Hydrobiologia 285: 41-48, 1994.

A. Sasekumar, N. Marshall & D. J. Macintosh (eds), Ecology and Conservation of Southeast Asian Marine and 41
Freshwater Environments including Wetlands.
1994 Kluwer Academic Publishers. Printed in Belgium.

Zoogeography and biodiversity of the freshwater fishes of

Southeast Asia

Mohd. Zakaria-Ismail
Department of Zoology, University of Malaya, 59100 Kuala Lumpur, Malaysia

Key words: zoogeographic regions, freshwater fishes, local extinction

Abstract

The ichthyofauna of the freshwater system of Southeast Asia is extremely diverse. A recent estimate of
about 1000 species is probably an understatement. More than 10 new species are being added to the
list annually. The distribution pattern of the Southeast Asian freshwater fishes can be divided into five
zoogeographic regions. The first one is the Salween basin in Burma, with fishes mainly of the Indian
subcontinent origin such as Amblypharyngodon atkinsoni, Bangana almorae and Brachydanio jayarami.
The second zoogeographic area is the Mekong, Chao Phraya and Mae Khlong drainages which harbour
fishes typical of the mainland of Southeast Asia such as Acanthorhodeus deignani, Barbichthys nitidus and
Cirrhinus siamensis. The Malay Peninsula is the third region whose species composition is heavily in-
fluenced by the Siamese (such as Homalopterasmithi, Tuberoschisturabaenzigeri and Botia beauforti) and
Indonesian (such as Botia hymenophysa, Luciocephalus pulcher and Parosphromenus deissneri) elements.
The islands of Sumatra, Borneo and Java are the fourth zoogeographic area of fish distribution. These
islands show a high degree of endemism, especially in fishes of the family Belontiidae. Finally, the
freshwater system of the Philippines is the last zoogeographic region of Southeast Asia. The area is
characterized by the presence of closely related species of the cyprinids especially in Lake Lanao.
Currently, high diversity of these freshwater fishes is being threatened by land development, such as
deforestation, road construction and land expansion for plantation. Recent studies in the Gombak River
basin show the extermination of 41 per cent of native fish species from 1969 to 1990. This is probably
due to the construction of highways, logging, as well as land clearing for agriculture.

