber & deBeaufort, 1922: 357; Roberts, 1989: 172 (table only); Witte &Schmidt, 1992: 326; Kottelat et al., 1993: 161, pl. 75 (part); Ng,
1993: 289 (part); Kottelat & Ng, 1994: 74 (part); Kottelat &Lim, 1995: 248 (part); Tan & Ng, 2004a: 278, figs. 3, 7.
Betta climacura Vierke, 1988: 336; Witte & Schmidt, 1992: 326;
Kottelat et al., 1993: 161 (part); Ng, 1993: 290 (part); Kottelat& Ng, 1994: 74 (part).
Betta anabatoides (non Bleeker, 1851) - Regan, 1910: 780, pl. 78,
fig. 4 (part); Vinciguerra, 1926: 592; Kottelat & Lim, 1995: 248.
Betta unimaculata (non Popta, 1905) - Zakaria-Ismail, 1984:73;
Kottelat & Lim, 1995: 248.
Betta spec. affin. taeniata (non Regan, 1910) - Brown & Brown,
1987:160, pl. 12.
Material examined. – Holotype - Sarawak: BMNH 1895.7.2:44,
31.6 mm SL juvenile; Sungai Akar, coll. C. Hose.

a

Others - BMNH 1868.6.9:18-20, 2 ex., 46.4-56.1 mm SL; Sarawak,
coll. M. Doria. — BMNH 1889.7.31:4-5, 2 ex., 58.5-62.5 mm SL;
Baram, coll. C. Hose. — BMNH 1891.1.27:26-27, 2 male ex., 78.879.7 mm SL; Baram, coll. C. Hose. — BMNH 1893.3.6:139, 1
female ex., 52.0 mm SL; Poeh, coll. A. Everett. — BMNH
1893.3.6:141-144, 4 ex., 49.5-67.5 mm SL; north Borneo, coll. A.
Everett. — BMNH 1894.8.3:28-30, 3 ex., 52.1-54.4 mm SL; Baram
River, coll. C. Hose. — BMNH 1895.7.2:43, 1 male ex., 61.1 mm
SL; Baram district, coll. C. Hose. — ZRC 37993, 1 ex., 37.9 mm
SL; Marudi Airport-Lubok Nibong road, at 12.1 km, coll. M. Kottelat& T. H. T. Tan, 19 Jun.1994. — ZRC 37967, 1 ex., 52.5 mm SL;
Sungai Tebu at 8 km, Daro-Matu road, coll. M. Kottelat & T. H.
T. Tan, 14 Jun.1994. — ZRC 37972, 1 ex., 52.2 mm SL; Parit
Nyadok, 200 m after 10 km stone, Daro-Matu road, coll. M. Kottelat& T. H. T. Tan, 14 Jun.1994. — ZRC 37858, 3 ex., 19.3-56.1 mm
SL; Sungai Nibong, ca. 1 km north of Durin ferry on Sri AmanSibu road, coll. M. Kottelat et al., 15 May.1994. — ZRC 37929, 2
ex., 39.5-50.9 mm SL; outskirts of Sibu, Sungai Teku, 5.2 km northbFig. 17. a) Betta akarensis - BMNH 1895.7.2:44, holotype, 31.6mm SL, b) ZRC 40928, 51.8 mm SL.

67

Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei
33085, 5 ex., 28.4-50.4 mm SL; Belait District, two streams nearold padi fields ca. 200m downstream of Kampung Melilas, coll. H.
H. Tan et al., 12 May.1996. — ZRC 40936, 2 ex., 23.6-53.7 mm
SL; Bandar Seri Begawan, Mentiri waterfalls, Sungai Mengkubau
downstream and tributaries (4°57’54.8”N 115°1’7.8”E), coll. H.H.
Tan et al., 14 May.1996. — ZRC 40937, 39 ex., 18.9-73.8 mm SL;
Tutong District, Sungai Ratuan Uluh, ca. 200m behind basecamp(Pankalang Kabo) at Tasik Merimbun (4°34’52.9”N
114°41’24.4”E), coll. H. H. Tan et al.,15 and 18 May.1996. — ZRC
40938, 13 ex., 24.8-63.9 mm SL; Tutong District, Sungai Siong
Mitom, tributary to Tasik Merimbun (4°35’16.3”N 114°40’42.5”E),
coll. H. H. Tan et al., 16 May.1996. — ZRC 40939, 1 ex., 37.4 mm
SL; Tutong District, Sungai Merimbun, outflow from Tasik
Merimbun (4°36’5.7”N 114°40’41.1”E), leads to Sungai Tutong,coll. H. H. Tan et al.,16-17 May.1996. — ZRC 40940, 1 ex., 32.3mm SL; Tutong District, Logon Uluh Boyoh, Bawang Ulop
(4°36’33.7”N 114°26’55.2”E), coll. H. H. Tan et al., 17 May.1996.
— ZRC 40941, 41 ex., 19.1-69.0 mm SL; BRM uncat., 10 ex., 28.151.4 mm SL; FMNH 105717, 10 ex., 26.5-54.9 mm SL; Belait
District, Sungai Mau tributary, draining out of peatswamp forest(4°33’41.5”N 114°29’41.7”E), coll. H. H. Tan et al., 19 May.1996.

Description. – General appearance is shown in Figs. 17a-b,
46f. Body relatively stout; caudal fin lanceolate; dorsal andanal fins pointed; pelvic fin filamentous. Mature males from
Brunei (ZRC 35437, 35439) with concave head profile, but
only for large males. Large males from Sarawak without
concave head profile. Vertebrae 2 + 8-9 + 20-21 = 31-32(mode 31, n = 11). Meristic and morphometric data in Table
4. Maximum known size 79.7 mm SL (BMNH 1891.1.27:
26-27).
Coloration. – For live coloration, see Fig. 46f (see Kubota
et al., 1996, identified as B. climacura). Fig. 7 shows
schematic head drawings with opercle patterns. Mature
malewith iridescent blue or green posterior half of bodyscales; pelvic fin with a white tip; caudal-fin transverse bars
distinctly present in males; black distal margin on anal fin.
Female with less iridescence on body and without caudal-fin
transverse bars. During mating, female with central andsecond central stripes on body converging into one stripe
ending on caudal peduncle.

Diagnosis. – Betta akarensis is distinguished from the othermembers of the B. akarensis species group by the following
combination of characters: opercle with uninterrupted second
postorbital stripe; yellow eye when alive; anal-fin rays 2830 (mode 28); subdorsal scales 5-6 (mode 5); lateral scales
31-33 (mode 32); predorsal scales 22-25 (mode 23);
postdorsal scales 10-12 (mode 11); preanal length 43.4-48.5%
SL; head length 29.8-34.9% SL; length of anal-fin base 49.656.8% SL.

Distribution. – Betta akarensis occurs in Brunei (Bandar Seri
Bagawan, Belait and Tutong Districts) and Sarawak (Miri,
Marudi, Sungai Akar, Sungai Tebu, Parit Nyadok, Sungai
Nibong, Sungai Teku, Mukah) (Fig. 18).
Comparative notes. – Betta akarensis is distinguished fromthe other members in the species group in having: opercle
with uninterrupted second postorbital stripe (Fig. 7a); yellow

Fig. 18. Distribution map of B. akarensis (circle), B. balunga (triangle), B. chini (square), B. pinguis (asterix), B. aurigans (diamond), B.
ibanorum (upside-down triangle), B. obscura (star).

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
eye when alive (vs. orange in B. balunga); slightly morevertebrae than B. chini (total 31-32, mode 31, vs. total 2931, mode 30); more lateral scales than B. balunga (31-33,
mode 32, vs. 29-30); dorsal-fin origin above 16-18th lateralscale (vs. 18-19th in B. pinguis); greater postdorsal lengththan B. pinguis (19.9-25.1% SL, vs. 18.0-20.9); smaller
preanal length than B. balunga (43.4-48.5% SL, vs. 49.252.2); smaller body depth at dorsal-fin origin than B. pinguis(25.1-30.9% SL, vs. 30.6-31.6); greater orbital diameter thanB. pinguis (24.5-32.3% HL, vs. 22.9-25.0); relatively smaller
lower jaw length than B. ibanorum (8.8-10.7% SL vs. 9.312.0; 25.4-34.4% HL vs. 28.0-36.1).

scales 21-24; postdorsal scales 10-12; preanal length 48.552.6% SL; head length 32.6-37.5% SL; length of anal-finbase 47.8-55.4% SL.
Description. – General appearance is shown in Figs. 19, 46g.
Body relatively stout; caudal fin lanceolate, dorsal and anal
fins pointed, pelvic fin filamentous. Vertebrae 2 + 8-9 + 1921 = 30-31 (n=2). Meristic and morphometric data in Table
4. Maximum known size 49.3 mm SL (CAS/SU 33203).
Coloration. – See Fig. 46g for the live coloration of theKalimantan specimen (see also Kubota et al., 1996: 45). Males
have both dorsal and caudal transverse bars. For comparisonof opercle patterns of members of the B. akarensis group,
see Fig. 7. The specimens from Kalimantan retain a slightyellow body coloration when preserved (ZRC 35497); andin life, had iridescent blue on body scales.

Remarks. – Please refer to Tan & Ng (2004a) for more details.

Betta balunga Herre, 1940
(Figs. 19, 46g)

Distribution. – Betta balunga is found in Tawau, Sabah andthe Sebuku and Mahakam basins, Kalimantan Timur (Inger& Chin, 1962; Ng, 1993; unpublished data) (Fig. 18).

Betta balunga Herre, 1940: 44; Inger & Chin, 1962: 157; Witte &Schmidt, 1992: 324 (key only); Kottelat et al., 1993: 161(comments only); Ng, 1993: 292 (discussion).

Field notes. – The Sebuku specimens were caught from forest
streams with moderate current, in very shallow water (5-10cm deep) amongst plant roots and submerged leaf litter along
the banks. Betta albimarginata was also caught from the same
stream (M. Kottelat, pers. comm.). Betta balunga is an oral
brooder (Rehwinkel, 1995). The Mahakam specimens wereobtained slow flowing black water streams, from amongst
submerged stream bank vegetation and submerged leaf litter.

Material examined. – Holotype - Sabah: - CAS/SU 33203, 49.3mm SL; Borneo: Sabah, Tawau, Sungai Balung, coll. A. W. C. T.
Herre, 23 Jan.1937.
Others - CAS 17361, 2 ex., 33.3-46.7 mm SL; Borneo: Sabah,
Tawau, Sungai Balung, coll. A. W. C. T. Herre, 1937. Kalimantan
Timur: ZRC 35497, 1 ex., 37.2 mm SL, CMK 9451, 8 ex., 17.233.4 mm SL; Borneo: Kalimantan Timur, Sebuku basin, near
Pembeliangan, creek along road from base camp to Semunad,draining into Sungai Tikung (4°0’48”N 117°2’54”E), coll. M.
Kottelat & P. McKee, 10 Feb.1993.

Comparative notes. – Betta balunga is distinguished fromthe other members of the species group in having: reddishorange iris of eye (vs. light yellow or green when alive); more
subdorsal scales than B. pinguis (6-7, vs. 5-51/2, mode 51/2);
fewer lateral scales than B. akarensis and B. pinguis (29-30,
vs. 31-33); dorsal-fin origin above 15-16th lateral scale (vs.
18-19th in B. pinguis); anal-fin origin below 6-61/2th lateralscale (vs. 7th in B. pinguis); smaller total length than B.
pinguis (132.8-141.3% SL, vs. 145.5-153.5); greater preanallength than B. akarensis, B. balunga, B. chini and B. ibanorum(48.5-52.6% SL, vs. 42.6-49.4); greater head length than B.
pinguis (32.6-37.5% SL, vs. 31.3-31.7); smaller length ofanal-fin base than B. chini and B. pinguis (47.8-55.4% SL,
vs. 54.5-58.5); greater lower jaw length than B. pinguis (10.210.4% SL, vs. 9.3-9.9); greater orbital diameter than B.
pinguis (25.4-33.3% HL, vs. 22.9-25.0).

Betta cf. balunga - ZRC 46332, 3 ex., 31.9-42.3 mm SL; Kalimantan
Timur: Mahakam basin; black water stream along road km 124 from
Kota Bangun to Samarinda (km 9.5 to Kota Bangun) (00º14.60’S
116º38.48’E), coll. H. H. Tan & D. Wowor, 10 Nov.1999. — ZRC
46333, 50 ex., 9.2-43.4 mm SL; Kalimantan Timur: Mahakam basin;clear water stream 10 km before Muara Badak (00º19.62’S
117º20.93’E), coll. H. H. Tan & D. Wowor, 11 Nov.1999.

Diagnosis. – Betta balunga is distinguished from the othermembers of the B. akarensis group by the following
combination of characters: opercle with interrupted second
postorbital stripe; eye colour orange when alive; anal-fin rays
27-30; subdorsal scales 6-7; lateral scales 29-30; predorsal

Remarks. – Herre (1940) described B. balunga on the basisof one specimen (CAS/SU 33203) of 49.3 mm SL. The useof the presence of spines in anal or dorsal fin and small
differences in the number of spines in fin ray counts todistinguish between species as used by Herre (1940) is now
regarded as untenable (see Tan & Tan, 1996). The materialof B. balunga is scarce and more material is required before
the species status can be ascertained (Kottelat et al., 1993:
161). The specimens from the Mahakam drainage differ fromthe holotype in the following aspects: fewer anal-fin rays (27vs. 30); fewer predorsal scales (21 vs. 23); smaller predorsal
length (66.7% SL, vs. 71.7); and smaller length of anal-fin

Fig. 19. Betta balunga – CAS/SU 33203, holotype, 49.3 mm SL(taken by Susan Middleton).

69Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei
base (47.8% SL, vs. 55.4). The significance of thesedifferences, however, cannot be ascertained due to the low
number of specimens from the type locality and its immediate
vicinity. For the time being, these specimens are regarded asconspecific (see Ng, 1993: 292).

shallow water (less than 50 cm depth). This species is an oral
brooder (Dickman, 1995).
Comparative notes. – Betta chini differs from the othermembers of the species group in having: yellow eye when
alive (vs. orange of B. balunga); more subdorsal scales thanB. akarensis (51/2-7, mode 6, vs. 5-6, mode 5); dorsal-fin
origin above 16-17th lateral scale (vs. 18-19th in B. pinguis);fewer vertebrae than B. akarensis (total 29-31, mode 30, vs.
total 31-32, mode 31); smaller total length than B. pinguis(137.2-144.6% SL, vs. 145.5-153.5); greater postdorsal lengththan B. pinguis (19.9-23.3% SL, vs. 18.0-20.9); smaller
preanal length than B. balunga and B. ibanorum (42.6-45.7%
SL, vs. 45.8-52.2); smaller body depth at dorsal-fin origin
than B. pinguis (26.3-29.0% SL, vs. 30.6-31.6); greater lengthof anal-fin base than B. balunga and B. ibanorum (54.8-58.5%
SL, vs. 47.8-55.8); greater orbital diameter than B. pinguis(26.2-30.8% HL, vs. 22.9-25.0); smaller snout length than B.
pinguis (20.0-24.4% HL, vs. 25.0-27.8); smaller height at
pectoral-fin origin than B. pinguis (47.6-59.3% HL, vs. 58.162.0). Fig. 7 shows schematic head drawings with the opercle
patterns of B. akarensis, B. balunga, B. chini, B. ibanorum
and B. obscura.

Betta chini Ng, 1993
(Figs. 20, 46h)
Betta chini Ng, 1993: 290, figs. 1-3; Kottelat & Ng, 1994: 74.
Material examined. – Holotype - Sabah: - ZRC 35086, 51.0 mm
SL male; Borneo: Sabah, Beaufort area, peat swamps, along logging
trail, ca. 12 km from Beaufort, on road from Kota Kinabalu to
Beaufort (5°33’06"N 115°50’23"E), coll. P. K. L. Ng & R. B.
Stuebing, 29 Dec.1992.
Paratypes - ZRC 35087-093, 7 ex., 17.4-54.2 mm SL; - SBM uncat.,2 ex., 31.0-33.3 mm SL; same locality data as holotype. — ZRC
35096-35102, 7 ex., 22.8-45.5 mm SL; Borneo: Sabah, Beaufort
area, Sungai Mawao, peat swamps, coll. R. B. Stuebing, 11 Jan.1993.
— ZRC 35094-95, 2 ex., 25.4-47.3 mm SL; Borneo: Sabah, Beaufort
area, Kampung Lumat, coll. R. B. Stuebing, 7 Jan.1993.

Diagnosis. – Betta chini is distinguished from the othermembers of the B. akarensis group by the following
combination of characters: opercle with interrupted second
postorbital stripe; eye colour yellow when alive; anal-fin rays
26-30 (mode 28); subdorsal scales 51/2-7 (mode 6); lateralscales 30-31 (mode 31); predorsal scales 22-24 (mode 23);
postdorsal scales 9-11 (mode 10); preanal length 42.6-45.7%
SL; head length 31.4-33.3% SL; length of anal-fin base 54.858.5% SL.

Remarks. – Betta chini is one of the smallest species of theB. akarensis group. Ng (1993) has provided a detailed
comparison with the other members of the B. akarensis group.

Betta ibanorum Tan & Ng, 2004
(Figs. 21, 47a)
Betta ibanorum Tan & Ng, 2004: 273, figs. 3, 7; Grinang & Lim,
2004: 292.
Betta anabatoides - Regan, 1910: 780, pl. 78, fig. 4 (part).
Betta akarensis (non Regan, 1910) - Witte & Schmidt, 1992: 326(part); Kottelat et al., 1993: 161, pl. 75 (part); Ng, 1993: 290(part); Kottelat & Ng, 1994: 74 (part); Kottelat & Lim, 1995:
248 [part].
Betta climacura (non Vierke, 1988) - Kottelat et al., 1993: 161 (part);Ng, 1993: 289 (part); Kottelat & Ng, 1994: 74 (part).
Betta spec. affin. taeniata (non Regan, 1910) - Brown & Brown,
1987: 169, pl. 13.

Description. – General appearance is shown in Figs. 20, 46h.
Body relatively stout, caudal fin lanceolate; dorsal and anal
fins pointed; pelvic fin filamentous. Vertebrae 2 + 8 + 19-21= 29-31 (mode 30, n = 8). Meristic and morphometric data
in Table 4. Maximum known size 54.2 mm SL (ZRC 35087).
Coloration. – See Fig. 46h for live coloration and Ng (1993)
for more details (see also Kubota et al., 1996: 46).
Distribution. – Betta chini is apparently restricted to the peatswamps of Beaufort in Sabah (Ng, 1993) (Fig. 18).

Material examined. – Holotype - Sarawak: - ZRC 41584, 65.0 mm
SL male; Borneo: Sarawak, Kuching, Bako National Park, Bukit
Gondol, coll. N. Sivasothi et al., 28 Jun.1994.

Field notes. – Betta chini was collected from badly disturbed
and logged peat swamps. Specimens were collected from

Paratypes - ZRC 37750, 2 males, 53.9-65.1 mm SL, 1 female, 43.8mm SL, same locality data as holotype. — ZRC 37757, 22 ex., 9.4-

Fig. 20. Betta chini – ZRC 35086, male holotype, 51.0 mm SL.

Fig. 21. Betta ibanorum – ZRC 41584, male holotype, 65.0 mm SL(right side, reversed).

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
76.1 mm SL; Borneo: Sarawak, Kuching, Bako National Park, Tajor
waterfalls, coll. T. H. T. Tan & A. T. C. Wong, 29 Jun.1994.

1991: 29, identified as B. climacura). Body dark brown
dorsally, lighter ventrally. Mature male with iridescent green
to blue posterior half on body scales, dorsal and caudal
transverse bars. Female with possess less iridescence andwithout caudal transverse bars. Juvenile with a distinct black
central stripe on body and a black caudal peduncle spot. A
narrow distinct chin-bar in all specimens. Opercle with little
or no pattern (Fig. 7).

Others - ZRC 37764, 7 ex., 17.1-51.7 mm SL; Borneo: Sarawak,
Kuching, Bako National Park, Sungai Serait at goldmine, coll. N.
Sivasothi et al., 30 Jun.1994. — ZRC 37755, 3 juv., 16.2-18.3 mm
SL; Borneo: Sarawak, Kuching, Bako National Park, Sungai Nipah,
coll. D. S. L. Chung et al., 29 Jun.1994. — ZRC 39273, 2 males,
75.3-77.4 mm SL, 1 female, 66.1 mm SL; Borneo: Sarawak, Bako
National Park, coll. NUS Honours class 1994-95 (kept in aquaria),
Jun.1994. — ZRC 35435-36, 2 males, 71.5-80.8 mm SL; Borneo:
Sarawak, blackwater stream at km-stone 7 from Kuching to Batu
Kawa (1°31’N 110°18’E), coll. P. K. L. Ng et al., 3 Jul.1992. —ZRC 27830-35, 6 ex., 12.4-75.3 mm SL; Borneo: Sarawak, 7 km
Kuching to Batu Kawa road, coll. P. K. L. Ng et al., 3 Jul.1992. —ZRC 37838, 4 juv., Borneo: Sarawak, Matang Wildlife Centre, small
forest creek 100 m beyond Sungai Rayu, coll. M. Kottelat et al., 5
May.1994. — ZRC 39584, 13 ex., 22.9-50.2 mm SL; Borneo:
Sarawak, Batu Kawa-Matang area, Taman Koperkasa (1°34’42.0"N
110°16’24.7"E), ca. 2 km after bridge over blackwater river, ca. 10km after Kuching town, coll. P. K. L. Ng et al., 14 Jan.1996. —ZRC 39585, 2 ex., 24.5-32.9 mm SL; Borneo: Sarawak, Batu KawaMatang area (1°34’27.1"N 110°17’34.9"E), ca. 50 m before
blackwater river in flooded peat swamp forest, access via first righttrack after bridge over blackwater river, coll. P. K. L. Ng et al., 14
Jan.1996. — ZRC 39586, 1 ex., 27.2 mm SL; Borneo: Sarawak, ca.
50 m into peatswamp forest and ditch on left side of road
(1°10’50.0"N 110°39’22.1"E), ca. 8 km towards Gedong after
turnoff from Serian-Sri Aman road, coll. H. H. Tan & S. H. Tan,
16 Jan.1996. — ZRC 40942, 33 ex., 13.6-70.5 mm SL; SM uncat.,
5 ex., 31.1-54.7 mm SL; CMK 13123, 5 ex., 30.0-63.9 mm SL;
Borneo: Sarawak, peatswamp forest, 11.4 km towards Gedong fromturnoff on Serian-Sri Aman road (1°12’07.8”N 110°39’51.2”E), coll.
P. K. L. Ng et al., 31 Aug.1996. — ZRC 40943, 20 ex., 25.2-50.4mm SL; Borneo: Sarawak, peatswamp forest, 38.1 km towards
Simunjan, from Kuching-Sri Aman road (1°18’55.3”N
110°51’43.6”E), coll. H. H. Tan & S. H. Tan, 4 Sep.1996. — ZRC
40944, 27 ex., 12.0-64.8 mm SL; SM uncat., 5 ex., 29.0-50.8 mm
SL; CMK 13124, 5 ex., 22.7-53.2 mm SL; Borneo: Sarawak, watercascade from Gunung Santubong, ca. 1 km before Damai Beach
Resort (1°44’10.1”N 110°19’8.6”E), coll. H. H. Tan & S. H. Tan,
5 Sep.1996.

Distribution. – Betta ibanorum is found in Bako National
Park, in the adjacent areas around Kuching and appears to be
restricted to southern Sarawak (Fig. 18). It is confined east
towards Bau area (e.g. Matang, Serian, Gedong, Simunjan,
Sri Aman) and is replaced by B. lehi in the area west of Bau(e.g. Lundu).
Comparative notes. – Betta ibanorum differs from the othermembers of the species group in having: opercle without
second postorbital stripe (Fig. 7); yellow eye when alive (vs.
orange of B. balunga); largest adult size (80.8 mm SL); dorsalfin origin above 15-17th lateral scale (vs. 18-19th in B.
pinguis); fewer predorsal scales than B. pinguis (21-23, mode
22, vs. 23-24, mode 24); greater preanal length than B. chini(45.8-49.4% SL, vs. 42.6-45.7); smaller preanal length thanB. balunga (45.8-49.4% SL, vs. 49.2-52.2); greater headlength than B. pinguis (31.7-36.4% SL, vs. 31.3-31.7); smaller
body depth at dorsal-fin origin than B. balunga and B. pinguis(23.3-28.1% SL, vs. 28.0-31.6); smaller length of anal-finbase than B. chini and B. pinguis (50.0-55.8% SL, vs. 54.558.5); smaller orbital diameter than B. balunga (22.2-30.5%
HL, vs. 29.5-33.3); greater snout length than B. balunga (22.929.5% HL, vs. 20.0-21.2). Betta ibanorum can bedifferentiated from B. aurigans by the following characters:
blueish iridescence on scales (vs. gold scales); fewer analfin rays (27-29, mode 28, vs. 29-30, mode 30); fewer lateralscales (30-32, mode 30, vs. 33-331/2, mode 331/2); fewer
predorsal scales (21-23, vs. 24-26); fewer postdorsal scales(10-11, mode 11, vs. 11-13, mode 13); and smaller length ofanal –fin base (50.0-55.8, vs. 55.4-58.1% SL). Fig. 7 shows
schematic head drawings with the opercle patterns of B.
akarensis, B. balunga, B. chini and B. ibanorum.

Diagnosis. – Betta ibanorum is distinguished from the othermembers of the B. akarensis group by the following
combination of characters: opercle without second postorbital
stripe; eye colour yellow when alive; anal-fin rays 27-29(mode 28); subdorsal scales 5-6 (mode 51/2); lateral scales
30-32 (mode 30); predorsal scales 21-23 (mode 22);
postdorsal scales 10-11 (mode 11); preanal length 45.8-49.4%
SL; head length 31.7-36.4% SL; length of anal-fin base 50.055.8% SL.

Remarks. – For more details, please refer to Tan & Ng (2004).

BETTA UNIMACULATA GROUP
Betta macrostoma Regan, 1910
(Figs. 22a-b, 47b-c)

Description. – General appearance is shown in Figs. 21, 47a.
Body relatively slender, caudal fin broadly lanceolate; dorsal
and anal fins pointed; pelvic fin filamentous. Large male witha concave head profile. Rays of unpaired fins of males
elongated, and may appear filamentous. Vertebrae 2 + 9 +19-21 = 30-32 (mode 31, n = 9). Meristic and morphometricdata in Table 4. Maximum known size 80.8 mm SL (ZRC
35435).

Betta macrostoma Regan, 1910: 778, pl. 78, Fig. 3; Weber & deBeaufort, 1922: 355; Herre, 1940 (key only); Witte & Schmidt,
1992 (key only); Kottelat et al., 1993: 162, pl. 76; Kottelat &Lim, 1995: 248.
Material examined. – Holotype - North Sarawak: BMNH
1898.11.2:2, male, 60.0 mm SL; Borneo, coll. Hose, 1898.
Others – Brunei: ZRC 31793-94, 1 male - 69.8 mm SL, 1 female- 59.0 mm SL; Borneo: Brunei, Belait District, Sungai Rampayoh,coll. S. C. Choy, Apr.1992. ZRC 40945, 8 ex., 9.0-59.7 mm SL;

Coloration. – For live coloration, refer to Fig. 47a and Kottelat
et al. (1993: pl. 75, identified as B. akarensis) (see Linke,

71Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei
Borneo: Brunei, Belait District, Implasi Batu, small hill stream ca.
750 m after the hot spring, coll. H. H. Tan et al., 10 May.1996. —ZRC 40987, 1 juv., 14.6 mm SL; Belait District, Sungai Jaung,
stream below hot spring, coll. H. H. Tan et al., 10 May.1996. —ZRC 40946, 8 ex., 14.7-67.0 mm SL; BRM uncat., 2 ex., 55.8-56.5mm SL; Borneo: Brunei, Belait District, Labi, Rampayoh, Sungai
Mendaram (entrance to falls: 4°20’30.7”N 114°26’55.2”E; cascade:4°20’20.0”N 114°27’7.7”E), coll. H. H. Tan et al., 19 May.1996.

in Belait district, Brunei. They were found in shallow (ca. 30
cm deep) near stagnant pools along a clayey stream bed. There
were only an adult pair or an adult with several juveniles ineach pool sampled. There were no other fish in this locality.
The pH was 5.7.
Betta macrostoma was also collected at the Mendaram
waterfalls, Rampayoh in Labi area, Belait district. The terrain
was rocky, with fast flowing clear water, pH 4.4. Dipteris
fern (Polypodiaceae) was growing along the banks of thestreams. Betta macrostoma was in small pools in tributaries
and in the quiet pools along the main confluence of the river.
Only one adult or several juveniles could be located in eachof these microhabitats. The only other fish was Rasbora tubbi(Cyprinidae). Betta macrostoma is an oral brooder (Ettrich,
1992; Kubota et al., 1996: 54).

Diagnosis. – Betta macrostoma is distinguished from theother members of the B. unimaculata group group by thefollowing combination of characters: dorsal fin ocellus (Fig.
8a); red body colour; transverse black and red bands on caudal
fin; opercle with large red blotch in mature male; vertebrae
31-32 (mode 32); dorsal-fin rays 9-11 (mode 10), anal-fin
rays 25-28 (mode27); pectoral-fin rays 13-14; lateral scales
32-33 (mode 33); subdorsal scales 6-8 (mode 7); predorsal
scales 21-24 (mode 23); predorsal length 62.3-66.7% SL; headlength 30.4-33.3% SL; length of anal-fin base 43.2-49.2%
SL; length of dorsal-fin base 12.2-15.7% SL; lower jaw length38.0-54.0% HL.

Distribution. – Betta macrostoma is presently known from
Belait District, Brunei Darussalam; and has been reported
from the area around Marudi, North Sarawak (pers. observ.)(Fig. 23).

Description. – General appearance is shown in Figs. 22a-b,
47b-c. Body slender; head large; caudal, dorsal and anal finsrounded; pelvic fin falcate with short filamentous soft ray.
Vertebrae 2-3 + 8 + 21-22 = 31-32 (mode 32, n=11). Meristic
and morphometric data in Table 5. Maximum known size
67.0 mm SL (ZRC 40946).

Comparative notes. – Betta macrostoma is distinguished fromthe other members of the species group by the following
characters: dorsal fin ocellus (vs. absence) (Fig. 8a);transverse black and red bands on caudal fin (vs. absence);
opercle with large red blotch in mature males (vs. absence);
more dorsal-fin rays (9-11 vs. 6-9), fewer anal-fin rays (2528 vs. 27-31); more subdorsal scales (6-8 vs. 4-61/2); dorsalfin origin above 15-17th lateral scale (vs. 17-21st); smaller
predorsal length (62.3-66.7% SL vs. 67.7-73.0); smaller
length of anal-fin base (43.2-49.2% SL vs. 45.9-54.5); greater
length of dorsal-fin base (12.2-15.7% SL vs. 7.3-11.4); greater
lower jaw length (38.0-54.0% HL vs. 27.3-38.3).

Coloration. – For live coloration, refer to Figs. 47b-c (seealso Linke, 1991: 38; Kubota et al., 1996). Mature male withbright red body, red opercle blotch surrounded by black,ocellus on dorsal fin (Fig. 8a) and transverse black and redbars on caudal fin, anal fin red with black distal margin, pelvic
fin with white tip. Females and juveniles with black central
and second central stripes on body and a black caudal
peduncle spot (Fig. 1). Preserved male specimens still withsome red on body, mostly faded to yellow and light brown.
Black stripe pattern on a light brown body distinct inpreserved female and juvenile.

Remarks. – Betta macrostoma is a very distinctly coloured
species with long jaws that extend beyond orbital towards
the opercle in mature male. There are a few populations inwhich the dorsal fin ocellus is present (Regan, 1910;
Yamazaki, 1995) and some others apparently absent (Linke,
1991; Kottelat et al., 1993). The intensity of coloration,
however, differs among populations. Tentatively, allpopulations are regarded as conspecific.

Field notes. – Betta macrostoma was obtained in clear watersa

Betta ocellata de Beaufort, 1933
(Figs. 24a-b, 47d-e)
Betta ocellata de Beaufort, 1933: 35; Herre, 1940: 43; Inger & Chin,
1962: 159; Witte & Schmidt, 1992: 327; Martin-Smith & Tan,
1998: 593.
Betta unimaculata: (non Popta, 1905) - Inger, 1955: 70; Inger &bChin, 1962: 159; Lim & Wong, 1994: 62; Chin & Samat,
1995: 29.
Material examined. – Holotype - Sabah: - BMNH 1959.7.7:1, 70.0
mm SL; Borneo: Sabah, Sandakan, Bettotan.
Others - BMNH 1893.5.30:52, 1 ex., 50.1 mm SL; Borneo: Sabah,
Bongon, coll. A. Everett. — ZRC 37642, 26 ex., 23.4-60.0 mm SL;
Borneo: Sabah, Kampung Batu Puteh (Kinabatangan basin), shaded

Fig. 22. a) Betta macrostoma - BMNH 1898.11.2:2, male holotype,60.0 mm SL, b) ZRC 40945, 59.8 mm SL.

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
forest stream in SAFODA, coll. K. K. P. Lim et al., 7-13 Apr.1994.
— ZRC uncat., 3 ex., 62.1-68.9 mm SL; Borneo: Sabah, Kampung
Batu Puteh (Kinabatangan basin), shaded forest stream in SAFODA,
coll. K. K. P. Lim et al., 7-13 Apr.1994 (kept in aquaria for oneyear). — FMNH 44901, 5 ex., 15.6-46.5 mm SL; Borneo: Sabah,
Sandakan, Sandala Estate, North road, coll. Fisheries Department,
8 Oct.1948. — FMNH 44902, 4 ex., 28.0-50.8 mm SL; Borneo:
Sabah, Sandakan, mile 6 on North road, coll. J. A. Tubb, 25 Jul.1950.
— FMNH 44903, 1 ex., 42.2 mm SL; Borneo: Sabah, Sandakan,mile 7 on North road, coll. Fisheries Department, 31 Jul.1950. —FMNH 44904, 2 ex., 43.1-46.7 mm SL; Borneo: Sabah, Sandakan,
Tenosa, coll. J. A. Tubb, 9 Dec.1947. — FMNH 51634, 10 of 16ex., 33.9-55.0 mm SL; Borneo: Sabah, Kinabatangan, clear water
tributary of Sungai Kretam Kechil below falls, coll. R. F. Inger, 11
May.1950. — FMNH 51635, 5 of 9 ex., 43.6-53.8 mm SL; Borneo:
Sabah, Kinabatangan, southeast end of Dewhurst Bay, west fork ofGaja River above rapids, coll. R. F. Inger, 30 May.1950. — FMNH
51638, 5 of 11 ex., 38.8-58.5 mm SL; Borneo: Sabah, Kinabatangan,
southeast end of Dewhurst Bay, west fork of Gaja River, 0.25 miles
upstream from station 11, coll. R. F. Inger, 3 Jun.1950. — FMNH
51639, 5 ex., 26.8-61.7 mm SL; Borneo: Sabah, Sandakan, tributaryof Sapagaya River, coll. R. F. Inger, 16 Jul.1950. — FMNH 51641,
9 ex., 12.1-67.8 mm SL; Borneo: Sabah, Kinabatangan, southeast
end of Dewhurst Bay, tributary of Sungai Kretam Kechil, coll. R.
F. Inger, 15 Jun.1950. — FMNH 68430, 5 of 10 ex., 54.1-79.8 mm
SL; Borneo: Sabah, Sandakan, Sepilok Forest Reserve, coll. R. F.
Inger, 9 Apr.1956. — FMNH 68435, 5 of 13 ex., 48.6-67.4 mm SL;
Borneo: Sabah, Kinabatangan, Deramakot camp, hill stream below
falls, coll. R. F. Inger & P. K. Chin, 2 May.1956. — FMNH 68436,
10 ex., 43.7-71.0 mm SL; Borneo: Sabah, Kinabatangan, Deramakot
camp, below waterfall, coll. R. F. Inger, 3 May.1956. — FMNH
68439, 5 ex., 29.3-60.4 mm SL; Borneo: Sabah, Tawau, Pulau
Sebatik, coll. R. F. Inger, 28 May.1956. — FMNH 68440, 1 ex.,32.8 mm SL; Borneo: Sabah, Tawau, Kalabakan, Sungai Tibas camp,
Sungai Tawan, coll. R. F. Inger, 6 Jun.1956. — FMNH 68441, 1
ex., 24.5 mm SL; Borneo: Sabah, Tawau, Kalabakan, Sungai Tibas
camp, ca. 1 km upstream of Sungai Tawan, coll. R. F. Inger, 8
Jun.1956. — FMNH 68442, 2 ex., 18.1-30.4 mm SL; Borneo: Sabah,
Tawau, Brantian River estate, semi-isolated pool, coll. R. F. Inger,
8 Jul.1956. — ZRC 40947, 6 ex., 35.1-56.8 mm SL; Borneo: Sabah,
Lahad Datu, Danum Valley, Kallang Sebaru stream, tributary of
Sepat Kalisun stream, km 64.5 from main road to Danum Valley
Field Centre (4°58’04.8”N 117°48’56.5”E), coll. H. H. Tan et al.,1 Oct.1996. — ZRC 40948, 3 ex., 21.4-44.9 mm SL; Borneo: Sabah,
Lahad Datu, Danum Valley, Sepat Kalisun stream, coll. H. H. Tan
et al., 1 Oct.1996. — ZRC 40949, 1 ex., 42.7 mm SL; Borneo: Sabah,
Lahad Datu, Danum Valley, Sungai Paulm Tambun, tributary ofSungai Segama, upstream of Danum Valley Field Centre, coll. H.
H. Tan et al., 1 Oct.1996. — ZRC 40950, 1 ex., 59.1 mm SL; Borneo:
Sabah, Lahad Datu, Danum Valley, small tributary of Sungai Bole,ca. 500 m into coupe 93 (4°58’17.8”N 117°51’48.0”E), coll. H. H.
Tan et al., 2 Oct.1996. — ZRC 40951, 7 ex., 23.3-40.5 mm SL;
Borneo: Sabah, Lahad Datu, Danum Valley, cabin stream right, km
50 on road to Danum Valley Field Centre, drains from Bukit
Rafflesia (4°59’08.5”N 117°54’05.1”E), coll. H. H. Tan et al., 2
Oct.1996. — ZRC 40952, 3 ex., 33.7-50.4 mm SL; Borneo: Sabah,
Lahad Datu, Danum Valley, West six, forest stream 600 m into
conservation area, tributary of Sungai Segama, coll. H. H. Tan etal., 4 Oct.1996. — ZRC 40953, 4 ex., 34.4-60.3 mm SL; Borneo:
Sabah, Lahad Datu, Danum Valley, West eight, forest stream 800
m into conservation area, tributary of Sungai Segama, coll. H. H.
Tan et al., 4 Oct.1996. — ZRC 40954, 6 ex., 14.1-35.7 mm SL;
Borneo: Sabah, Tawau, stream by Air Panas, near base of Tawau
Hills park (4°18’03.0”N 117°54’20.7”E), coll. H. H. Tan et al., 5
Oct.1996. —- ZRC 40955, 3 ex., 19.6-52.4 mm SL; Borneo: Sabah,
Tawau, Jalan Sin On Jaya, swampy area (4°16’07.6”NFig. 23. Distribution map of B. macrostoma (circle), B. ocellata (triangle), B. gladiator (upside-down triangle), B. unimaculata (square),B. patoti (star), B. pallifina (diamond).

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Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei
117°54’25.6”E), coll. H. H. Tan et al., 5 Oct.1996. — ZRC 40956,
70 ex., 9.7-59.4 mm SL; SBM uncat., 10 ex., 20.9-42.7 mm SL;
ZMA 121.677, 10 ex., 24.7-46.9 mm SL; BMNH 1997.4.3.6-15,
10 ex., 26.3-45.8 mm SL; RMNH 33086, 10 ex., 26.8-45.4 mm SL;
FMNH 105718, 10 ex., 27.5-41.9 mm SL; Borneo: Sabah, Tawau,
stream at km 76 to Semporna, km 32 to Tawau (4°20’17.4”N
118°05’32.3”E), coll. H. H. Tan et al., 5 Oct.1996. — ZRC 40957,
52 ex., 11.7-63.7 mm SL, CMK 13016, 5 ex., 40.0-48.2 mm SL;
Borneo: Sabah, Tawau, stream at km 77 to Semporna, km 31 toTawau (4°19’37.8”N 118°05’36.8”E), coll. H. H. Tan et al., 5
Oct.1996. — ZRC 40958, 1 ex., 44.9 mm SL; Borneo: Sabah,
Tawau, Sungai Nasib Katib, ca. km 66.5 to Semporna, km 41.5 to
Tawau (4°22’41.2”N 118°07’55.6”E), coll. H. H. Tan et al., 6
Oct.1996. — ZRC 40959, 25 ex., 25.0-65.5 mm SL; Borneo: Sabah,
Tawau, Jalan Madai, Gua Madai (limestone outcrop), Sungai
Matarid runs through caves (4°43’08.7”N 118°09’14.7”E), coll. H.
H. Tan et al., 6 Oct.1996. Kalimantan Timur: CMK 9540, 1 ex.,24.2 mm SL; Borneo: Kalimantan Timur, Sungai Sebuku basin,
Sungai Sanul, tributary of Sungai Tikung, ca. km 7 on RMK logging
road from Semunad to Apas (4°2’25”N 116°58’46”E), coll. M.
Kottelat & P. McKee, 14 Feb.1993.