Introduction graphic regions, with the publication of his clas-

Southeast Asia, which includes Burma, the Indo-
Chinese and Malayan peninsulas, the Indo-
Malayan archipelago, and the Philippines, has
been a focus of zoogeographic interest. Although
the foundations of biogeography can be traced
to the writing of Georges Louis Leclerc, comte de
Buffon and Augustin Pyramus de Candolle
(Nelson, 1978), it was Alfred Russel Wallace, a
British naturalist and codiscover of the theory of
evolution, who elaborated the concept of zoogeo-
sic work, The Geographical Distribution of Ani-
mals, in 1876.
Wallace (1860) proposed a hypothetical bound-
ary between the Oriental and Australian faunas
when he said 'we may consider it established that
the Strait of Lombok marks the limit and abruptly
separates two of the great zoological regions of
the globe'. Later, Huxley (1868) named the
boundary Wallace's Line and modified its limits
on the basis of bird studies placing the Philippines
east of the line within the Australian realm. As the
42
fauna of the region became better known, many
examples of animal groups crossing Wallace's line
were noted. For this reason, most zoogeographers
today do not take Wallace's line literally as an
exact boundary between the Oriental and Aus-
tralian fauna but rather a boundary of major fau-
nal break separating the relatively rich Oriental
continental fauna from the depauperate Austra-
lian island fauna. This demarcation is apparent
with respect to the distribution of freshwater
fishes.
Myers (1949, 1951) developed an ecological
classification of freshwater fishes based on their
tolerance to salt water. The primary division fish
families are those whose members have little salt
tolerance and are confined to freshwaters, al-
though salt tolerance of individuals within a fam-
ily does vary. Secondary freshwater fishes are
salt-tolerant families, whose origin can be traced
to marine relatives, that live chiefly in freshwaters
but sometimes enter the sea and can survive there
for a limited time. The peripheral division fresh-
water fishes are those that occur in freshwaters
but are actually semi-marine, or migratory, or de-
rived from marine families. Although these divi-
sions are not absolute and do not apply strictly to
whole taxonomic units, their usefulness in zoo-
geography cannot be denied.
Freshwater fishes of Southeast Asia are di-
verse. Unfortunately, up-to-date references on
these fishes are scarce, especially at the generic
revision. Recent generic revision works include
Sontirat (1976) on Cyclocheilichthys, Karnasuta
(1981) on Osteochilus, Roberts (1982a, 1982b,
1983, 1986) on Wallago, Chaca, Bagarius and
Macrognathus,Roberts & Vidthayanon (1991) on
Pangasius,Parenti (1989) on Phallostetus, Siebert
(1991) on Acanthopsoides, and Burridge (1992) on
Acanthophthalmus. More revisional works are re-
quired, such as on Lobocheilos, Mystacoleucus,
Neolissochilus and 'Puntius', before we can esti-
mate the number of species present in the area
with some degree of confidence.
The objective of this paper is to summarize
patterns of distribution and diversity of the fresh-
water fishes of Southeast Asia based on current
and historical data of fish taxonomy. An example
of changes of fish diversity and its probable cause
in Peninsular Malaysia is also presented.
Materials and methods
Observations and speculations on zoogeography
and diversity of the freshwater fishes of Southeast
Asia are primarily based on the evidence pre-
sented by various workers such as Anderson &
Collette (1991), Inger & Chin (1962), Kottelat
(1985, 1989, 1990), Ng & Lim (1990), Parenti
(1989), Roberts (1980, 1982a, 1982b, 1986, 1989),
Smith (1945), Taki (1974), Uwa & Magtoon
(1986) Weber & de Beaufort (1913, 1916), and
Zakaria-Ismail (1987, 1989). Studies on changes
in the species composition in the Gombak River,
Peninsular Malaysia were carried out in 1985 and
1990 to provide evidence of the impact of land
development on fish communities. Data of these
studies were compared with the baseline infor-
mation of Bishop (1973). Fish diversity in the
Kerling River, Peninsular Malaysia was deter-
mined by exhaustive sampling with an electro-
shocker to observe some pattern of dominance of
the various groups of fish in a forest that was
logged forty years ago.
Results and discussion
There are many ways to classify the pattern of fish
distribution of Southeast Asia. Kottelat (1989)
for example identified seven zoogeographic re-
gions for the freshwater fishes of Southeast Asian
mainland on the basis of the main river systems.
They are the Salween, Mae Khlong, Chao Phraya
and Mekong river drainages and Annam, south-
east Thailand and the Malay Peninsula. For the
purpose of this paper, only five main zoogeo-
graphic regions are used (Fig. 1). They are the
Salween basin in Burma, the Indo-Chinese pen-
insula which includes the Mekong, Chao Phraya
and Mae Khlong river drainages in Thailand and
Kampuchea, the Malay Peninsula, the Indo-
Malayan archipelago, and the Philippines.
The Salween River, with a length and drainage