B unimaculata). Mature male with iridescent blue opercle
scales when alive, blackish when preserved, with a single
iridescent scale on upper region of head. Faintly iridescentscales on body; specimens from Kinabantangan basin withvery little iridescence; male from Tawau with full body
iridescence; specimens from Danum valley and Tawau withirregular black marks along flank. Body light to dark brownwhen alive, light brown when preserved. Caudal peduncle
spot below lateral scale series near end of hypural plate,present in both sexes and juveniles (Fig. 1). Female with very
few or no iridescent opercle scales. When alive, faint or
distinct dorsal, caudal and anal fin transverse bars; faint whenpreserved.
Distribution. – Betta ocellata is restricted to northeastern
Borneo, found only in Sabah (Bettotan, Sandakan,
Kinabatangan, Lahad Datu, Tawau and Bongon) and thenorthern part of East Kalimantan (Sebuku basin) (de Beaufort,
1933; Inger & Chin, 1962; pers. observ.; M. Kottelat, pers.
comm.) (Fig. 23).

Diagnosis. – Betta ocellata is distinguished from the othermembers of the B. unimaculata species group by thefollowing combination of characters: distinct black spot nearcaudal peduncle; body brownish; opercle blue; uniform caudal
fin coloration; vertebrae 33-34 (mode 33); anal-fin rays 3031 (mode 30); dorsal-fin rays 8; pectoral fin rays 12-13 (mode13); subdorsal scales 5-51/2 (mode 51/2); lateral scales 32-34(mode 33); predorsal scales 24; predorsal length 68.8-71.9%
SL; head length 31.4-33.3% SL; length of anal-fin base 52.254.5% SL; length of dorsal-fin base 10.5-11.3% SL; lower
jaw length 27.3-33.1% HL.

Field notes. – Specimens collected from Sabah were mainly
from headwater habitats. The fishes were usually obtained
from ravines and isolated elevated pools. A single adult andseveral juveniles are usually obtained from the submerged
leaf litter. Adults may take cover under floating leaves. They
are excellent jumpers, which may explain their presence inisolated elevated pools. Their diet consist mainly of terrestrial
insects and occassional aquatic invertebrates (Inger, 1955;
Inger & Chin, 1965).
Specimens collected within the Kinabatangan basin were fromlowland forest streams and ox-bow lakes. The water in thesehabitats was murky. This species inhabits the shallow partsof ox-bow lakes among the flooded grass banks and among
the submerged leaf litter and tree roots in the forest streams.
Other species collected from the forest streams included:
Rasbora sumatrana, Puntius sealei, P. banksi (Cyprinidae),
Pangio mariarum, Lepidocephalichthys sandakanensis(Cobitidae), Nemacheilus olivaceus (Balitoridae), Ompok
sabanus (Siluridae), Clarias teijsmanni (Clariidae) andTrichogaster trichopterus (Osphronemidae).

Description. – General body shape and appearance as in Figs.
24a-b, 47d-e. Body relatively stocky, broad but relatively
pointed head; caudal fin rounded, dorsal and anal finsrelatively pointed, pelvic fin filamentous. Mature males witha steep rise behind eye. Vertebrae 2 + 9 + 22-23 = 33-34(mode 33, n=13). Meristic and morphometric data in Table
5. Maximum known size 79.8 mm SL (FMNH 68430).
Coloration. – For live coloration, see Figs. 47d-e (see also
Linke, 1991: 73; Kubota et al., 1996: 50, both identified asSpecimens from Lahad Datu (Danum valley) were in stagnant
or slow moving clearwater pools along headwater streams.
Syntopic species include Garra borneensis (Cyprinidae),
Homaloptera stephensoni, Gastromyzon danumensis, G.
lepidogaster, G. cf. punctulatus, Protomyzon griswoldi, P.
whiteheadi, Nemacheilus olivaceus (Balitoridae) (MartinSmith & Tan, 1998). Specimens from Tawau were from quiet
murky pools along the coast in oil palm plantations. Numerous
specimens of B. ocellata inhabit these quiet pools and it is
the numerically dominant species.

ab

The figure in Kottelat et al. (1993: pl. 77) shows an adultmale with full body iridescence. Through selective breeding,a blue form has been developed (Linke, 1991: 74). Breeding
report shows that, as in B. ocellata, the female does not help
in the transfer of eggs to the male for oral-brooding, the malepicks up the fertilised eggs and oral-brood without any

Fig. 24. a) Betta ocellata - BMNH 1959.7.7:1, holotype, 70.0 mm
SL, b) ZRC 37642, 56.6 mm SL.

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
holotype. — ZRC 40961, 6 ex., 21.8-51.8 mm SL; same locality
data as holotype, 23 May.1996.

participation of the female (van den Nieuwenhuizen, 1995;
Tan, 1997). This behaviour, along with the open mouth
display, are uniquely found in the B. unimaculata group.

Others - UMS uncat., 14 ex., 21.6-41.1 mm SL; SBP uncat., 14 ex.,23.0-38.3 mm SL; same locality data as holotype. — SBM uncat.,
6 ex., 24.7-52.2 mm SL; UMS uncat., 6 ex., 24.3-40.7 mm SL; SBP
uncat., 6 ex., 23.0-41.6 mm SL; same locality data as holotype, 23
May.1996. — ZRC 40962, 6 ex., 12.5-50.8 mm SL, SBM uncat.,
6 ex., 14.1-24.5 mm SL, UMS uncat., 6 ex., 17.0-25.1 mm SL, SBP
uncat., 6 ex., 17.7-22.9 mm SL; Borneo: Sabah, Maliau Basin, small
stream near camp 1988 (ca. 5°14’N 116°53’E), ca. 7 km northeastof basecamp 1996 (ca. 5°14’N 116°53’E), coll. S. H. Tan, T. H. T.
Tan et al., 15-17 May.1996. — SBM uncat., 3 ex., 15.7-23.4 mm
SL, UMS uncat., 2 ex., 15.8-16.1 mm SL, SBP uncat., 2 ex., 14.314.9 mm SL; Borneo: Sabah, Maliau Basin, small streams at righttrail ca. 3 to 4 km into Jalan Babi towards Maliau falls, coll. S. H.
Tan, T. H. T. Tan et al., May.1996.

Comparative notes. – Betta ocellata is distinguished fromthe other members of the species group in the following
characters: greater body depth at dorsal-fin origin than B.
macrostoma and B. gladiator (21.6-24.5% SL, vs. 18.4-21.1);
greater length of anal-fin base than B. macrostoma and B.
gladiator (52.2-54.5% SL, vs. 43.2-52.8); greater height at
pectoral-fin origin than B. macrostoma and B. gladiator (16.718.7% SL, vs. 10.6-16.5). Betta ocellata is further
distinguished from B. unimaculata in the following
characters: more pointed lateral head profile and more conelike dorsal head profile (vs. blunter and more bulbous headprofile, Fig. 23b); caudal-fin transverse bars faint or absent(vs. distinct); posterior extremity of maxilla at vertical throughfront margin of orbit (vs. behind front margin, Figs. 23c-d);more subdorsal scales (51/2, vs. 5); more lateral scales (33,
vs. 32); more anal-fin rays (30-31, mode 30, vs. 27-30, mode
28 or 29); greater length of dorsal-fin base (10.5-11.3% SL,
vs. 7.9-10.4); greater head width (21.0-24.0% SL, vs. 19.621.8).

Diagnosis. – Betta gladiator is distinguished from the othermembers of the B. unimaculata group in: no anal-fin basesheath scales; no dorsal-fin spine; body yellowish brown,
opercle blueish; caudal fin plain brown ; vertebrae 33-34;
anal-fin rays 28-31 (mode 30); dorsal-fin rays 7-9 (mode 8);
pectoral-fin rays 16-17 (mode 17); subdorsal scales 5-61/2(mode 6); lateral scales 34-36 (mode 35); predorsal scales
24-27 (mode 26); predorsal length 68.0-72.8% SL; headlength 28.5-31.3% SL; length of anal-fin base 48.1-52.8%
SL; length of dorsal-fin base 7.3-11.4% SL; lower jaw length28.3-38.3% HL.

Remarks. – Betta ocellata was described by de Beaufort(1933) on the basis of a single specimen 99 mm TL from
Bettotan, near Sandakan. He described the fish as having “...
a black blotch ... and bordered anteriorly by a white area,
forming an indistinct ocellus, below the median line at baseof caudal ...” (de Beaufort, 1933: 36). He also noted that thisspecimen has no anal spines.Fresh specimens that we
examined have one or two anal-fin spines and the white areaaround the black spot at caudal peduncle is indistinct. Inger(1955) commented that B. unimaculata was the only species
found in the headwaters of Sungai Pinang, which drains into
Sungai Kretam Kechil. But all these specimens actually are
B. ocellata, whose type locality is not very far northwest of
the area actually sampled by Inger.

Description. – General body shape and appearance as in Figs.
25a-b, 47f. Body slender, head broad and blunt; caudal fin
rounded with distal part of rays protruding, dorsal and anal
fins bluntly pointed, pelvic fin filamentous. Vertebrae 2-3 +8 + 22-23 = 33-34 (mode 33 or 34, n = 10). Meristic andmorphometric data in Table 5. Maximum known size 55.5mm SL (ZRC 40960).
Coloration. – For live coloration, see Fig. 47f. When alive,
dorsal half of body dark brown, ventral half lighter brown orslight yellowish. Male with dark blue iridescent opercle, witha little yellowish-green iridescence on body; when preserved,
opercle appearing black. Female without dark pigmented

Inger & Chin (1962: 159) synonymised B. ocellata and B.
unimaculata on the argument that the placement of the eye
above the maxilla, the segmentation of the first anal-fin ray
and the colour at the base of the caudal fin fell within thenormal variation and coloration of populations examined bythem. They had not examined the type material of B.
unimaculata and a comparison of the type material of both
species shows that there are clear differences between thetwo species (see above).

a

Betta gladiator, new species(Figs. 25a-b, 47f)b

Material examined. – Holotype - SBM uncat., 45.5 mm SL; Borneo:
Sabah, Maliau Basin, northeast of basecamp 1996 (ca. 5°14’N
116°53’E), stream at right trail ca. 2 km into Jalan Babi towards
Maliau falls, coll. S. H. Tan, T. H. T. Tan et al., 22 May.1996.
Paratypes - ZRC 40960, 9 ex., 13.3-55.5 mm SL; RMNH 33088,
2 ex., 42.0-48.5 mm SL; CMK 13125, 2 ex., 37.7-49.2 mm SL;
SBM uncat., 13 ex., 26.0-42.9 mm SL; same locality data as

Fig. 25. a) Betta gladiator – ZRC 40960, paratype, 56.0 mm SL,b) ZRC 40960, paratype, 39.2 mm SL, without dorsal fin.

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Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei
opercle when alive or preserved. Pelvic fin filamentous, tip
white, rest iridescent whitish. Anal fin with blueish subdistal
band, edge iridescent green. Dorsal, caudal and anal-fin
transverse bars present. Caudal peduncle black blotch present
in both sexes and juveniles Preserved specimens uniformly
dark brown dorsally and lighter brownish yellow ventrally.

BETTA PICTA GROUP
Betta taeniata Regan, 1910
(Figs. 26a-b, 47g-h)
Betta taeniata Regan, 1910: 781, pl. 78, fig. 1; Weber & de Beaufort,
1922: 362 (part); Herre, 1940: 45 (part); Vierke, 1984: 58, 1988:
335; Witte & Schmidt, 1992: 3245 (key only); Kottelat et al.,
1993: 163, pl. 77; Kottelat & Lim, 1995: 248.
Betta spec. affin. edithae & affin. foerschi: Brown & Brown, 1987:
168, pl. 14.

Distribution. – Betta gladiator is currently known only from
the Maliau basin, a near circular plateau enclosed by highelevations in the Sabah inland, part of the Kinabantangan
basin (Fig. 23) (S. H. Tan & Ng, 1997, and Martin-Smith etal., 1998; as B. unimaculata).

Material examined. – Lectotype - Sarawak: - BMNH 1893.3.6:147,
42.1 mm SL male; Borneo: Sarawak, Sungai Senah, coll. A. Everett.

Field notes. – Specimens were collected from heath forest,in acidic water of pH 4.2-4.8. The only syntopic fishes are
Puntius sealei (Cyprinidae) and Hemibagrus aff. baramensis(Bagridae). Betta gladiator is an aggressive and territorial
species. Single adults were found about one metre apart,usually together with a small number of juveniles. Specimens
kept in captivity fought continuously and had to be kept
separately, otherwise they would have killed each other (pers.
observ.).

Paralectotypes - BMNH 1893.3.6:148-150, 3 ex., 33.8-37.2 mm SL,
same locality data as lectotype.
Others - ZRC 39266, 4 ex., 31.4-42.2 mm SL, CMK 13020, 2 ex.,36.4-38.1 mm SL, SM uncat., 2 ex., 26.6-34.4 mm SL; Borneo:
Sarawak, Sungai Kuhas (1°09’10.0"N 110°29’22.7"E), 5.8 km into
right side-road at 6.9 km into Tebelu Tebakang turnoff after Serian
town, coll. P. K. L. Ng et al., 5 Sep.1995. — ZRC 39581, 6 ex.,15.2-39.6 mm SL; CMK 13017, 2 ex., 25.5-35.2 mm SL; SM uncat.,2 ex., 25.4-31.6 mm SL; Borneo: Sarawak, tributary of Sungai Kuhas(1°9’23.1"N 110°29’29.9"E), 0.5 km into Kampung Lanchang, 5.8
km into right track of side-road at 6.9 km into Tebelu Tebakang
turnoff after Serian town, coll. P. K. L. Ng et al., 14 Jan.1996. —ZRC 40963, 7 ex., 26.9-40.5 mm SL; Borneo: Sarawak, tributaryof Sungai Kuhas (1°9’23.1"N 110°29’29.9"E), 0.5km into Kampung
Lanchang, 5.8 km into right side-road at 6.9 km into Tebelu
Tebakang turnoff after Serian town, coll. P. K. L. Ng et al., 31

Etymology. – From the Latin gladiator for warrior or fighter,alluding to the aggressive behaviour of the species. Used asa noun in apposition.
Comparative notes. – Betta gladiator is distinguished fromthe other members of the species group in the following
characters: more pectoral fin rays (16-17, vs. 11-14); morelateral scales (34-36, vs. 31-341/2); flat lateral head profile
and a bulbous dorsal head profile (head width 64.5-77.3%
HL, vs. 56.4-74.2); more vertebrae than B. macrostoma andB. unimaculata (33-34, vs. 30-32); more subdorsal scales thanB. ocellata and B. unimaculata (5-61/2, vs. 4-51/2); fewer
subdorsal scales than B. macrostoma (5-61/2, vs. 6-8); dorsalfin origin above 19-21st lateral scale (vs. 15-19th in B. patoti
and B. macrostoma); more predorsal scales than B. ocellata
and B. macrostoma (24-27, vs. 21-24); more lateral scales
than B. unimaculata and B. macrostoma (34-36, vs. 31-33);
greater total length than B. ocellata (135.4-139.2% SL, vs.
131.9-135.6); greater predorsal length than B. macrostoma(68.0-72.8% SL, vs. 62.3-66.7); smaller head length than B.
ocellata (28.5-31.3% SL, vs. 31.4-33.3); smaller body depth
at dorsal-fin origin than B. ocellata (18.4-20.4% SL, vs. 20.524.5); greater pelvic-fin length (29.0-35.7% SL, vs. 21.6-27.6)
and smaller length of anal-fin base than B. macrostoma (48.152.8% SL, vs. 43.2-49.2); smaller length of anal-fin base thanB. ocellata (48.1-52.8% SL, vs. 52.2-54.5); smaller length of
dorsal-fin base than B. macrostoma (7.3-11.4% SL, vs. 12.215.7); smaller height at pectoral-fin origin than B. ocellata(10.6-16.5% SL, vs. 16.7-18.7).

abRemarks. – Betta gladiator seems to be most closely allied
to B. ocellata, a common species in the lowlands of Sabah.
The isolated geography of the Maliau Basin has probably
facilitated the evolution of this species. Specimens without
dorsal fins (three out of 150 specimens) could be due toinbreeding or other unknown factors (Fig. 25b).

Fig. 26. a) Betta taeniata - BMNH 1893.3.6:147, male lectotype(uppermost), 42.1 mm SL, BMNH 1893.3.6:148-150, 3
paralectotypes (2 depicted), 33.8-37.2 mm SL, b) ZRC 39581, 36.6mm SL.

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Aug.1996. — ZRC 40964, 2 ex., 21.7-27.6 mm SL; Borneo:
Sarawak, Sungai Kuhas (1°09’10.0"N 110°29’22.7"E), 5.8 km into
right side-road at 6.9 km into Tebelu Tebakang turnoff after Serian
town, coll. P. K. L. Ng et al., 31 Aug.1996. — ZRC 40988, 3 ex.,14.8-38.2 mm SL; Borneo: Sarawak, Sungai Semabang, 21.7 km
towards Simunjan from Kuching-Sri Aman road (1°12’46.4"N
110°55’40.4"E), coll. H. H. Tan & S. H. Tan, 4 Sep.1996.
Kalimantan Barat: CMK 11758, 7 ex., 13.9-54.9 mm SL; ZRC
39271, 5 ex., 23.3-45.2 mm SL; ZRC 39272, 2 males, 38.8-41.2mm SL; Borneo, Indonesia: Kalimantan Barat, Kapuas basin, Sungai
Palat at Pala Hulu (Kec. Siberuang, Kampung Renyai Hulu), km
101 on road from Sintang to Putussibau, coll. M. Kottelat, 16
Jun.1995. — CMK 10546, 4 ex., 12.2-52.2 mm SL; Borneo,
Indonesia: Kalimantan Barat, Kapuas basin, Sungai Palat at Pala
Hulu (Kec. Siberuang, Kampung Renyai Hulu), km 99 on road from
Sintang to Putussibau, coll. M. Kottelat, 16 Sep.1993.

Description. – General body shape and appearance as in Figs.
26a-b, 47g-h. Meristic and morphometric data of type andfresh material in Table 6. Body stout and short, head relatively
pointed. Unpaired fins rounded, anal fin maybe bluntly
pointed, pelvic fin with first simple ray filamentous but short,extending to sixth anal ray (including spines). Vertebrae (typematerial) 2 + 8 + 18 = 28 (mode 28, n = 3), (fresh material)
2 + 8 + 17-19 = 27-29 (mode 28, n = 8). Maximum known
size 54.9 mm SL (CMK 11758).
Coloration. – For live coloration, see Figs. 47g-h (see also
Linke, 1991: 68; Kubota et al., 1996: 36). Mature male withiridescent blue to green opercle scales when alive, blackish
when preserved. Iridescent scales on body faint, concentrated
on dorsal half of body when alive, not visible when preserved.
In mature male, unpaired fins brownish-red, distal margin of
anal and caudal fins dark blue with iridescent light blue borderanteriorly and a thin iridescent blue edge. Dorsal fin with
distal iridescent green edge. Iridescent blue edges on analand caudal fins most obvious when male in antagonistic
behaviour or freshly preserved. Pelvic fin with distal half offilamentous ray white, rest brownish-red. Preserved maleretaining distinct dark fin edges, faint in female both aliveand preserved. Female with more distinct black markings or
spots on unpaired fins when alive, faint or absent whenpreserved. Juvenile with central and second central black
stripes on body. A black caudal peduncle spot also present.
Preserved specimens dark brown dorsally and light brownventrally, fins brownish to hyaline, with distinct dark edges.

Diagnosis. – Betta taeniata is characterised by a relatively
weakly contrasted sexual dimorphism. Mature males havemore intense coloration, but when disturbed they are similar
to the females. In both sexes, the opercle has blue to green
iridescent scales depending on angle of reflected light. The
anal and caudal fins (and sometimes the dorsal fin) arebordered distally by a dark blue edge, followed by aniridescent blue or green border when live (Fig. 4); only a
blackish distal border is visible when preserved. Betta taeniata
is distinguished from the other members of the B. picta species
group by the following combination of characters: anal-fin
rays 23-26 (mode 26); subdorsal scales 5-6 (mode 6); lateralscales 271/2-30 (mode 29); caudal peduncle depth 14.8-19.7%
SL; head length 30.3-34.5% SL; body depth at dorsal-fin
origin 25.3-30.1% SL; length of anal-fin base 47.9-52.7%
SL; interorbital width 29.2-36.2% HL.

Distribution. – Betta taeniata is apparently restricted to hill

Fig. 27. Distribution map of B. taeniata (circle).

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Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei
Singapore and Peninsular Malaysia is erroneous. Examinationof specimens collected and identified by Herre and depositedin ZRC reveals that they are all B. pugnax . The specimen
from Sarawak listed by Brown & Brown (1987: 168, pl. 14)
as “Betta spec. affin. edithae & affin. foerschi” is almost
certainly B. taeniata.

streams of southern Sarawak and northern part of Kalimantan
Barat (Fig. 27).
Field notes. – Most of the fresh samples of B. taeniata werecollected from a fast flowing clear water hill stream near
Serian, south Sarawak (pH 7.0). The substratum was quartz
gravel littered with rocks of variable size. The width of thestream was about 10 m at its widest and water depth wasfrom 5 cm to 80 cm. No aquatic vegetation was present. Betta
taeniata was collected from the more stagnant stretches ofthe stream, among the submerged leaf litter, tree roots andoverhanging bank vegetation. Syntopic species include:
Barbodes collingwoodi, Paracrossochilus vittatus, Puntius
kuchingensis, P. banksi, P. orphoides, P. sealei, Rasbora
sarawakensis, R. caudimaculata (Cyprinidae), Gastromyzon
cf. fasciata, G. cf. punctulatus, Homaloptera tweediei,
Nemacheilus saravacensis (Balitoridae), Glyptothorax cf.
major (Sisoridae), Clarias teijsmanni (Clariidae),
Hemirhamphodon kuekenthali (Hemiramphidae), Channa
lucius (Channidae) and Macrognathus maculatus(Mastacembelidae). The Iban name for this species is ikant’pachi. This species is an oral brooder (pers. observ.).

Roberts (1989: 173) listed B. taeniata from the Kapuas basin.
His specimens had anal ray counts of II, 26-29, which hecommented as being agreeable to the description of B. edithae.
Examination of Roberts’ material deposited in MZB (MZB
3867, MZB 3870) confirms that it actually is B. edithae.

BETTA SPLENDENS GROUP
Betta imbellis Ladiges, 1975
(Figs. 28, 48a-b)
Betta imbellis Ladiges, 1975: 263; Kottelat, 1989: 19 (list only);
Schaller & Kottelat, 1989: 36; Ng et al., 1992: 28, Fig. 100;
Witte & Schmidt, 1992: 26 (key); Kottelat et al., 1993: 162, pl.
76; Vidthayanon et al., 1997: 54; Vidthayanon, 2002: 105.
Betta splendens Regan, 1910: 782 (part) (see Schaller & Kottelat,
1989, for details).
Betta splendens (non Regan, 1910) - Herre, 1940: 46; Tweedie, 1952:
75; Alfred, 1963: 147; 1966: 50 (part).

Comparative notes. – Betta taeniata is distinguished fromthe other members of thespecies group by the following ofcharacters: greater mode of anal-fin rays than B. picta and B.
simplex (26, vs. 22-24); greater mode of subdorsal scales thanB. picta and B. simplex (6, vs. 5-5 1/ 2); lower mode oftransverse scales than B. simplex (91/2,, vs. 111/2,); greater modeof lateral scales than B. picta and B. simplex (29, vs. 28);
dorsal-fin origin above modal 14th lateral scale (vs. 15th in
B. simplex); relatively smaller predorsal length than B. simplex(62.4-67.7% SL, vs. 66.7-69.8); smaller head length than B.
simplex (30.3-34.5% SL, vs. 34.6-37.2); greater body depth
at dorsal-fin origin than B. picta (25.3-30.1% SL, vs. 21.525.5); smaller body depth at dorsal-fin origin than B. simplex(25.3-30.1% SL, vs. 29.3-32.2); greater length of anal-fin basethan B. picta (47.9-52.7% SL, vs. 42.6-48.4) (data of B.
simplex from Kottelat, 1994).

Material examined. – Malaysia: ZRC 32130-54, 25 ex., 12.5-36.5mm SL; Malaysia: Penang, Bayan Lepas, Kampung S. Tiram, ditch
off airport, coll. H. H. Tan & S. H. Tan, 9 Jun.1993. — ZRC 32235262, 28 ex., 13.0-32.6 mm SL; Malaysia: Penang, Kampung Terang,
coll. H. H. Tan & S. H. Tan, 10 Jun.1993. — ZRC 437, 11 ex.,
Malaysia: Kedah, Sungai Patani; E. D. B. Wolf, 1951. — ZRC 442,
9 ex., Malaysia: Kedah, Alor Star; E. L. B. Wolfe, 1951. — ZRC
38418, 1 ex., Malaysia: Perak, Krian, stream flowing from Bukit
Merah Reservoir, coll. H. H. Tan et al., 21 Dec.1994. — ZRC 39540,
1 ex., Malaysia: Perak, ca. 11 km before Gerik, trubutary of Sungai
Kulim, coll. H. H. Tan et al., 19 Nov.1995. — ZRC 18842-47, 6
ex., 19.0-28.9 mm SL males, Malaysia: Selangor, 32 km RawangKuala Selangor road, coll. P. K. L. Ng, 24 Aug.1991. — ZRC 1446467, 4 ex., Malaysia: Selangor, north Selangor peatswamp forest,
stream near padi field on west edge of swamp forest, coll. P. K. L.
Ng, 15 May.1991. — ZRC 24656-57, 2 ex., 25.3-26.4 mm SL males,
Malaysia: Terengganu, Rantau Abang, swamp at 56 km KuantanKuala Terengganu road; T. H. T. Tan & D. S. L. Chung, 3-5
Aug.1992. — ZRC 17295-99, 5 ex., 22.3-25.1 mm SL; Malaysia:
Terengganu, Rantau Abang, stream at 1 km west off East Coast
Highway about 53 km south of Kuala Terengganu, coll. K. K. P.
Lim, 10-11 Sep.1991. — ZRC 40965, 4 ex., 14.0-23.3 mm SL;
Malaysia: Terengganu, Rantau Abang, 56 km to Kuala Terengganu,
coll. P. K. L. Ng et al., 16 May.1995. — ZRC 14776-832, 57 ex.,12.8-31.3 mm SL; Malaysia: Johor, Ayer Hitam, in ditch at
Kampung Botak Parit, coll. P. K. L. Ng et al., 23 May.1991. —Remarks. – Regan (1910:78) described B. taeniata on thebasis of four syntypes. The condition of the syntypes is ratherpoor, but they fortunately still allow to recognize enough
diagnostic characters. The best preserved specimen (BMNH
1893.3.3:147, 42.1 mm SL) is here designated as thelectotype. It is a male, as shown by the dark anal distal margin.
Weber & de Beaufort’s (1922: 362) described B. taeniata as
having the anal penultimate rays in old mature specimens
strongly produced and reaching the extremity of the caudal
fin and prolonged median rays in the caudal fin and II, 2025 anal-fin rays. These characters do not fit with those of theB. picta group (see earlier). The distribution cited by Weber& de Beaufort (1922) includes Sumatra, where B. falx andB. pugnax do occur (pers. observ.) and it seems that Weber& de Beaufort’s material might be a mixture of several
species.
Herre (1940: 45) stated having obtained B. taeniata in
Singapore, Johore, Pahang, Perak (Malay Peninsula) andKuching (Sarawak). His identification of specimens from

Fig. 28. Betta imbellis – ZRC 32235, 28.1 mm SL male.

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
ZRC 22148-172, 25 ex., 16.9-32.2 mm SL; Malaysia: Johor, Pontian,Parit Sikom, coll. K. K. P. Lim & D. S. L. Chung, 14 May.1992.
— ZRC 14839-144, 6 ex., 18.9-28.8 mm SL; Malaysia: Johor, Ayer
Hitam, coll. P. K. L. Ng & K. K. P. Lim, 23 May.1991. Singapore:
ZRC 37595, 3 ex., 20.5-32.6 mm SL; Singapore: stream at Lorong
Banir, coll. K. K. P. Lim et al., 24 May.1994. Sumatra: ZRC 40966,
17 ex., 12.1-27.0 mm SL, CMK 13126, 5 ex., 20.3-24.8 mm SL;
Indonesia: Sumatra, North Sumatra, ca. 80 km south of Medan, Sei
Rampah, Pantai Berdagai, pond at Kampung Nagur (3°31’14.0”N
99°10’32.2”E), coll. H. H. Tan et al., 15 Jun.1996. Sabah: ZRC
40967, 12 ex., 8.5-24.4 mm SL; Borneo: Sabah, Tawau, Jalan Sin
On Jaya, swampy area (4°16’07.6”N 117°54’25.6”E), coll. H. H.
Tan et al., 5 Oct.1996 (most probably introduced).

Description. – General body shape and appearance as in Figs.
28, 48a-b. Body slender (body depth at dorsal-fin origin 24.230.1% SL); head small (HL 25.8-30.9% SL); caudal fin
rounded with distal red crescent margin when live, yellowish
when preserved (Fig. 5a); pelvic fin falcate; dorsal fin
rounded, anal fin pointed, anal fin reaching to half or moreof caudal fin. Meristic and morphometric data in Table 7.
Maximum known size 36.5 mm SL (ZRC 32130).
Coloration. – For live coloration, see Figs. 48a-b (see Linke,
1991: 34; Kubota et al., 1996: 16). Male with head darkbrown, opercle with iridescent blue scales, eye with reflective
green top. Body light brown with posterior half of body scales
iridescent blue. Dorsal fin iridescent blue, with transverse bars
distinct; caudal fin with red distal margin in a crescent, rays
iridescent blue; anal fin iridescent blue, posterior ray tips red;
pelvic fin falcate, red with white tip; pectoral fin hyaline.
Female less colourful, lacking iridescence and red caudal fin
crescent. Throat with chin-bar, opercle pale gold. Body with
distinct central and second central stripes, with caudal
peduncle spot. Fins yellowish to hyaline, dorsal fin withtransverse bars.

Diagnosis. – Betta imbellis is distinguished from othermembers of the B. splendens group by the following
combination of characters: a bright red crescent on distal edgeof caudal fin (vs. faint crescent or absence in B. splendens
and B. smaragdina); no caudal transverse bars (vs. presencein B. smaragdina and B. stiktos); dorsal transverse bars on
distal two-thirds of dorsal fin (vs. whole dorsal fin in B.
smaragdina); iridescent blue opercle scales (vs. parallel red
vertical opercle bars in B. splendens and green iridescent
opercle scales in B. smaragdina); dark brown to black body(vs. light brown body in B. smaragdina and B. stiktos); lesslateral scales than B. stiktos (30-32, vs. 33-34); more slenderbody than B. splendens (body depth 24.2-30.1% SL, vs. 27.132.2); more backwards position of dorsal fin than in B.
splendens (dorsal-fin origin above lateral scales 15-17, mode
16, vs. 14-16, mode 14); relatively smaller length of dorsalfin base than B. splendens (12.9-18.9% SL, vs. 16.7-23.8);
smaller height at pectoral-fin origin than B. splendens (14.818.9% SL, vs. 18.5-20.7).

Distribution. – Betta imbellis is widely distributed in
Peninsular Malaysia (Fig. 29). This species is probably native
to the northern part of Sumatra in Medan (pers. observ.;
Schaller & Kottelat, 1989). Its presence in Sabah is mostprobably due to introduction. Betta imbellis is apparently feralin Singapore (Ng et al., 1993).
Field notes. – Betta imbellis is often found in stagnant pools
and ditches, in clear to brown water.

Fig. 29. Distribution map of B. imbellis (circle).

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Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei
10 ex., ZRC 21051-21052, 2 ex.,22.4-28.4 mm SL; Sumatra: Riau
Province, Pulau Padang, Sungai Ponder, 12 km east of Karau, onroad to Meribur, coll. M. Kottelat, 12 Feb.1991. — ZRC 38257, 2
ex., 26.1-27.9 mm SL; Sumatra: Jambi Province, coll. M. Kottelat,
May.1994. — ZRC 38611, 4 ex., 17.0-24.5 mm SL; Sumatra: Jambi
Province, Danau Rasau, M. Kottelat & H. H. Tan, 1-2 Jun.1994.

Remarks. – There is only one indigenous species in the study
area: B. imbellis. This species has had a long and confusedtaxonomic history. Cantor (1850) listed “B. pugnax var.” asa domesticated form of B. pugnax from Siam (Bangkok,
Thailand). This is the species known today as B. splendens.
Regan (1910: 782) described B. splendens on the basis of
two lots of specimens (total nine specimens) from Bangkok
and Pinang (Pulau Penang). Schaller & Kottelat (1989)
examined eight of the syntypes and identified the Bangkok
specimens as B. splendens and the Penang specimens as B.
imbellis. They also designated a lectotype from among the
Bangkok syntypes (BMNH 1898.11.8:95, 38.6 mm SL). Betta
splendens has one junior subjective synonym, B. abbreviata
Pellegrin, 1925. Schaller & Kottelat (1989) commented thatthe author of B. abbreviata was Blanc (1963), but this isincorrect. The term variety was used in the past to namespecimens which were not clearly defined. The variety
described by Pellegrin (1925) was a captive stock of
domesticated origin from Indochina. According to the Code(Article 45.6.4, ICZN, 1999), Blanc’s use of name makes it
valid, as it is used in a way equivalent to a subspecies. The
correct citation of the author of B. abbreviata is thus Pellegrin(1925).

Diagnosis. – Betta coccina is distinguished from the othermembers of the B. coccina group by the following unique
combination of characters: small iridescent green lateralblotch in male, missing in female; dark red body colour; noparallel vertical iridescent gold opercle bars; pelvic fin falcatewith black filamentous tip, rest red; median fins with discrete
iridescent green spots; dorsal-fin rays 9-12; anal-fin rays 2729; subdorsal scales 7-8; lateral scales 31-32; predorsal scales
18-21 (adapted from Vierke, 1979; Witte & Schmidt, 1992;
Ng & Kottelat, 1992).

Ladiges (1975) described B. imbellis from Kuala Lumpur,
Selangor. In literature before 1975, B. imbellis had been
identified as B. splendens , viz. Herre (1940), Tweedie (1952)
and Alfred (1963, 1966). Herre (1940) comments on B.
splendens from Mandai (Singapore) probably refer to B.
imbellis; although there are feral B splendens in Singapore(Lim & Ng, 1990; Ng et al., 1993; pers. observ.). Tweedie(1952) and Alfred (1963, 1966) listed B. splendens from
Kedah, Terengganu, Penang and Singapore. Their specimens
in ZRC are all B. imbellis.

Betta livida Ng & Kottelat, 1992
(Figs. 32, 48e)

Remarks. – See Ng & Kottelat (1992) for a comparative
diagnosis. Vierke (1979) described B. coccina on the basisof aquarium specimens from Jambi, Sumatra. Recent
collections yielded specimens mainly from peat swamps inJambi (pers. observ.). Specimens from Peninsular Malaysia
are from the aquarium trade. These specimens were reportedlycollected from the Segamat-Muar area in Johor (Fig. 31), an
information which seems reliable. For live coloration, see
Figs. 48c-d, Linke (1991: 30) and Kubota et al. (1996: 20).

Betta coccina (non Vierke, 1979) - Witte & Schmidt, 1992: 307.
Betta livida Ng & Kottelat, 1992: 177; Tan & Tan, 1994: 44.
Betta, new species - Ng et al., 1992: 28; Fig. 13.
Material examined. – Holotype - Malaysia: - ZRC 15287, 29.0mm SL (38.8 mm TL) male, blackwater stream, north Selangor peatswamp forest, 800 m from 45 km mark, on road from Tanjung Malim
to Sungai Besar, coll. 1991 Honours Year class, 18 Jun.1991.

BETTA COCCINA GROUP
Betta coccina Vierke, 1979
(Figs. 30, 48c-d)Paratypes - All localities in blackwater streams or pools, northSelangor peat swamp forest, Peninsular Malaysia. — ZRC 1528815289, 1 male, 30.2 mm SL; 1 female, 31.2 mm SL; 800 m from45 km mark, on road from Tanjung Malim to Sungai Besar, coll.
1991 Honours Year class, 18 Jun.1991. — CMK 8032, 25 ex., 23.329.9 mm SL; Peninsular Malaysia: Selangor, north Selangor peatswamp forest, blackwater stream at 47 km milestone on road fromSungei Besar to Tanjung Malim, coll. 1991 Honours Year class, 19
Jun.1991. — ZRC 22764-22767, 2 males, 32.3-35.6 mm SL; 2females, 28.9-36.3 mm SL; on road from Tanjung Malim to Sungai
Besar, coll. P. K. L. Ng, Jun.1991. — ZRC 14439, 1 male, 29.8 mm
SL; northwestern boundary of north Selangor peat swamp forest,
coll. P. K. L. Ng et al., 15 May.1991. — ZRC 18836-18840, 5 ex.,19.6-29.0 mm SL; 43 km milestone, on road from Tanjung Malim
to Sungai Besar, coll. P. K. L. Ng, 14 Sep.1991. — ZRC 1525415273, 20 ex., 14.8-29.5 mm SL; 50 km milestone, on road from
Sungai Besar to Tanjung Malim, coll. 1991 Honours Year class, 18
Jun.1991. — ZRC 15209-15228, 20 ex., 16.9-27.5 mm SL; 200 m
from 45 km milestone, on road from Tanjung Malim to Sungai Besar,coll. 1991 Honours Year class, 17 Jun.1991. — ZRC 22768, 1 male,33.1 mm SL; ZRC 22829, 1 female, 27.3 mm SL; between 32-33
km milestone, in pool surrounded by Pandanus, Rawang-Kuala
Lumpur road, coll. P. K. L. Ng, Nov.1991. — ZRC 18851-18853,

Betta coccina Vierke, 1979a: 288; Schaller, 1985: 348, 1986: 299;
Kottelat, 1989: 19 (list only); Witte & Kottelat, in Kottelat, 1991:
279; Witte & Schmidt, 1992: 313, 326 (key); Ng & Kottelat,
1992: 178; Kottelat et al., 1993: 161, pl. 75; Tan & Tan, 1994:
44.
Material examined. – Malaysia: ZRC 28616-635, 20 ex., 18.8-28.7mm SL; Malaysia: Johor, Segamat-Muar area, purchased from theaquarium trade, coll. P. K. L. Ng, Oct.1992. Sumatra: CMK 7290,

Fig. 30. Betta coccina – ZRC 28616, 27.3 mm SL.

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
3 ex., 13.4-22.9 mm SL; Rawang-Kuala Lumpur road, 32 km
milestone, coll. P. K. L. Ng et al., 24 Aug.1991.

median fins with discrete iridescent green spots; dorsal-fin
rays 9-13; anal-fin rays 25-31; subdorsal scales 7; lateralscales 30-31; predorsal scales 18-21 (from Ng & Kottelat,1992).

Others - All localities in blackwater streams or pools, north Selangor
peat swamp forest, Peninsular Malaysia. — ZRC 14440-14451, 12
ex., northwestern boundary of north Selangor peat swamp forest,
coll. P. K. L. Ng et al., 15 May.1991. — ZRC 15290-15302, 13 ex.,15.6-30.1 mm SL; 800 m from 45 km mark, on road from Tanjung
Malim to Sungai Besar, coll. 1991 Honours Year class, 18 Jun.1991.
— ZRC 22769-22789, 21 ex., 17.7-34.0 mm SL; road from Tanjung
Malim and Sungai Besar, coll. P. K. L. Ng et al., Jun.1991. — ZRC
17962, 1 ex., 29.0 mm SL; 200 m from 37 km milestone, on roadfrom Sungai Besar to Tanjung Malim, coll. P. K. L. Ng et al., 25
Aug.1991. — ZRC 17842, 1 ex., 21.6 mm SL; 39 km milestone, onroad from Sungai Besar to Tanjung Malim, coll. P. K. L. Ng et al.,24 Aug.1991. — ZRC 15274-15286, 13 ex., 34 km milestone, Sungai
Besar to Tanjung Malim road, coll. 1991 Honours Year class, 17
Jun.1991. — ZRC 18037, 1 ex., 650 m from 35 km milestone, onroad from Sungai Besar to Tanjung Malim, coll. P. K. L. Ng, 25
Aug.1991. — ZRC 15204-15208, 5 ex., 15.5-26.4 mm SL; 700 m
from 41 km milestone, on road from Sungai Besar to Tanjung Malim,coll. 1991 Honours Year class, 19 Jun.1991. — ZRC 16696-16699,
4 ex., on road from Sungai Besar to Tanjung Malim, coll. 1991
Honours Year class, 17-19 Jun.1991. — ZRC 15202-15203, 2 ex.,stream adjacent to Sungai Tengi, coll. 1991 Honours Year class, 20
Jun.1991.