Fig. 1. Major zoogeographic regions of the freshwater fishes
of Southeast Asia. ICP, Indo-Chinese peninsula; IMA, Indo-
Malayan archipelago; M, Mindanao; MP, Malay Peninsula;
S, Salween basin.
area of about 2800 km and 330 km 2 respectively,
is the second largest river system in Southeast
Asia. This area has little in common in terms of
the cyprinid fish composition with other zoogeo-
graphic regions in Southeast Asia. The majority
of species in the area are of Indian Sub-continent
origin such as Amblypharyngodon atkinsoni, Ban-
gana almorae and Brachydaniojayarami.
In his recent study of the zoogeography of
Southeast Asian mainland freshwater fishes,
Kottelat (1989) indicated that the Salween basin
is the eastern limit of the range of several genera
widely distributed in India (e.g. Amblypharyn-
godon, Aspidoparia and Chagunius) as well as the
western limit of the range of many Southeast
Asian genera (e.g. Albulichthys, Amblyrhynchich-
thys and Barbichthys). Only those genera that have
wide distribution is Asia such as Notopterus and
Mystus, montane forms such as Garra and Neo-
43
lissochilus, and genera whose systematics are
poorly known such as Puntius are found on both
sides of the border.
The Indo-Chinese zoogeographic region har-
bours fishes of typical of the mainland of South-
east Asia such as Acanthorhodeus deignani, Bar-
bichthys nitidus and Cirrhinussiamensis. This area
also shows a sizeable gap in the distribution of
some genera of catfishes. The angler catfish of the
genus Chaca comprises two species. Chaca chaca
occurs in Burma and also India, while Chaca ban-
kanensis is present only in the southern part of the
Malay Peninsula, Sumatra and Borneo. A simi-
lar pattern of distribution can also be observed in
Amblyceps mangois which is present in India and
the Malay Peninsula but not in the Indo-Chinese
peninsula.
The Malay Peninsula is the third zoogeographic
region. Species composition is heavily influenced
by Siamese as well as Indonesian elements. Many
Bornean and Sumatran species have their north-
ernmost distribution limit in the peninsula. As an
example, Luciocephalus pulcher is not known to
exist in the Indo-Chinese peninsula, but is com-
monly found in the Malay Peninsula especially in
blackwaters. Other species with a similar pattern
of distribution are Osteochilus spilurus, Parosph-
romenus deissneri and Sphaerichthys osph-
romenoides. Several Indonesian species are also
found only in the Malay Peninsula but not par-
ticularly restricted to the blackwaters; they are
Botia hymenophysa, Chaca bankanensis, Chela
maassi, Crossocheilus langei, Doryichthys deokha-
toides, Mystus bimaculatus, Osteochilus enneaporos,
Oxygaster anomalura, and Vaillantella maassi.
Several Siamese as well as Indian species have
their southernmost distribution limit in the pen-
insula. These species include Amblyceps mangois,
Botia beauforti, Garracambodgiensis, Glyptothorax
callopterus, Homaloptera smithi, Mystacoleucus
chilopterus, Nemacheilus masyae, Paralaubuca
typus, Puntioplites proctozysron, Rasbora borape-
tensis and Tuberoschistura baenzigeri. Thus, the
Malay Peninsula has become an area of overlap
for the Indian, Indo-Chinese and Indonesian
fishes.
Based on available literature, I estimate more
44
than 1000 species of fishes present in the fresh-
water system of Southeast Asia. The Salween
basin has some 150 species representing 77 gen-
era, of which 55% are shared with the lower
Mekong, 60 % with the middle Mekong, 60 % with
the Chao Phraya, 87 % with the Irrawaddy, 77 %
with the Brahmaputra, and 71 % with the Ganges.
Thus, the fish diversity is comparable to that of
the Indian sub-continent. Two genera (2.5 %) are
endemic to the basin, and both are known only
in In16 Lake (Kottelat, 1989).
The Indo-Chinese peninsula is probably the
most diverse zoogeographic area in Southeast
Asia. The Mekong is the longest and has the
largest drainage area of all rivers in Southeast
Asia. High diversity of fishes in the Mekong is
expected. More than 500 species (Smith, 1945;
Taki, 1975; Rainboth et al., 1976; Kottelat, 1985;
Mai & Nguyen, 1988) are present in the area.
Kottelat (1989) estimated some 300 species of
freshwater fishes from the Indo-Chinese parts of
the Mekong and Chao Phraya basins alone. Of
the 103 genera found in the region, 67% are in
common with the Sundaic forms, whereas only
47 % of the genera have a strong affinity with the
Salween basin. Forty-eight percent of the species
are endemic to both rivers. A high degree of en-
demism suggests that the basin has not had water
Poecilidae
Hemiramphidae 9.6%
Channidae
Fig. 2. Ichthyofauna composition of the Pusu River, Peninsular Malaysia.
exchange with other systems for a long period of
time.
Great diversity of the ostariophysan fishes is
found in the Indo-Chinese peninsula. Many stud-
ies have shown that the area has the highest con-
centration of this group of fishes especially the
Balitoridae (Kottelat, 1989), Sisoridae (Roberts,
1983) and Cobitidae (Sawada, 1982). Although
the Indo-Chinese peninsula is probably not the
centre of origin of ostariophysan fishes (Regan,
1922; Myers, 1967; Roberts, 1975), it is probably
a place of rapid speciation of these fishes.
Although the Malay Peninsula is only slightly
larger than the Salween basin, its diversity is much
higher (260 vs. 150). According to Kottelat (1989)
15% of the species are shared with the Salween
basin, 47% with Chao Phraya, 44% with the
Mekong and 66% with the Indo-Malayan archi-
pelago. Peninsular Malaysia, the southern half of
the Malay Peninsula, harbours more than 200
species (Zakaria-Ismail, 1989) four of which
are endemic. They are Neolissochilus hendersoni,
Poropuntius birtwistlei, P. smedleyi, Parosph-
romenus nagyi and P. paludicola. These endemic
species are confined either to montane streams or
blackwater swamps.
The Indo-Malayan archipelago is the fourth
zoogeographic region. The area, which includes
. _ _- _....__. -A
ae 34."/0%
Clariidae 9.6%
ae 9.6%
Belontiidae 9.6%