Remarks. – See Ng & Kottelat (1992) for more details. The
B. cf. coccina from Rawang, Selangor of Witte & Schimdt(1992) is B. livida. Betta livida is known only from the peatswamp forest in Selangor (Fig. 31). For live coloration, seeFig. 48e and Kubota et al. (1996: 22). This species is a bubble
nest brooder (Smith, 1996).

Betta persephone Schaller, 1986
(Figs. 33, 48f)
Betta persephone Schaller, 1986: 298; Kottelat, 1989: 19 (list only);
Schaller & Kottelat, 1989: 36; Witte & Kottelat, in Kottelat,
1991: 279; Witte & Schmidt, 1992: 308, 325 (key); Ng &Kottelat, 1992: 179; Tan & Tan, 1994: 42.
Material examined. – Malaysia: ZRC 28596-615, 20 ex., 17.5-25.9mm SL, ZRC uncat., 57 ex., 16.8-25.5 mm SL; Malaysia: Johor,
Segamat-Muar area, purchased from aquarium trade, coll. P. K. L.
Ng, Oct.1992. — ZRC 24443-452, 10 ex., 14.7-24.4 mm SL; - ZRC
24691-96, 6 ex., 20.8-25.0 mm SL; Malaysia: Johor, ca. 3 km northof Ayer Hitam, north bank of Sungai Sambrong, in swamp forest,
coll. K. K. P. Lim & D. G. B. Chia, 6 Aug.1992. — ZRC 38255,
2 ex., 19.9-21.4 mm SL; Malaysia: Johor, Ayer Hitam, in swamp
forest, coll. H. H. Tan & D. M. Armitage, Apr.1994.

Diagnosis. – Betta livida is distinguished from the othermembers of the B. coccina group by the following unique
combination of characters: small iridescent green lateralblotch in both male and female, disappearing with age; redbody colour; parallel vertical iridescent gold bars on opercle;
pelvic fin falcate, with wide, iridescent green tip, rest red;Diagnosis. – Betta persephone is distinguished from the otherFig. 31. Distribution map of B. coccina (circle), B. livida (triangle), B. persphone (square), B. tussyae (upside-down triangle), B. brownorum(star), B. miniopinna (diamond), B. burdigala (asterix), B. rutilans (hollow circle).

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Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei
Kampung Mahkota) road, coll. P. K. L. Ng, J. B. Tay & D. Chia,
20 Oct.1991. — ZRC 22795-22797, 3 ex., CMK 8036, 3 ex., pool
along road from Mersing to Pekan, about 80 km before Kuantan,
3°10’31.8"N 103°26’41.8"E, coll. P. K. L. Ng & M. Kottelat, 9Mar.1992. — ZRC 22790-22794, 5 ex., CMK 8040, 6 ex., pool
along road from Mersing to Pekan, 73 km before Kuantan,
3°18’13.0"N 103°25’25.2"E, coll. P. K. L. Ng & M. Kottelat, 9Mar.1992. — ZRC 22798-22802, 5 ex., CMK 8055, 6 ex., stream
across road from Mersing to Pekan, 69 km before Kuantan, coll. P.
K. L. Ng & M. Kottelat, 9-10 Mar.1992. — ZRC 22803-22805, 3
ex., CMK 8073, 2 ex., 16 km milestone, on road from Kuantan to
Mersing, 3°42’47.3" 103°16’28.5"E, coll. P. K. L. Ng & M. Kottelat,
9 Mar.1992. — ZRC 22806-22812, 7 ex., CMK 8108, 7 ex., stream,road from Pekan to Mersing, 1 km north of 3°22’04.1"N103°25’13.8"E, coll. P. K. L. Ng & M. Kottelat, 10 Mar.1992. —ZRC 22813-22828, 16 ex., CMK 8115, 17 ex., road from Pekan to
Mersing, 3°22’04.1"N 103°25’13.8"E, coll. P. K. L. Ng & M.
Kottelat, 10 Mar.1992.

members of the B. coccina group by the following unique
combination of characters: no lateral blotch; blackish bodycolour; no parallel vertical iridescent gold bars on opercle;
pelvic fin falcate with white filamentous tip, rest black to
reddish; median fins covered diffusely with light green
iridescence; dorsal-fin rays 9; anal-fin rays 24-27; subdorsal
scales 6-7; lateral scales 27-29; predorsal scales 17-18(adapted from Schaller, 1986; Witte & Schmidt, 1992; Tan& Tan, 1994).
Remarks. – See Ng & Kottelat (1992) and Tan & Tan (1994)
for comparative diagnosis. Schaller & Kottelat (1989)
completed the listings of the type material. This species isrestricted to the peat swamps in southwestern Johor (Fig. 31).
It is typically found in shallow pools among submerged leaf
litter. Syntopic osphronemids include B. bellica, B. imbellis
and Trichopsis vittata. The closest allied species is B.
miniopinna which occupies a similar habitat in Pulau Bintan,
Riau Archipelago (Tan & Tan, 1994). Betta persephone
exhibits kinship care (Witte & Schmidt, 1992; pers. observ.).
For live coloration, see Fig. 48f, Linke (1991: 40) and Kubota
et al. (1996: 26).

Diagnosis. – Betta tussyae is distinguished from the othermembers of the B. coccina group by the following unique
combination of characters: no lateral blotch; brick-red bodycolour; faint parallel vertical iridescent gold bars on opercle;
pelvic fin falcate, with narrow white tip, rest red; median finswithout iridescent green spots; dorsal-fin rays 11-12; analfin rays 27-28; subdorsal scales 8; lateral scales 30-31(adapted from Schaller, 1985; Witte & Schmidt, 1992; Ng &Kottelat, 1992).

Betta tussyae Schaller, 1985
(Figs. 34, 48g)
Betta tussyae Schaller, 1985: 350; 1986: 300; Kottelat, 1989: 20(list only); Schaller & Kottelat, 1989: 35; Witte & Kottelat, inKottelat, 1991: 279; Witte & Schmidt, 1992: 307, 326 (key);
Ng & Kottelat, 1992: 179; Tan & Tan, 1994: 44.

Remarks. – See Ng & Kottelat (1992) for comparative
diagnosis. Schaller & Kottelat (1989) clarified the status of
the type material. This taxon is restricted to eastern Pahang,where it the coastal peat swamps (Fig. 31). It occurs
syntopically with B. waseri. For live coloration, see Fig. 48g,
Linke (1991: 71) and Kubota et al. (1996: 23). Betta tussyae
is a bubble nest brooder (Smith, 1994).

Material examined. – Holotype - ZRC 38451/CMK 5947, 36.5 mm
SL; Malaysia: Pahang, 17 km south of Pekan, or about 77 km southof Kuantan on road along east coast, ca. 1 km from sea, coll. P.
Nagy, 1984.
Others - Peninsular Malaysia: Pahang. — ZRC 18493-18495, 2
males, 24.1-26.4 mm SL, 4 females, 22.7-27.9 mm SL; in pool,
about 500 m before 10 km milestone, Kuantan to Gambang (nearBetta brownorum Witte & Schmidt, 1992
(Figs. 35, 48h)
Betta brownorum Witte & Schmidt, 1992: 312, 326 (key); Ng &Kottelat, 1992: 178; Kottelat et al., 1993: 161, pl. 75; Tan &Tan, 1994: 44; Kottelat & Lim, 1995: 248.
Betta spec. affin. coccina (non Vierke, 1979) - Brown & Brown,
1987: 169, pl. 11.
Material examined. – ZRC 39582, 60 ex., 8.4-25.9 mm SL, CMK
13018, 10 ex., 10.5-22.3 mm SL, SM uncat., 10 ex., 10.0-23.3 mm
SL; Borneo: Sarawak, ca. 200 m into peat swamp forest on left sideof road (1°12’08.7"N 110°39’52.2"E), ca. 11 km towards Gedong
after turnoff from Serian-Sri Aman road (after 78 km to Kuchingfrom Gedong mark), coll. H. H. Tan & S. H. Tan, 16 Jan.1996. —Fig. 32. Betta livida – ZRC 15287, holotype, 29.0 mm SL.

Fig. 33. Betta persephone – ZRC 24443, 24.0 mm SL.

Fig. 34. Betta tussyae – ZRC 25066, 30.2 mm SL.

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
pH 4.0, coll. P. K. L. Ng et al., 9 Mar.1992. — ZRC 35403-08, 6
ex., 39.6-80.3 mm SL; Malaysia: Pahang, about 100 m south of 68
km stone, on road from Johor Bahru to Kuantan, near Pekan, wellshaded peat swamp forest stream, blackwaters, pH 3.4, coll. P. K.
L. Ng et al., 19 Oct.1992. — CMK 8057, 4 ex., 20.6-44.0 mm SL;
same locality as previous, coll. M. Kottelat et al., 9 Mar.1992. —ZRC 18617-20, 4 ex., 20.5-33.2 mm SL; Malaysia: Pahang, about
200 m north of 16 km stone, on road from Johor Bahru to Kuantan,
coll. P. K. L. Ng et al., 20 Oct.1991. — ZRC 25907, 1 ex., 56.6 mm
SL, CMK 8074, 2 ex., ZRC 25210-11, 2 ex., 21.9, 32.0 mm SL;
Malaysia: Pahang, stream at 16 km stone, on road from Mersing to
Kuantan, coll. P. K. L. Ng et al., 9 Mar.1992. — CMK 3189, 1 ex.,31.1 mm SL; Malaysia: Kuantan, Sungai Soi; P. Nagy, May.1981.
— ZRC 24370, 1 ex., 19.7 mm SL; Malaysia: Terengganu, near
Rantau Abang, 56 km stone, on road from Kuantan to Kuala
Terengganu, coll. P. K. L. Ng et al., 18 Mar.1992.

ZRC 39583, 3 ex., 18.6-19.4 mm SL; Borneo: Sarawak, ca. 50 m
into peatswamp forest and ditch on left side of road (1°10’50.0"N
110°39’22.1"E), ca. 8 km towards Gedong after turnoff from SerianSri Aman road, coll. H. H. Tan & S. H. Tan, 16 Jan.1996. — ZRC
40968, 1 ex., 21.8 mm SL; Borneo: Sarawak, peatswamp forest at
Balai Ringin, ca. 0.5 km after Sungai Kerang towards Sri Aman
(1°03’01.3”N 110°45’38.1”E), coll. H. H. Tan & S. H. Tan, 5
Sep.1996.

Diagnosis. – Betta brownorum is distinguished from the othermembers of the B. coccina group by the following unique
combination of characters: large iridescent green lateral blotch
in both male and female; reddish-magenta body colour; noparallel vertical iridescent bars on opercle; pelvic fin falcate,with white filamentous tip, rest red; median fins without
iridescent green spots; dorsal-fin rays 10-11; anal-fin rays
26-28; subdorsal scales 6-7; lateral scales 291/2-31; predorsal
scales 18-20 (adapted from Witte & Schmidt, 1992; Ng &Kottelat, 1992).

Diagnosis. – Betta waseri is distinguished from the othermembers of the B. waseri group by the unique suite ofcharacters: black tear-drop shape marking on throat (Fig. 3a);opercle usually with gold scales, without iridescent green
scales; lower margin of opercle black; no dorsal and caudal
transverse bars; hyaline anal-fin edge; predorsal scales 231/12-24 (mode 24); subdorsal scales 6-6 /2 (mode 6); postdorsal
scales 101/2-12 (mode 11); lateral scales 311/2-321/2 (mode 32);
dorsal-fin rays 9-10 (mode 9); anal-fin rays 28-31 (mode 30);
head length 28.6-33.3% SL; body depth 23.1-27.6% SL (fromNg & Kottelat, 1994).

Remarks. – See Witte & Schmidt (1992) for more details.
Betta brownorum was collected from the same habitat typeas B. persephone. The head profile is also similar to that ofB. persephone. Betta brownorum is a bubble nest brooder
which exhibits kinship care; the older siblings from theprevious batch take care of the younger ones (Witte &Schmidt, 1992). The lateral green spot of B. brownorum isconsistently larger than that of B. livida. Syntopic species
collected from Gedong area include Rasbora kalochroma,
Boraras maculatus (Cyprinidae), Clarias teijsmanni, C. cf.
nieuhofii and Encheloclarias baculum (Clariidae). Betta
brownorum is found from Sibu to Matang and Kuching (Fig.
31). For live coloration, see Fig. 48h and Linke (1991: 50,
identified as B. spec. affin. coccina).

Remarks. – There has been some confusion about the statusof B. waseri. Schmidt (1988) had synonymised B. waseri with
B. macrophthalma (type locality Singapore), the type of whichis actually a juvenile specimen of B. pugnax. Witte & Schmidt(1992) regarded B. macrophthalma as a junior synonym of
B. pugnax without providing reasons. This synonymy,
however, was confirmed and elaborated on by Tan & Tan(1996). Witte & Schmidt (1992) also placed B. waseri into
the B. anabatoides group. Ng & Kottelat (1994) redescribed
B. waseri from fresh material. See Fig. 37 for the distribution
map. For live coloration, see Fig. 49a, Linke (1991: 37,
identified as B. macrophthalma) and Kubota et al. (1996: 48).

BETTA WASERI GROUP
Betta waseri Krummenacher, 1986
(Figs. 36, 49a)
Betta waseri Krummenacher, 1986: 177; Kottelat et al., 1992: 11(list only); Witte & Schmidt, 1992: 324 (key only); Ng &Kottelat, 1994: 595.

aMaterial examined. – Holotype - ZMZ 129201, male, 90.8 mm SL;
Malaysia: Pahang, Kuantan, 22.5 km from Kuantan, coll. W. Schar,
Aug.1985 (radiographs and photographs examined only).
Others - ZRC 35402, 80.7 mm SL male; Malaysia: Pahang, 16 km
stone, Johor Bahru to Kuantan road, shallow blackwater stream,b

Fig. 36. a) Betta waseri – ZMZ 129.201, holotype, 90.0 mm SL(taken by Maurice Kottelat), b) ZRC 35403, 78.5 mm SL.

Fig. 35. Betta brownorum – ZRC 39582, 22.4 mm SL.

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Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei
Betta hipposideros Ng & Kottelat, 1994
(Figs. 38, 49b)

Estate), coll. K. K. P. Lim, 14 Sep.1991. — ZRC 18029-18034, 6
ex., 29.5-47.3 mm SL; north Selangor peat swamp forest, 500 m
from 36 km stone, on road to Tanjung Malim, coll. P. K. L. Ng etal., 25 Aug.1991. — ZRC 27761, 1 ex., 64.0 mm SL; north Selangor
peat swamp forest, 650 m from 35 km stone, on road from Sungai
Besar to Tanjung Malim, coll. P. K. L. Ng et al., 18 Sep.1992. —ZRC 28550, 1 ex., 25.2 mm SL; north Selangor peat swamp forest,
200 m from 45 km stone, road to Sungai Besar, coll. P. K. L. Ng
et al., 18 Sep.1992. — CMK 10037, 16 ex., 37.8-77.8 mm SL; northSelangor peat swamp forest, stream at 43 km stone on road from
Tanjung Malim to Sungai Besar, coll. P. K. L. Ng et al., 19 Jun.1991.
— CMK 10038, 10 ex., 27.9-59.6 mm SL; north Selangor peatswamp forest, stream at 0.2 km from 45 km stone on road from
Tanjung Malim to Sungai Besar, coll. P. K. L. Ng et al., 17 Jun.1991.
— ZRC 28515-28517, 69.1-77.9 mm SL; peat swamp forest, northshore of Sungai Bernam; P.K.L. Ng et al., 19 Sep.1992. Sumatra:
ZRC 39630, 2 male ex., 62.1-67.1 mm SL; Indonesia, Sumatra: Riau,
peat swamp draining into Sungai Bengkwan, tributary of Indragiri
River; H. H. Ng, S. H. Tan et al., Jun.1995.

Betta aff. waseri (non Krummenacher, 1986) - Ng et al., 1992: 28,
Fig. 14, 15, 98.
Betta hipposideros Ng & Kottelat, 1994: 597.
Material examined. – Holotype - Malaysia: - ZRC 18688, 69.0mm SL male; Malaysia: Selangor, north Selangor peat swamp forest,
39 km stone, on road from Sungai Besar to Tanjung Malim, coll.
P. K. L. Ng et al., 24 Aug.1991.
Paratypes - Malaysia: Selangor - ZRC 18689-18705, 17 ex., 33.471.1 mm SL; north Selangor peat swamp forest, 39 km stone, onroad from Sungai Besar to Tanjung Malim, coll. P. K. L. Ng et al.,24 Aug.1991. — ZRC 15351, 1 ex., 32.0 mm SL; north Selangor
peat swamp forest, stream at 34 km stone, on road to Tanjung Malim;
Zoology Honours Class, 17 Jun.1991. — ZRC 15352, 1 ex., 25.9mm SL; north Selangor peat swamp forest, stream at 50 km stone,on road to Tanjung Malim (United Plantations Estate); Zoology
Honours Class, 18 Jun.1991. — ZRC 15353, 1 ex., 43.1 mm SL;
north Selangor peat swamp forest, stream at 43 km stone, on roadto Sungai Besar; Zoology Honours Class, 18 Jun.1991. — ZRC
15354, 15355, 2 ex., 27.5, 37.0 mm SL; north Selangor peat swamp
forest, stream 700 m from 41 km stone, on road to Tanjung Malim;
Zoology Honours Class, 19 Jun.1991. — ZRC 15356, 15357, 2 ex.,35.3, 38.7 mm SL; north Selangor peat swamp forest, 47 km stone,on road from Sungai Besar to Tanjung Malim; Zoology Honours
Class, 19 Jun.1991. — ZRC 15358-15361, 4 ex., 25.8-39.6 mm SL;
north Selangor peat swamp forest, 800 m from 45 km stone, onroad to Sungai Besar; Zoology Honours Class, 18 Jun.1991. — ZRC
17097, 1 ex., 54.6 mm SL; north Selangor peat swamp forest, stream
at 50 km stone, on road to Tanjung Malim (United Plantations

Diagnosis. – Betta hipposideros is distinguished from theother members of the B. waseri group by the unique suite ofcharacters: black horse-shoe shaped marking on throat (Fig.
3b); opercle usually with gold scales, without iridescent green
scales; lower margin of opercle brown; dorsal and caudal-fin
transverse bars; hyaline anal fin edge; predorsal scales 21-24(mode 23); subdorsal scales 61/2-7 (mode 61/2); postdorsal
scales 9-10 (mode 91/2); lateral scales 31-32 (mode 32); dorsalfin rays 9-10 (mode 10); anal-fin rays 28-31 (mode 30); headlength 28.3-34.1% SL; body depth 23.8-33.1% SL (from Ng& Kottelat, 1994).

Fig. 37. Distribution map of B. waseri (circle), B. hipposideros (triangle), B. tomi (square), B. pi (upside-down triangle), B. spilotogena(diamond), B. renata (asterix).

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
Remarks. – See Ng & Kottelat (1994) for details. The
specimens from Sumatra were only observed in blackwater
habitats (Fig. 37). They can be found in the slow-flowing tostagnant shallow parts of the peat swamp forest amongst the
thick leaf litter and overhanging vegetation. Syntopic
osphronemids present are: Belontia hasseltii, Betta coccina,
B. aff. fusca, B. simorum, Parosphromenus sumatranus,
Sphaerichthys osphromenoides and Trichogaster leerii. This
new record and extends the range of B. hipposideros to
Sumatra. For live coloration, see Fig. 49b and Kubota et al.
(1996: 47).

members of the B. waseri group by the unique suite ofcharacters: black semi horse-shoe shaped marking on throat(Fig. 3c); iridescent blueish opercle scales; lower margin of
opercle black; no dorsal and caudal-fin transverse bars; broadand distinct dark green anal-fin margin; predorsal scales 2123 (mode 23); subdorsal scales 5-51/2 (mode 5); postdorsal
scales 10-101/2 (mode 10); lateral scales 31-32 (mode 31);
dorsal-fin rays 9-10 (mode 9); anal-fin rays 28-31 (mode 30);
head length 29.3-37.1% SL; body depth 24.1-29.8% SL (fromNg & Kottelat, 1994).
Remarks. – See Ng & Kottelat (1994) for details. See Fig.
37 for the distribution map. For live coloration, see Fig. 49c
and Kubota et al. (1996: 47).

Betta tomi Ng & Kottelat, 1994
(Figs. 39, 49c)
Betta tomi Ng & Kottelat, 1994: 603.

Betta pi Tan, 1998
(Figs. 40, 49d)

Material examined. – Holotype - Malaysia: - ZRC 35409, 70.4mm SL male; Malaysia: Johor, tributary of Sungai Mupor, about 15
km on road from Kota Tinggi to Mersing, about 1°52’N 103°56’E,
freshwater swamp forest, coll. P. K. L. Ng, Aug.1992.

Betta pi Tan, 1998: 285, figs. 3-4; Vidthayanon, 2002: 106.
Material examined. – Holotype - ZRC 40289, 54.9 mm SL male;
Thailand: Narathiwat Province, Mae Nam Tod Deng, about 6 km
north of Sungai Kolok, open area in swamps ca. 100-200 m south
before bridge, coll. M. Kottelat et al., 1-3 Nov.1995.

Paratypes - ZRC 35410, 1 ex., 37.6 mm SL; same data as holotype.
— CMK 9753, 6 ex., 16.8-60.4 mm SL; same locality as holotype,coll. M. Kottelat, P. K. L. Ng & K. K. P. Lim, 22 Jan.1991. — ZRC
35411-35416, 6 ex., 24.8-64.9 mm SL; Malaysia: Johor, tributaries
of Sungai Mupor, freshwater swamp forest, coll. P. K. L. Ng et al.,13 Jan.1993.

Paratypes - ZRC 40290, 48.5 mm SL male, 43.2 mm SL female,
CMK 12030, 3 ex., 20.0 - 41.1 mm SL; same locality data as
holotype. — ZRC 40291, 6 ex., 35.9-60.2 mm SL, CMK 13025, 3
ex., 35.9-53.0 mm SL; Thailand: Narathiwat Province, peatswamps
in Mae Nam Tod Deng, near Sungai Kolok, coll. K. Kubota et al.,Mar.1996. — ZRC 40292, 18 ex., 19.6-43.4 mm SL; Thailand:
Narathiwat Province, peatswamps in Mae Nam Tod Deng, near
Sungai Kolok, coll. K. Kubota et al., Dec.1995.

Others - ZRC 18059-18073, 15 ex., 11.8-20.9 mm SL; Malaysia:
Johor, north of Mersing, 177 km milestone, on road from Johor
Bahru to Kuantan, coll. P. K. L. Ng et al., 19 Oct.1991. — ZRC
19079-19131, 53 ex.; Malaysia: Johor, Mawai district; M. W. F.
Tweedie, 1935. Singapore: ZRC 1097, 6 ex.; Singapore: Mandai
road; M. W. F. Tweedie, 1937.

Diagnosis. – Betta pi is distinguished from the other membersof the B. waseri group by the unique suite of characters: blackpi (p) shaped marking on throat (Fig. 3g); opercle usuallywith gold scales, without green scales; lower margin of
opercle black, black spots on opercle; no dorsal and caudalfin transverse bars; broad and distinct dark blue anal margin;
predorsal scales 24-25; subdorsal scales 51/2-6 (mode 6);
postdorsal scales 10-12 (mode 11); lateral scales 32-33 (mode33); dorsal-fin rays 8-10 (mode 9); anal-fin rays 30-32 (mode31); head length 32.3-34.4% SL; body depth 26.4-29.7% SL(from Tan, 1998).

Diagnosis. – Betta tomi is distinguished from the otherFig. 38. Betta hipposideros – ZRC 18689, male holotype, 69.0 mm
SL.

Remarks. – The type locality of this species is in the drainage
of Sungai Kolok in Thailand. Sungai Kolok makes the border
between Thailand and malaysia and the species is expected
to also occur in the peatswamp forests of Kelantan (Fig. 37).

Fig. 39. Betta tomi – ZRC 35409, male holotype, 70.4 mm SL.

Fig. 40. Betta pi – ZRC 40289, male holotype, 54.9 mm SL.

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Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei
121.694, 38 ex., 14.9-31.2 mm SL; Indonesia: Java, Tjinjiroean,
after Poentjak Gedeh, ca. 1600m (Gunung Gede: ca. 25 km southeastof Bogor); Kerkhoven, 1921. – MHNG 2090-92, 3 ex., 23.6-30.0mm SL; Indonesia: Java, Sukabumi (ca. 40 km southeast of Bogor);
Walsh, July 1930. – ZMA 121.693, 2 ex., 28.8 mm SL; Indonesia:
Java, Bandoeng (Bandung); Huysmans. – ZMA 121.689, 1 ex., 24.5mm SL; Indonesia: Java, Bandoeng (Bandung), ca. 700m; I.
Jacobson, 1934. – RMNH 26940, 4 ex., 31.8-35.3 mm SL; Indonesia:
Java, Bezoeki (Besoki) (ca. 7°16’S 109°E, ca. 1050 km southeastof Bogor); J. Semmelink, 1864-65. – RMNH 13698-13699, 2 ex.,32.6-35.2 mm SL; Indonesia: Java, Ambarawa (ca. 7°8’S 110°20’E);
J. Sybrandi, 1933. – RMNH 10740, 4 ex., 24.9-34.2 mm SL;
Indonesian Archipelago; from P. Bleeker’s collections. Betta falx
- Sumatra: MZB 9308, holotype, 32.7 mm SL, male; Indonesia:
Sumatra, Jambi, Sungai Alai, km 19.5 on Muara Bungo - Muara
Tebo road (bridge at Sungai Alai: 1°28’42.6”S 102°18’31.7”E); H.
H. Ng & S. H. Tan, 22 Jun.1995. — ZRC 40974, paratypes, 20 ex.,17.2-33.0 mm SL, ZMA 121.673, paratypes, 6 ex., 21.8-31.0 mm
SL, MZB 9307, paratypes, 4 ex., 17.5-26.3 mm SL, RMNH 33087,
paratypes, 5 ex., 19.7-26.7 mm SL; same data as holotype. — ZRC
38571, paratypes, 44 ex., 13.8-32.4 mm SL, CMK 11119, paratypes,
44 ex.; Indonesia: Sumatra, Jambi, Sungai Alai at km 28 on Muara
Bungo - Muara Tebo road, between half hour downriver of bridge
to ca. 1 hour upriver, including small tributaries and Danau Gresik,
coll. M. Kottelat & H. H. Tan, 30-31 May.1994. — ZRC 38254,
paratypes, 3 ex., 31.7-36.1 mm SL; Indonesia: Sumatra, Jambi,
Sungai Alai, coll. M. Kottelat & H. H. Tan, May.1994. — ZRC
42496, paratypes, 10 ex., 17.0-34.9 mm SL, MHNG 2593.95,
paratypes, 5 ex., 29.7-30.1 mm SL; Indonesia: Sumatra, Jambi,
Sungai Alai, km 19.5 km on Muara Bungo - Muara Tebo road (bridge
at Sungai Alai: 1°28’42.6”S 102°18’31.7”E), coll. H. H. Tan & H.
H. Ng, 22 Jul.1997.

See Tan (1998) for more details. For live coloration, see Fig.
49d.

COMPARATIVE MATERIAL EXAMINED
Betta splendens group. Betta splendens - Thailand: ZRC Group
no. 57, Cat. no. 443, 40 ex., 27.9-39.8 mm SL males; Thailand:
Bangkok; Milton, 12 Oct.1962. — ZRC 39580, 1 ex., 42.4 mm SL
male; Thailand: Bangkok, purchased from Chatukchak market, coll.
P. K. L. Ng, 21 Sep.1995. — Holotype - Betta splendens abbreviata- MNHN 1925-82, 1 ex., 36.3 mm SL; Indochine; D. Coyon. Betta
smaragdina - Thailand - ZRC 37597, 2 male ex., 32.4-34.5 mm
SL, 1 female, 27.1 mm SL; Thailand: Nakhon Ratchasima; K.
Kubota, 1993. — ZRC 39301, 7 ex.; Thailand: Nakhon Ratchasima;
K. Kubota, Dec.1995. — ZRC, 11 ex.; Thailand: Nakhon
Ratchasima; K. Kubota et al., 1996.
Betta edithae group. Betta edithae - Kalimantan Selatan: Holotype- SMF 18712, 55.3 mm SL, male; Borneo, Kalimantan Selatan:
Banjarmasin, Barito delta; J. Vierke, Aug.1983. Paratype - SMF
18713, 56.2 mm SL, female; same locality data as holotype. Others- ZRC 40969, 11 ex., 20.5-31.6 mm SL; Borneo, Kalimantan Selatan:
Banjarmasin, stream at 55 km to Martapura on Rantau-Martapura
road; H. H. Ng & O. K. S. Chia, 6 Jun.1996. Kalimantan Barat:
MZB 3867, 10 ex., 21.7-37.4 mm SL; Borneo, Kalimantan Barat:
24 km NE of Pontianak, small forest stream flowing into Sungai
Sepatah, a tributary of Sungai Mandor (0°07.5’N 109°30’E), coll.
T. R. Roberts, 10 Jul.1976. — MZB 3870, 1 ex., 34.1 mm SL;
Borneo, Kalimantan Barat: Sungai Sekilap, tributary to Sungai
Mempawah, ca. 64 km N of Pontianak and 38 km by road NE of
Andjongan (0°33.5’N 109°22.5’E), coll. T. R. Roberts, 14 Jul.1976.
Banka: ZRC 31031-31044, 14 ex., 10.6-34.9 mm SL; Indonesia,
Sumatra: Banka, ca. 25 km north of Koba, stream between Desa
Kurau and Desa Balilik, outlet of an abandoned mine, coll. M.
Kottelat et al., 3 Mar.1993. — ZRC 30865-30899, 17 ex., 10.728.0 mm SL; Indonesia, Sumatra: Banka, Kwsn. Sungai Liat,
Kampung Jelit, 3 km from main road (Sungai Liat) to Tanjung
Persema beach, coll. M. Kottelat et al., 2 Mar.1993. — ZRC 3515935161, 3 ex., 19.4-32.1 mm SL; Indonesia, Sumatra: Banka, 4 km
north of Bikang village on road from Koba to Toboali, coll. M.
Kottelat et al., 3 Mar.1993. Bintan: ZRC 37551, 4 ex., 24.3-42.5mm SL; Indonesia, Sumatra: Bintan, stream at km 47 Tanjung UbanTanjung Pinang road, coll. H. H. Tan et al., 27 Apr.1994. — ZRC
40970, 6 ex., 18.6-39.6 mm SL; Indonesia, Sumatra: Bintan, northernpart of island, coll. P. K. L. Ng et al., May.1993. — ZRC 34721742, 22 ex., 10.4-35.7 mm SL; Indonesia, Sumatra: Bintan, northernpart of island; N. Sivasothi et al., 11 May.1993. — ZRC 34877883, 7 ex., 17.3-33.6 mm SL; Indonesia, Sumatra: Bintan, ditch fromreservoir near Gunung Bintan Besar; T. H. T. Tan et al., 14
May.1993.

Betta foerschi group. Betta strohi: - Kalimantan Tengah:
Paratypes - ZRC 32126-32129, 4 ex., 33.0-42.4 mm SL; Borneo:
Kalimantan Tengah, Nataik Sedawak (2°41’S 111°13’E), about 30
km south of Sukamara; H. Stroh, Oct.1987. Betta rubra: Sumatra:
MSNG 13019a, lectotype, 28.6 mm SL, MSNG 13019, 4
paralectotypes, 26.9-31.4 mm SL; Sumatra Barat: Siboga; E.
Modigliani, 1886. — BMNH 1893.5.29:1, 1 paralectotype, 33.3 mm
SL; Indonesia: Sumatra, Lake Toba; Modigliani. — MZB 4784, 3ex., 23.4-35.0 mm SL, ZRC 42497, 2 ex., 33.4-35.4 mm SL;
Sumatra: Aceh, Aceh Barat, Alur Sungai Iamueselatan; H. B. Munaf& M. Toha, 13 Dec.1982.
Betta anabatoides group. Betta anabatoides - Kalimantan Selatan:
ZRC 40971, 2 ex., 47.2-55.3 mm SL; Borneo: Kalimantan Selatan,
Banjarmasin, stream at 55 km to Martapura on Rantau-Martapura
road; H. H. Ng & O. K. S. Chia, 6 Jun.1996. Kalimantan Tengah:
ZRC 22842-43, 2 ex., 46.7-55.9 mm SL, CAS 96958, 4 ex., 35.759.4 mm SL; Borneo: Kalimantan Tengah, Mentaya basin, Sungai
Ramban, 22 km west of Sampit on road to Pembuanghulu, coll. T.
R. Roberts, 11 Jun.1992. — MZB 5326, 1 ex., 60.2 mm SL; Borneo:
Kalimantan Tengah, Kotim, Kec. Pulan Malan, Sungai Banjang;
A. Saim, 13 Mar.1984. Kalimantan Barat: MZB 3846, 1 ex., 58.0mm SL; Borneo, Kalimantan Barat: 24 km NE of Pontianak, small
forest stream flowing into Sungai Sepatah, a tributary of Sungai
Mandor (0°07.5’N 109°30’E), coll. T. R. Roberts, 10 Jul.1976.

Betta picta group. Betta picta - Java: ZMA 102.149, 10 ex., 22.331.6 mm SL, ZRC 40973, 2 ex., 29.9-32.0 mm SL; Indonesia: Java,
ponds near Trogon (ca. 6°21’S 106°34’E, ca. 30 km northwest ofBogor); M. Weber, 1888. – ZMA 121.691, 52 ex., 13.3-32.8 mm
SL; Indonesia: Java, Buitenzorg (Bogor); M. Weber, 1888. – MZB1325, 37 ex., 13.8-32.2 mm SL, ZRC 42495, 5 ex., 28.0-32.1 mm
SL; Indonesia: Java, Bogor, Tjidjeruk, Sawahbera; Sukardi, 24
Aug.1970. – RMNH 10447, 3 ex., 29.3-32.4 mm SL; Indonesia:
Java, Buitenzorg (Bogor); M. Weber. – RMNH 15794, 1 ex., 36.4mm SL; Indonesia: Java, Buitenzorg (Bogor), Tjiblagoeng; A.
Bushkiel, 1935. – ZMA 121.675, 30.9 mm SL, male; ZMA 121.589,
13 ex., 19.5-34.3 mm SL, ZRC 40972, 2 ex., 29.9-31.6 mm SL;
Indonesia: Java, Tjipanas (Cipanas: ca. 6°43’S 107°2’E, ca. 30 km
southeast of Bogor) near Sindanglaja; M. Weber, 1888. – ZMA

Betta albimarginata group. Betta albimarginata: Kalimantan
Timur: Holotype - ZRC 38485, 27.9 mm SL male; Borneo,
Indonesia: Kalimantan Timur, Sungai Sebuku basin, Sungai Sanul,a tributary of Sungai Tikung, at about km 7 on logging road from
Semunad (4°04.08’N 117°00.4’E) to Apas (3°59.96’N
116°59.32’E), coll. M. Kottelat & P. McKee, 14 Feb.1993. Paratypes- ZRC 35121-22, 2 ex., 21.8-23.8 mm SL, same locality data as
holotype. Betta channoides: Kalimantan Timur: Paratype - ZRC

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35165, 1 ex., 17.2 mm SL; Borneo, Indonesia: Kalimantan Timur,
Mahakam River basin, unnamed blackwater stream entering
Mahakam River on left side near Mujub (0°01’S 115°43’E), coll.
M. Kottelat, 3 Aug.1991.

LITERATURE CITED
Alfred, E. R., 1958. The food and habitat of the fighting-fish Betta
pugnax. Malayan Nature Journal, 12: 161-163.
Alfred, E. R., 1961a. Singapore fresh-water fishes. Malayan Nature
Journal, 15: 1-19, pl. 1-4.

Betta dimidiata group. Betta dimidiata: Kalimantan Barat: MZB
3849, holotype, 30.6 mm SL; Borneo: Kalimantan Barat, Kapuas
basin, Sungai Seriang Hutan Kecil; T. R. Roberts, 12 Aug.1976. —ZRC 35498-99, 2 ex., 28.8-37.2 mm SL; Borneo: Kalimantan Barat,
Kapuas basin, coll. M. Kottelat, Sep.1993. — ZRC 37707, 3 ex.,33.6-40.0 mm SL; Borneo: Kalimantan Barat, Kapuas basin, coll.
M. Kottelat, Sep.1993. — ZRC 38782, 10 ex., 16.7-32.5 mm SL;
Borneo: Kalimantan Barat, Kapuas basin, Sungai Letang near
Kampung Kandung Suli, coll. M. Kottelat, 8 Jun.1995.

Alfred, E. R., 1961b. The Cantor collection of Malayan fresh-water
fishes. Bulletin of the Raffles Museum, Singapore, 26: 183-186.
Alfred, E. R., 1963. Notes on a collection of freshwater fishes from
Penang. Bulletin of the Singapore National Museum, 32: 143153, 2 pl.
Alfred, E. R., 1966. The fresh-water fishes of Singapore. Zoologische
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de Beaufort, L. F., 1933. On some new or rare species of Ostariophysi
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ACKNOWLEDGEMENTS

Blanc, M., 1963. Catalogue des types d’Anabantidae etd’Ophiocephalidae (Poissons téléostéens perciformes) encollection au Muséum national d’Histoire naturelle. Bulletin duMuséum National d’Histoire naturelle Paris, 35: 70-77.

We are grateful to the following who have helped us in many
ways small and large: Maurice Kottelat, for his
encouragements, loan of material (CMK) and patiently
reviewing the manuscript several times; Ralf Britz, for his
helpful discussions and reviewing the manuscript; Kelvin K.
P. Lim, for his advice and use of his maps; Darrell Siebert(BMNH), David Catania and Susan Middleton (CAS), Barry
Chernoff and Mary Ann Rogers (FMNH), Sonia Muller and
Claude Weber (MHNG), Guy Duhamel and Patrice Pruvost(MNHN), Giulianno Doria (MSNG), Agus Tjakrawidjaja, Ike
Rachmatika, Renny Hardiaty and Daisy Wowor (MZB),
Lipke B. Holthuis and Martin van Oijen (RMNH), Lynne
Parenti and Susan Jewett (USNM), Hans Nijssen and Isaäc
Isbrücker (ZMA), and C. M. Yang (ZRC), for the loan ofmaterial; Tyson R. Roberts, for the discussions andsuggestions; Marina Wong and staff (BRM), for help with
the Brunei trip; Charles M. U. Leh (SM), for help with theSarawak collections, especially for arranging for permits for
Bako; Leo Chai, previously head of Sarawak Forestry
Department, for permission to conduct surveys in Bako; P.
K. Chin, Alvin Wong of Sabah Fisheries, Anna Wong (SBM),
for help with the Sabah trip; Keith M. Martin-Smith and John
Salleh bin Otong (DVFC), for help with the Danum trip;
Katsuma Kubota, for providing material from Thailand;
Gunawan ‘Thomas’ Kasim, for providing material from
Sumatra; Koji Yamazaki and Günther Ettrich, for kindly
providing colour slides of various Betta species; David M.
Armitage, for donation of phototanks; S. H. Tan for his manykind and unselfish radiographing sessions; Research
Associate of RMBR – Maurice Kottelat, Systematics andEcology Laboratory staff, postgraduates, satellites andassociated lab denizens (Oliver K. S. Chia, Daphne S. L.
Chung, W. K. Goh, Y. Y. Goh, Robert Kerle, O. Y. Kok,
Adrian H. B. Loo, H. H. Ng, N. Sivasothi, Diana G. B. Tan,
Cheryl G. S. Tan, Tommy H. T. Tan, Alvin T. C. Wong,
Darren C. J. Yeo) for their help in the field and criticisms.
Last but not least, THH would like to dedicate this work to
his late father, thank his parents and sister who have tolerated
and nurtured his interests throughout the years. This fieldwork conducted had been partly funded by Research Grants
RP 950326, RP 960314 and RP3982327 to P. K. L. Ng by
the National University of Singapore.

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Ng, P. K. L. & M. Kottelat, 1992. Betta livida, a new fighting fish
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Schaller, D. & M. Kottelat, 1989. Betta strohi sp. n., ein neuer
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Ng, P. K. L. & M. Kottelat, 1994. Revision of the Betta waseri
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Schmidt, J., 1988. Wasers grosser maulbrütender Kampffisch Betta
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Ng, P. K. L., J. B. Tay, K. K. P. Lim & C. M. Yang, 1992. The
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Sim, C. H., 2002. A field guide to the fish of Tasek Bera Ramsar
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Ng, P. K. L., J. B. Tay & K. K. P. Lim, 1994. Diversity andconservation of blackwater fishes in Peninsular Malaysia,
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Smith, A., 1994. Spawning and brood care in Betta tussyae.
Anabantoid Association of Great Britain. Labyrinth, 78: 1-4.
Smith, A., 1996. Care and breeding of the jealous Betta. Anabantoid
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van den Nieuwenhuizen, A., 1993. Betta bellica. Die Aquarien- undTerrarien-Zeitschrift, 46: 365-369.

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Tan, H. H., 1997. Notes on the breeding of Betta pulchra Tan &Tan, 1996; B. ocellata de Beaufort, 1933; and B. spilotogena
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Perugia, A., 1893. Di alcuni pesci raccolti in Sumatra dal Dott. ElioModigliani. Annali del Museo Civico di Storia Naturale
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Tan, H. H., 1998. Two new species of the Betta waseri group(Teleostei: Osphronemidae) from central Sumatra and southern
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Popta, C. M. L., 1905. Suite des descriptions préliminaires desnouvelles espèces de poissons recueillies au Bornéo central par
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Leyden Museum, 25: 171-186.

Tan, H. H. & M. Kottelat, 1998a. Two new species of Betta(Teleostei: Osphronemidae) from the Kapuas basin, Kalimantan
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41-51.

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Tan, H. H. & M. Kottelat, 1998b. Redescription of Betta picta(Teleostei: Osphronemidae) and description of B. falx n. sp. from
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Rehwinkel, J., 1995. Care and breeding of Betta balunga Herre 1940

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APPENDIXBETTA AKARENSIS GROUP

Tweedie, M. W. F., 1936. A list of the fishes in the collection of
the Raffles Museum. Bulletin of the Raffles Museum, Singapore,
12: 16-28.

Betta obscura, new species(Fig. 41)

Tweedie, M. W. F., 1952a. Notes on Malayan fresh-water fishes.
3. The anabantoid fishes. Bulletin of the Raffles Museum,
Singapore, 21: 63-76.

Material examined. – Holotype - MZB 9331, 60.2 mm SL; Borneo,
Kalimantan Tengah: Sungai Barito basin, Desa Kerendan, Sungai
Lahei and tributaries, coll. D. J. Siebert, A. Tjakrawidjaja & A.
Mun’im, 23-28 Sep.1992 (DS-3-1995).

Tweedie, M. W. F., 1952b. Notes on Malayan fresh-water fishes.
Bulletin of the Raffles Museum, Singapore, 24: 63-75.
Tweedie, M. W. F., 1953. Malayan aquarium fishes. Malayan Nature
Journal, 8: 47-51, pls. 9-10.

Paratypes - BMNH 1996.3.22:253-260, 9 ex., 28.9-69.1 mm SL, ZRC 41578, 5 ex., 33.0-62.6 mm SL, - MZB 9332, 1 ex., 65.5 mm
SL; same data as holotype. — BMNH 1996.3.22:43-44, 2 ex., 55.763.1 mm SL; Borneo, Kalimantan Tengah: Sungai Barito basin,
Sungai Bunes, a small tributary of Sungai Kerendan (a tributary ofSungai Lahei) above (ca. 1 km) Desa Kerendan, coll. D. J. Siebert,
A. Tjakrawidjaja & A. Mun’im, 24 Sep.1995 (DS-2-1995). —BMNH 1991.9.18:26-28, 3 ex., 26.1-51.1 mm SL; Borneo,
Kalimantan Tengah: Sungai Barito basin, Sungai Murung near
Project Barito basecamp, coll. D. J. Siebert, A. Tjakrawidjaja & O.
Crimmen, 11-13 Feb.1991 (DS-17-1991). — BMNH 1991.9.18:3237, 6 ex., 20.8-54.5 mm SL; Borneo, Kalimantan Tengah: Sungai
Barito basin, Sungai Barito at Muara Laung and market at Puruk
Cahu, coll. D. J. Siebert, A. Tjakrawidjaja & O. Crimmen, 20-21
Feb.1991 (DS-20-1991). — MZB 6941, 3 ex., 23.3-57.6 mm SL;
Borneo, Kalimantan Tengah: Sungai Mongkumuh and Sungai

Vidthayanon, C., 2002. Peat swamp fishes of Thailand. Office ofEnvironmental Policy and Planning, Bangkok, Thailand, 136pp.
Vidthayanon, C., J. Karnasuta & J. Nabhitabhata, 1997. Diversity
of Freshwater Fishes in Thailand. Office of Environmental
Policy and Planning, Bangkok, Thailand, 102 p.
Vierke, J., 1979a. Betta coccina nov. spec., ein neuer Kampffischvon Sumatra. Das Aquarium, 13: 288-289.
Vierke, J., 1979b. Betta anabatoides und Betta foerschi spec. nov.,
zwei Kampffische aus Borneo. Das Aquarium, 13: 386-388.
Vierke, J., 1984. Betta taeniata Regan, 1910 und Betta edithae spec.
nov., zwei Kampffische von Süd-Borneo. Das Aquarium, 18:
58-63.

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Bahadeng, tributary of Sungai Barito, 1.5 km from Muara Laung,coll. D. J. Siebert, A. Tjakrawidjaja & O. Crimmen, Feb.1991.

Etymology. – From the Latin adjective obscurus meaning
indistinct and inconspicuous, referring to the fish in general.

Others - BMNH 1996.3.22: 260-261, 2 ex., 26.5-38.4 mm SL; samedata as holotype. — BMNH 1996.3.22:45-47, 3 ex., 17.4-32.3 mm
SL; Borneo, Kalimantan Tengah: Sungai Barito basin, Sungai Bunes,a small tributary of Sungai Kerendan (a tributary of Sungai Lahei)
above (ca. 1 km) Desa Kerendan, coll. D. J. Siebert, A. Tjakrawidjaja& A. Mun’im, 24 Sep.1995 (DS-2-1995). — BMNH 1996.3.22:3940, 2 ex., 20.7-28.8 mm SL; Borneo, Kalimantan Tengah: Sungai
Barito basin, Sungai Barito at Muara Laung (0°35’34”S
114°44’12”E), coll. D. J. Siebert, A. Tjakrawidjaja & O. Crimmen,
8 Jul.1992 (DS-7-1992). — MZB 6793, 1 ex., 30.7 mm SL; Borneo,
Kalimantan Tengah: Barito Hulu, Sungai Sapen, tributary of Sungai
Joloi, upstream of Sungai Busang, coll. D. J. Siebert, A.
Tjakrawidjaja & O. Crimmen, 7 Feb.1991.

Comparative notes. – Betta obscura is distinguished fromthe other members of the species group by the following
characters: opercle pattern less dense (Fig. 7f) than in mostother members, except B. ibanorum and B. aurigans; fewer
vertebrae than in B. akarensis and B. pinguis (28-30, vs. 3132); greater mode of subdorsal scales (61/2, vs. 5-6); fewer
lateral scales than B. akarensis, B. aurigans and B. pinguis(28-30, vs. 31-331/2); fewer lateral scales modally than B.
akarensis, B. chini, B. pinguis and B. ibanorum (29, vs. 3032); dorsal-fin origin above 14-15th lateral scales (vs. 1619th of B. akarensis, B. chini and B. pinguis); fewer predorsal
scales than B. akarensis, B. chini and B. pinguis (19-21, vs.
22-25); fewer predorsal scales modally (20, vs. 21-24); greater
preanal length than B. chini and B. pinguis (46.1-50.7% SL,
vs. 42.6-46.3); greater body depth than B. ibanorum (28.732.7% SL, vs. 23.3-28.1); relatively greater body depth than
B. chini (28.7-32.7% SL, vs. 26.3-29.0); smaller orbital
diameter than B. balunga and B. chini (21.1-25.3% HL, vs.
26.2-33.3).

Diagnosis. – Betta obscura is distinguished from the othermembers of the B. akarensis group by the following
combination of characters: opercle with second postorbital
stripe; anal-fin rays 26-29 (mode 28); subdorsal scales 51/27 (mode 61/2); lateral scales 28-30 (mode 29); predorsal scales
19-21 (mode 20); postdorsal scales 10-11 (mode 11); preanallength 46.1-50.7% SL; head length 30.9-35.4% SL; lengthof anal-fin base 51.3-56.4% SL.

BETTA UNIMACULATA GROUP

Description. – General appearance is illustrated in Fig. 41.
Meristic and morphometric data in Table 4. Relatively shortand stout body (body depth 28.7-32.7% SL), relatively longand pointed head (HL 30.9-35.4% SL). Unpaired fins pointed,
dorsal and anal fins pointed with posterior rays elongated,caudal fin lanceolate with median rays elongated. Pelvic fin
falcate with long filamentous first ray (33.1-52.6% SL).
Maximum known size 69.1 mm SL (BMNH 1996.3.22:253260).

Betta unimaculata (Popta, 1905)(Figs. 42a-b, 49e)
Parophiocephalus unimaculatus Popta, 1905: 184; 1906: 10, pl. 1,
Figs. 1a, 1b.
Betta unimaculata - Regan, 1910: 779; Weber & de Beaufort, 1922:
355; Herre, 1940 (key only); Witte & Schmidt, 1992: 327;
Kottelat et al., 1993: 163, pl. 77.
Material examined. – Lectotype - RMNH 7660-A, 50.9 mm SL
male, Borneo, Kalimantan Timur: Kajan (Kayan basin), coll. M. le
Dr. A. W. Nieuwenhuis, Sep-Oct.1900.

Coloration. – Live coloration was not recorded, only
preserved specimens examined: Body uniformly brown withdarker dorsum. Preorbital stripe complete from snout througheye to operculum edge, lower preorbital stripe present.
Opercle markings relatively dense, chin-bar incomplete below
first preorbital stripe, a few (1-2) black marks below
postorbital stripe (Fig. 7f). Thin black distal margin on analfin. Dorsal and caudal-fin interradial membranes brownish
red, with vertical hyaline bar pattern in male, faint on anal
fin; only on dorsal fin membrane in female and juvenile.
Pelvic fin filament darkly pigmented, rest brownish-red.

Paralectotypes - RMNH 7659, 11 ex., 27.8-51.6 mm SL; Borneo,
Kalimantan Timur: Howong (upper Mahakam), coll. M. le Dr. A.
W. Nieuwenhuis, 1898. — RMNH 7660, 15 ex., 28.9-53.5 mm SL,
same locality data as lectotype.
Others - MNHN 58-180, 1 ex., 66.8 mm SL; Borneo, Kalimantan
Timur: Kajan; Pfeffer. — ZRC 46324, 50 ex., 15.0-55.2 mm SL;
Kalimantan Timur: Kayan basin; Usat Aran, small rocky hill stream
feeding ladang next to Sungai Pingai (02º00.67’N 115º08.97’E, 500
m asl), coll. H. H. Tan et al., 20 Nov.1999 [THH9991]. — ZRC
46325, 110 ex., 18.6-52.5 mm SL; CMK, 10 ex., 33.7-51.0 mm SL;
MZB 9347, 10 ex., 23.5-42.4 mm SL; Kalimantan Timur: Kayan
basin; Sungai Nap, feeder stream to Sungai Seba Ai (Sungai SebaAi, tributary to Kayan river - 01º59.86’N 115º06.77’E, 550 m asl),coll. H. H. Tan et al., 22 Nov.1999 [THH9995]. — ZRC 46326, 2
ex., 21.5-24.7 mm SL; Kalimantan Timur: Kayan basin; Sungai
Bako, tributary to Kayan river (01º59.02’N 115º06.91’E, 500 m asl),coll. H. H. Tan et al., 23 Nov.1999 [THH9996]. — ZRC 46327, 32
ex., 18.9-54.5 mm SL; Kalimantan Timur: Kayan basin; Sungai
Busang Matu, tributary to Kayan river ca. 500 m upstream of Data
Dian (01º59.89’N 115º07.81’E, 500 m asl), coll. H. H. Tan et al.,24 Nov.1999 [THH9997]. — ZRC 46328, 3 ex., 15.0-47.7 mm SL;
Kalimantan Timur: Kayan basin, Iwan; Sungai Panan, tributary to
Iwan river, ca. 60 mins. upstream from Data Dian (02º00.74’N
115º03.56’E, 530 m asl), coll. H. H. Tan et al., 26 Nov.1999

Distribution. – Betta obscura is presently known only fromthe upper Barito basin in Kalimantan Tengah (Fig. 18).

Fig. 41. Betta obscura – ZRC 41578, paratype, 62.6 mm SL.

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Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei[THH9999]. — ZRC 46329, 10 ex., 22.2-47.6 mm SL; Kalimantan
Timur: Kayan basin; Sungai Belanyan Tekan, tributary to Kayanriver (01º59.67’N 115º07.59’E, 500 m asl), coll. H. H. Tan et al.,26 Nov.1999 [THH99103]. — ZRC 46330, 35 ex., 18.9-54.2 mm
SL; Kalimantan Timur: Kayan basin; Sungai Nah, tributary to Kayanriver, ca. 20 mins upstream of fork to Iwan river (01º57.72’N
115º06.59’E, 550 m asl), coll. H. H. Tan et al., 27 Nov.1999[THH99104].

distal margin iridescent blue or white. Dorsal, caudal and anal
fins sometimes flushed with faint blue iridescence. Pelvic fin
filamentous, with first ray iridescent white, rest hyaline.
Pectoral fin base black, rest hyaline. Female with very little
or no iridescent opercle scales. All transverse bars on finsdistinct and iridescence absent in preserved material.
Distribution. – Betta unimaculata is known only from theupper Kayan (=Kajan) and Howong (Mahakam) drainages
in east Borneo (Popta, 1905:185, 1906: 10-11) (Fig. 23).

Diagnosis. – Betta unimaculata is distinguished from theother members of the B. unimaculata group group by thefollowing combination of characters: distinct black spot nearcaudal peduncle; body brownish; uniform caudal fin colour;
vertebrae 30-32 (mode 31); anal-fin rays 27-30 (mode 2829); dorsal-fin rays 6-8 (mode 8); pectoral fin rays 12-13(mode 12); subdorsal scales 4-5 (mode 5); lateral scales 3133 (mode 32); predorsal scales 24-26 (mode 24 ); predorsal
length 67.7-73.0% SL; head length 29.0-33.3% SL; lengthof anal-fin base 45.9-53.7% SL; length of dorsal-fin base 7.910.4% SL; lower jaw length 31.6-37.9% HL.

Comparative notes. – Betta unimaculata is distinguished fromthe other members of the species group in the following
characters: fewer anal-fin rays than B. ocellata, B. patoti andB. gladiator (27-30, vs. 30-33); fewer dorsal-fin rays than B.
macrostoma (6-8, vs. 9-11); fewer pectoral-fin rays than B.
gladiator (12-13, vs. 16-17); fewer subdorsal scales than B.
macrostoma and B. patoti (4-5, vs. 6-8); fewer lateral scales
than B. gladiator (31-33, vs. 34-36); fewer vertebrae than B.
ocellata, B. patoti and B. gladiator (30-32, vs. 33-34); smaller
predorsal length than B. macrostoma (67.7-73.0% SL, vs.
62.3-66.7); smaller length of anal-fin base than B. ocellata(45.9-53.7% SL, vs. 52.2-54.5); smaller length of dorsal-finbase than B. macrostoma and B. ocellata (7.9-10.4% SL, vs.
10.5-15.7); smaller head width than B. ocellata (19.6-21.8%
SL, vs. 21.0-24.0); smaller lower jaw length than B.
macrostoma (31.6-37.9% HL, vs. 38.0-54.0). Betta
unimaculata is further distinguished from B. ocellata in thefollowing characters: blunter lateral head profile and morebulbous dorsal head profile (vs. pointed and more conicalhead profile, Fig. 24c); distinct caudal-fin transverse bars (vs.
faint or absence); maxilla ends beyond anterior margin oforbit (vs. in front of anterior margin of orbit); fewer subdorsal
scales (5, vs. 51/2); fewer lateral scales (32 vs. 33); fewer analfin rays (27-30, mode 28 & 29, vs. 30-31, mode 30); smaller
length of dorsal-fin base (7.9-10.4% SL, vs. 10.5-11.3).

Description. – General body shape and appearance as in Figs.
42a-b, 49e. Body slender, broad and blunt head; caudal fin
rounded with median rays slightly elongated, tips of rays freeof interradial membrane, dorsal and anal fins relatively
pointed, pelvic fin filamentous. Vertebrae 2 + 8-9 + 20-21(total 30-32, mode 31) (n = 13). Meristic and morphometricdata in Table 5. Maximum known size 66.8 mm SL (MNHN
58-180).
Coloration. – For live coloration, see Fig. 49e. Mature malewith iridescent blue opercle scales when alive, blackish whenpreserved, with a single iridescent scale on dark upper regionof head, which is speckled with black dots. Area below eyewith 2 rows of iridescent blue scales, except region behind
eye and directly below that. Eye with a thin iridescent light
blue lower iris. Chin bar present. Body brown when alivewith irregular faint black bars on lower half, dark brown whenpreserved. Dorsal fin with iridescent white margin, with 56 transverse bars. Caudal fin with up to 8 transverse bars.
Large black rounded spot at lower half of caudal base, inboth sexes and juvenile (Fig. 1). Anal fin with black dots,

Remarks. – Popta (1905) described B. unimaculata on thebasis of 52 syntypes from two locations (Howong and Kajan).
We have examined most of this material. A specimen fromthe Kajan series, RMNH 7660-A (50.9 mm SL, male), is
designated as lectotype.

a

Betta patoti Weber & de Beaufort, 1922
(Figs. 43a-b, 49f-g)
Betta patoti Weber & de Beaufort, 1922: 359; Herre, 1940: 41 (keyonly); Witte & Schmidt, 1992: 323 (key only); Kottelat et al.,
1993: 162, pl. 76.
Material examined. – Lectotype - ZMA 112.513, 63.0 mm SL,
lectotype; Borneo, Kalimantan Timur: Balikpapan, rivulet 25 km
east of Bay of Balikpapan, coll. W. J. Tisot van Patot, 1911.

bParlectotypes - ZMA 121.679, 2 ex., 58.1-75.6 mm SL,
paralectotypes; same data as lectotype. — ZMA 112.510, 1 ex., 59.7mm SL, paralectotype; Borneo, Kalimantan Timur: Balikpapan,
freshwater; W. J. Tisot van Patot, 1911. — ZMA 112.511, 1 ex.,57.6 mm SL, paralectotype; Borneo, Kalimantan Timur: Balikpapan,
Sungai Manggar; W. J. Tisot van Patot, 1911. ZMA 112.512, 2 ex.,42.4-45.5 mm SL, paralectotypes; Borneo, Kalimantan Timur:
Balikpapan, Sungai Manggar; W. J. Tisot van Patot, 1911.

Fig. 42. a) Betta unimaculata – RMNH 7660A, lectotype, 50.9 mm
SL, b) ZRC 46327, 48.7 mm SL.

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
Others - ZMA 121.588, 2 ex., 55.9-71.3 mm SL; Borneo:
Balikpapan; L. Rutten, 1911. — RMNH 10440, 2 ex., 46.0-48.8mm SL; southeast Borneo, before 1921. — MZB 1769, 1 ex., 48.3mm SL; Borneo, Kalimantan Timur: Samarinda, Sungai Mahakam,coll. D. Darnaedi, Sep.1974. — ZRC 46322, 60 ex., 9.5-48.0 mm
SL; CMK, 10 ex., 15.8-41.9 mm SL; MZB 9346, 10 ex., 17.4-37.6mm SL; Kalimantan Timur: Mahakam basin; Taman Wisata Air
Terjun - Tanah Merah, Desa Lempabe, ca. 30 km north of Samarinda(00º26.00’S 117º13.99’E), coll. H. H. Tan & D. Wowor, 11
Nov.1999 [THH9976]. — ZRC 46323, 6 ex., 14.8-60.5 mm SL;
Kalimantan Timur: Mahakam basin; downstream of Taman Wisata
Air Terjun - Tanah Merah, Desa Lempabe, ca. 30 km north ofSamarinda (00º26.00’S 117º13.99’E), coll. H. H. Tan & D. Wowor,
2 Dec.1999 [THH99105].

eye slightly reddish. Region between preorbital stripe andchin bar light iridescent green. Upper and lower lips black.
Opercle faintly iridescent green. Dorsal fin with 5-7 fainttransverse bars. Caudal fin with 7-8 faint transverse bars on
dorsal half, median rays elongated, projecting beyond
interradial membrane. Anal fin darker on anterior region.
Pelvic fin filamentous with first ray white, rest hyaline.
Pectoral fin base dark brown, rest hyaline. Preserved
specimens with light brown body, dorsum darker, 13-14 blackbars on body. Transverse bars on fins present but faint.
Distribution.-Betta patoti is known only from the the southernpart of Kalimantan Timur, around Balikpapan and Samarinda(Fig. 23).

Diagnosis. – Betta patoti is distinguished from the othermembers of the B. unimaculata group by the following
combination of characters: 10-14 vertical dark bars on lower
half of body; body yellowish brown; lanceolated caudal finin adults; uniform light brown caudal fin; vertebrae 33-34(mode 33); anal-fin rays 30-33 (mode 31); dorsal-fin rays 89 (mode 8); pectoral-fin rays 12-14 (mode 13); subdorsal
scales 6-61/2 (mode 6); lateral scales 33-341/2 (mode 34);
predorsal scales 24-26 (mode 25); predorsal length 67.569.9% SL; head length 29.7-31.4% SL; length of anal-finbase 51.0-56.3% SL; length of dorsal-fin base 9.4-11.4% SL;
lower jaw length 32.9-38.5% HL.

Comparative notes. – Betta patoti is distinguished from theother members of the species group by the following
characters: caudal fin lanceolate in adults, which is shared tosome extent with B. pallifina (vs. rounded); 10-14 darkvertical bars on lower half of body (vs. absence); indistinct
pattern on opercle (vs. distinct dark and light opercle patternof B. macrostoma and B. pallifina); more vertebrae than B.
unimaculata, B. macrostoma (33-34, vs. 30-32); more analfin rays than B. macrostoma (30-33, vs. 25-28); fewer dorsalfin rays than B. macrostoma (8-9, vs. 9-11); fewer pectoralfin rays than B. gladiator (12-14, vs. 16-17); more subdorsal
scales than B. unimaculata and B. ocellata (6-61/2, vs. 4-51/2); dorsal-fin origin above 18-19th lateral scales (vs. 15-17thin B. macrostoma); greater predorsal length than B.
macrostoma (67.5-69.9% SL, vs. 62.3-66.7); smaller headlength than B. ocellata (29.7-31.4% SL, vs. 31.4-33.3); greater
length of anal-fin base than B. macrostoma (51.0-56.3% SL,
vs. 43.2-49.2); smaller length of dorsal-fin base than B.
macrostoma (9.4-11.4% SL, vs. 12.2-15.7); smaller bodyheight at pectoral fin than B. ocellata (12.7-16.6% SL, vs.
16.7-18.7); smaller head width than B. ocellata (18.0-20.7%
SL, vs. 21.0-24.0); smaller lower jaw length than B.
macrostoma (32.9-38.5% HL, vs. 38.0-54.0); lower range ofpredorsal length than all members (except B. macrostoma)(64.6-69.4% SL, vs. 67.5-73.0).
Betta patoti is distinguished from B. pallifina by having amore distinctly lanceolate caudal fin in adults (vs. lesspronounced); vertical dark bars on lower half of body (vs.
absence); no dark opercle pattern (vs. presence); no hyalineor whitish subdistal borders on anal and caudal fins in female(vs. presence); no vertical barring pattern on interradial
membranes of caudal and anal fins (vs. presence); slightly
more vertebrae (33-34, mode 33, vs. 31-33, mode 32); slightly
more anal-fin rays (30-33, mode 31, vs. 27-32, mode 30).

Description. – General appearance in Figs. 43a-b, 49f-g.
Meristic and morphometric data in Table 5. Body long andslender (body depth 20.0-22.6% SL); head blunt and broad(HL 29.7-31.4% SL, head width 18.0-20.7% SL, 57.9-67.2%
HL). Dorsal pointed, anal bluntly pointed, caudal strongly
lanceolate with median rays elongated. Pelvic withfilamentous first ray. Maximum known size 75.6 mm SL(ZMA 121.679).
Coloration. – For live coloration, see Figs. 49f-g. Body with
13-14 black bars on lower two-thirds, dorsum dark brown,
lateral light brown, ventrum light yellowish gold. Lower partof iris golden. Postorbital stripe distinct but interrupted.
Preorbital stripe distinct. Chin bar present, portion just belowab

Betta pallifina, new species(Figs. 44a-b, 49h)
Material examined. – Holotype - MZB 9328, 60.5 mm SL, male;
Borneo, Kalimantan Tengah: Muara Teweh, Kec. Laung Tuhup,
Desa Maruai, Rawa/Sungai Laung, tributary of Sungai Barito, coll.
D. J. Siebert, A. Tjakrawidjaja & O. Crimmen, 15-18 Jul.1992.

Fig. 43. a) Betta patoti – ZMA 112.513, lectotype, 63.5 mm SL, b)
ZRC 46323, 60.1 mm SL (right side, reversed).

Paratypes - MZB 6978, 60 ex., 20.4-66.3 mm SL; same locality
data as holotype. — BMNH 1996.3.22:106-138, 33 ex., 29.8-64.8

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Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei
mm SL, - ZRC 41576, 15 ex., 17.3-59.5 mm SL, - MZB 9329, 15
ex., 29.5-54.4 mm SL; Borneo, Kalimantan Tengah: Sungai Barito
basin, Sungai Laung at Desa Maruwei (0°21’59”S 114°44’6”E), coll.
D. J. Siebert, A. Tjakrawidjaja & O. Crimmen, 15-18 Jul.1992 (DS12-1992). — BMNH 1996.3.22:171-230, 60 ex., 23.2-62.3 mm SL,- ZRC 41577, 20 ex., 29.0-49.2 mm SL, - MZB 9330, 20 ex., 29.249.2 mm SL; Borneo, Kalimantan Tengah: Sungai Barito basin, Desa
Kerendan, Sungai Lahei and tributaries, coll. D. J. Siebert, A.
Tjakrawidjaja & A. Mun’im, 23-28 Sep.1992 (DS-3-1995). —BMNH 1996.3.22:48-57, 10 ex., 41.2-66.3 mm SL; Borneo,
Kalimantan Tengah: Sungai Barito basin, Sungai Idow above
waterfall (20 m high) ca. 7 km upstream from Sungai Kerendan(tributary of Sungai Lahei), coll. D. J. Siebert, A. Tjakrawidjaja &A. Mun’im, 26 Sep.1992 (DS-5-1995). — BMNH 1996.3.22:1-38,
38 ex., 15.3-61.0 mm SL; Borneo, Kalimantan Tengah: Sungai
Barito basin, Sungai Barito at Muara Laung (0°35’34”S
114°44’12”E), coll. D. J. Siebert, A. Tjakrawidjaja & O. Crimmen,
8 Jul.1992 (DS-7-1992).

Jul.1992. — MZB 7152, 8 ex., 23.4-45.9 mm SL; Borneo,
Kalimantan Tengah: Sungai Murung basecamp, Project Barito Ulu,coll. D. J. Siebert, A. Tjakrawidjaja & O. Crimmen, 11-13 Feb.1991.
— MZB 7075, 2 ex., 16.5-32.8 mm SL; Borneo, Kalimantan Tengah:
Laung Tuhup, Sungai Batu Iandai, tributary of Sungai Barito, 2 km
from Muara Laung, coll. D. J. Siebert, A. Tjakrawidjaja & O.
Crimmen, 8 Jul.1992. — MZB 7048, 4 ex., 28.5-35.3 mm SL;
Borneo, Kalimantan Tengah: Muara Teweh, Laung Tuhup, Desa
Maruei, Sungai Laung, tributary of Sungai Barito, coll. D. J. Siebert,
A. Tjakrawidjaja & O. Crimmen, 15-18 Jul.1992. — MZB 6917,
1 ex., 47.8 mm SL; Borneo, Kalimantan Tengah: Laung Tuhup,
DAS Barito, Sungai Maruai, tributary of Sungai Laung, coll. D. J.
Siebert, A. Tjakrawidjaja & O. Crimmen, 15 Jul.1992.

Diagnosis. – Betta pallifina is distinguished from the othermembers of the B. unimaculata group by the following
combination of characters: hyaline subdistal band on anal andcaudal fins of female; lanceolate caudal fin in adults; bodybrownish; lower half of caudal fin with dark distal edge;
vertebrae 31-33 (mode 32); anal-fin rays 27-32 (mode 30);
dorsal-fin rays 7-9 (mode 8); pectoral fin rays 11-14 (mode13); subdorsal scales 5-61/2 (mode 6); lateral scales 32-34(mode 33); predorsal scales 21-26 (mode 23); predorsal length64.6-69.4% SL; head length 26.8-33.4% SL; length of analfin base 47.4-57.7% SL; length of dorsal-fin base 9.3-12.5%
SL; lower jaw length 30.6-39.6% HL

Others - BMNH 1991.9.18:21-25, 5 ex., 22.5-52.1 mm SL; Borneo,
Kalimantan Tengah: Sungai Barito basin, Sungai Murung near
Project Barito basecamp, coll. D. J. Siebert, A. Tjakrawidjaja & O.
Crimmen, 11-13 Feb.1991 (DS-17-1991). — BMNH 1991.9.18:2931, 3 ex., 28.4-47.7 mm SL; Borneo, Kalimantan Tengah: Sungai
Barito basin, Sungai Barito at Muara Laung and market at Puruk
Cahu, coll. D. J. Siebert, A. Tjakrawidjaja & O. Crimmen, 20-21
Feb.1991 (DS-20-1991). — BMNH 1996.3.22:41-42, 2 ex., 41.942.4 mm SL; Borneo, Kalimantan Tengah: Sungai Barito basin,
Sungai Bunes, a small tributary of Sungai Kerendan (a tributary ofSungai Lahei) above (ca. 1 km) Desa Kerendan, coll. D. J. Siebert,
A. Tjakrawidjaja & A. Mun’im, 24 Sep.1995 (DS-2-1995). —BMNH 1996.3.22:76-105, 30 ex., 21.2-50.3 mm SL; same data as
holotype. — BMNH 1996.3.22: 231-253, 23 ex., 20.0-43.4 mm SL;
Borneo, Kalimantan Tengah: Sungai Barito basin, Desa Kerendan,
Sungai Lahei and tributaries, coll. D. J. Siebert, A. Tjakrawidjaja& A. Mun’im, 23-28 Sep.1992 (DS-3-1995). — BMNH 1996.3.22:
58-75, 18 ex., 9.0-44.1 mm SL; Borneo, Kalimantan Tengah: Sungai
Barito basin, Sungai Idow above waterfall (20 m high) ca. 7 km
upstream from Sungai Kerendan (tributary of Sungai Lahei), coll.
D. J. Siebert, A. Tjakrawidjaja & A. Mun’im, 26 Sep.1992 (DS-51995). — MZB 7019, 11 ex., 19.8-55.9 mm SL; Borneo, Kalimantan
Tengah: Laung Tuhup, Sungai Lubi, trubutary of Sungai Barito, 1
km from Muara Laung, coll. D. J. Siebert, A. Tjakrawidjaja & O.
Crimmen, 15 Jul.1992. — MZB 8237, 42 ex., 19.3-47.1 mm SL;
Borneo, Kalimantan Tengah: Laung Tuhup, Sungai Barito at Muara
Laung, coll. D. J. Siebert, A. Tjakrawidjaja & O. Crimmen, 8

Description. – General appearance in Figs. 44a-b, 49h.
Meristic and morphometric data in Table 5. Body slenderand long (body depth 20.5-23.0% SL); head broad and blunt(HL 26.8-33.4% SL, head width 17.7-20.8% SL, 56.4-70.2%
HL). Dorsal fin pointed, anal fin bluntly pointed, caudal fin
lanceolate with median rays elongated. Pelvic fin withfilamentous tip. Maximum known size 66.3 mm SL (BMNH
1996.3.22:48-57).
Coloration. – For live colour, see Fig. 49h. Both male andfemale specimens with patches of iridescent blue scales onbody and opercle. In preserved specimens, dorsum darker
brown than ventrum; vertical bar pattern on interradial
membranes of anal and caudal fins, absent on dorsal fin. Thinblack distal margin present on anal fin in male. A dark blotch
present just after hypural plate below lateral series on anterior
portion of caudal fin in both sexes and juvenile. Male with
distinct ‘pale cheek’ pattern on opercle, faint in female. Some
specimens with 4-6 irregular vertical bars across body. Female
sometime with distinct longitudinal reticulated pattern on
lower half of body, others more frequently with hyaline or
whitish (orangish iridescence when live) subdistal border withdark anal edge on anal fin and lower half of caudal fin. Pelvic
fin with filamentous tip black and rest darkly pigmented.

aDistribution. – Betta pallifina is currently known only fromthe upper Barito in Kalimantan Tengah (Fig. 23).

b

Field notes. – Betta pallifina is known only from headwater
habitats and is probably a paternal oral-brooder, similar tothe other members of B. unimaculata group. Syntopic Betta
present - B. obscura.
Fig. 44. a) Betta pallifina – ZRC 41577, male, 56.9 mm SL, b) ZRC
41577, female, 47.3 mm SL.

Etymology. – The name comes from the Latin pallidus

94THE RAFFLES BULLETIN OF ZOOLOGY 2005
meaning pale and finis meaning boundary, referring to theunique coloration of the bordered anal and caudal fins of thefemales. Used as a noun in apposition.

pelvic fin falcate; dorsal fin rounded, anal fin pointed, anal
fin reaching to half or more of caudal fin. Meristic andmorphometric data in Table 7. Maximum known size 28.2mm SL (CAS 94155).

Comparative notes. – Betta pallifina is distinguished fromthe other species in the species group by the following
characters: caudal fin lanceolate in adults (vs. rounded), acharacter shared only with B. patoti; female with hyaline towhitish subdistal margin (orange iridescence when live)
before dark edge (vs. absence); lower mode of vertebrae thanB. ocellata, B. patoti and B. gladiator (32, vs. 33-34); moreanal-fin rays than B. macrostoma (27-32, vs. 25-28); fewer
dorsal-fin rays than B. macrostoma (7-9, vs. 9-11); fewer
pectoral-fin rays than B. gladiator (11-14, vs. 16-17); greater
mode of subdorsal scales than B. unimaculata and B. ocellata(6, vs. 5-51/2); lower mode of subdorsal scales than B.
macrostoma (6, vs. 7); lower mode of lateral scales than B.
patoti and B. gladiator (33, vs. 34-35); dorsal-fin origin
modally above 18th lateral scale (vs. 19-20th in B. gladiator);anal-fin origin modally below 6th lateral scale (vs. 7-8th);and lower mode of predorsal scales than B. unimaculata, B.
ocellata, B. patoti and B. gladiator (23, vs. 24-26). In
preserved specimens, B. pallifina has an opercle patternsimilar to that of B. macrostoma in which a pale central regionof the opercle is surrounded by darkly pigmented area in bothmale (distinct) and female (faint).

Coloration. – Male with dark brown head; opercle black ,postorbital stripe present, chin-bar absent. Body light brown.
Dorsal fin with up to 5 rows of rounded black marks oninterradial membrane; caudal fin with up to 9 rows oftransverse bars; anal fin with thin black margin; pelvic fin
with extreme tip white, rest hyaline; pectoral hyaline. Body
with up to 7 vertical rows of dark marks on posterior edgeof scales. Female without black opercle, chin-bar andpostorbital stripe present. Body with distinct central andsecond central stripes, with caudal peduncle spot. Fins
yellowish to hyaline, dorsal and caudal fins with transverse
bars.
Distribution. – Betta stiktos is known from the Mekong
drainage in Cambodia.
Etymology. – The name comes from the Greek stiktos
meaning dappled, spotted, referring to the rows of diagnostic
round spots on the dorsal fin. Used as a noun in apposition.
Comparative notes. – Betta stiktos is distinguished from theother members of the species group by having: rounded blackspots on interradial membranes of dorsal fin (vs. thin verticalbars); up to 9 rows of dark transverse bars on caudal fin (vs.
absence); up to 7 vertical rows of dark marks on posterioredge of scales on body (vs. 8 or more); more lateral scales(33-34, vs. 29-31); more predorsal scales than B. imbellis (2425, mode 25, vs. 21-24, mode 23); smaller postdorsal lengththan B. smaragdina (17.5-20.1% SL, vs. 21.4-23.2); smaller
body depth than B. splendens (26.2-27.7% SL, vs. 27.1-32.2);
smaller caudal peduncle depth than B. splendens and B.
smaragdina (15.4-17.4% SL, vs. 18.1-20.3); smaller lengthof dorsal-fin base than B. splendens (15.2-15.6% SL, vs. 16.723.8); greater orbit diameter than B. splendens (27.7-29.7%
HL, vs. 22.7-26.6); smaller postorbital length than B.
splendens (45.9-50.5% HL, vs. 49.6-52.7); smaller interorbital
width than B. splendens and B. smaragdina (26.4-31.6% HL,
vs. 30.2-34.3).

BETTA SPLENDENS GROUP
Betta stiktos, new species(Fig. 45)
Material examined. – Holotype - CAS 204326, 27.3 mm SL male,;Cambodia: Mekong basin, small swampy stream from Stung Treng
to Ban Lung (ca. 3/4 to bridge over Tonle Srepok) (ca. 13°30’N
106°30’E), coll. T. R. Roberts, 16 Feb.1994.
Paratypes - CAS 94155, 19 ex., ZRC 43391 (ex CAS 94155), 3

ex., 16.7-28.2 mm SL; same locality as holotype.
Diagnosis. – Betta stiktos is distinguished from othermembers of the B. splendens group by the following
combination of characters: up to 5 rows of black rounded
marks on interradial membranes of dorsal fin; up to 9 rowsof transverse black bars on caudal fin; up to 7 vertical rowsof dark marks on posterior edge of scale on body; lateral scales
33-34 (mode 33), predorsal scales 24-25 (mode 25);
postdorsal length 17.5-20.1% SL; body depth 26.2-27.7% SL;
caudal peduncle depth 15.4-17.4% SL; length of dorsal-finbase 15.2-15.6% SL; orbit diameter 27.7-29.7% HL;
postorbital length 45.9-50.0% HL; interorbital width 26.431.6% HL.
Description. – General body shape and appearance in Fig.
45. Body slender (body depth at dorsal-fin origin 26.2-27.7%
SL); head small (HL 27.7-30.3% SL); caudal fin rounded;

Fig. 45. Betta stiktos – CAS 204326, holotype, 27.3 mm SL (rightside, reversed).

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Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei

a

e

b

f

c

g

d

h

Fig. 46. a) B. bellica – not preserved, ca. 50 mm SL (taken by Koji Yamazaki), b) B. pugnax – ZRC 39264, 59.6 mm SL, c) B. pulchra– not preserved, ca. 40 mm SL (taken by Koji Yamazaki), d) B. stigmosa – ZRC 43393, 37.5 mm SL, e) B. lehi – not preserved (taken by
Koji Yamazaki), f) B. akarensis – not preserved, ca. 60 mm SL, g) B. balunga – ZRC 35497, 39.2 mm SL, h) B. chini – ZRC 35086, holotype,51.0 mm SL.

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a

e

b

f

c

g

d

h

Fig. 47. a) B. ibanorum – not preserved, ca. 60 mm SL, b) B. macrostoma – not preserved (taken by Günther Ettrich), c) B. macrostoma– not preserved (taken by Koji Yamazaki), d) B. ocellata – ZRC 40956, ca. 40 mm SL, e) B. ocellata – not preserved, ca. 35 mm SL (taken
by Koji Yamazaki), f) B. gladiator – not preserved, ca. 50 mm SL, g) B. taeniata – not preserved, ca. 35 mm SL, h) B. taeniata – notpreserved, ca. 40 mm SL.

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Tan & Ng: Fighting fishes of Singapore, Malaysia and Brunei

a

e

b

f

c

g

d

h

Fig. 48. a) B. imbellis – not preserved, ca. 30 mm SL (taken by Koji Yamazaki), b) B. imbellis – not preserved, ca. 30 mm SL (taken by
Koji Yamazaki), c) B. coccina – not preserved, ca. 30 mm SL, d) B. coccina – not preserved, ca. 30 mm SL, e) B. livida – not preserved,
ca. 30 mm SL, f) B. persephone – not preserved, ca. 25 mm SL, g) B. tussyae – not preserved, ca. 30 mm SL, h) B. brownorum – notpreserved, ca. 25 mm SL.

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a

e

b

f

c

g

d

h

Fig. 49. a) B. waseri – ZRC 35402, 80.7 mm SL, b) B. hipposideros – ZRC 18689, paratype, 71.1 mm SL, c) B. tomi – ZRC 35411, paratype,64.9 mm SL, d) B. pi – not preserved, ca. 40 mm SL, e) B. unimaculata – ZRC 46324, ca. 45 mm SL, f) B. patoti – ZRC 46322, 48.2 mm
SL, g) B. patoti – not preserved, ca. 60 mm SL, h) B. pallifina – BMNH (taken by Darrell Siebert) (female above, male below).

99