the islands of Sumatra, Borneo and Java, has
more than 400 species. Some 290 species
(Roberts, 1989) inhabit the Kapuas River and it
is perhaps the most diverse fish community in the
region. High fish diversity is also seen in the
Baram River, in which more than 115 species of
fish were found (E. J. Crossman, pers. com.). The
presence of endemic species of the perciforms
such as Belontiidae, Helostomatidae, Osphrone-
midae, and Luciocephalidae suggest this area is a
centre of speciation of this perciform group.
The freshwater habitat of Southeast Asia is
dominated by the cyprinid fish. Some 70 genera
of cyprinids are endemic to the area. Many other
genera found here have a few representatives
living outside the region. Some regions of South-
east Asia, especially rivers of the Indo-Chinese
peninsula that traverse different faunal regions
have components of different cyprinid fauna. For
instance, the head waters of upper Mekong on the
Tibetan Plateau has greater numbers of high
Asian fauna. As it passes through Yunnan Pro-
vince, components of the East Asian fauna domi-
nate, while components of Southeast Asian fauna
Cyprinidae 53.8%
Hemiramphidae
Belontiidae 7.7%
Fig. 3. Ichthyofauna composition of the Kerling River, Peninsular Malaysia.
45
are more apparent throughout its middle and
lower reaches (Rainboth, 1991).
Using the freshwater fishes of Thailand as an
example (Smith, 1945), the cyprinoids consist of
more than 40% of the total fauna. In the study of
the ichthyofauna of the Mekong River, Taki
(1978) discovered that 49% of the fauna are cyp-
rinoid fishes, and the percentage of cyprinoids
increased progressively upstream, reaching 60%
in the upper middle Mekong. In a study of the two
small streams in Peninsular Malaysia, I observed
a similar pattern of cyprinid fish diversity. In the
Pusu River, a tributary of the Gombak River that
runs through old rubber estates, the fish commu-
nity is dominated by the cyprinid fishes followed
by the catfishes, snakeheads and some other
groups (Fig. 2). An increase in the percentage of
the cyprinid fish was observed in the Kerling
River, a second order stream that runs through a
secondary forest which was logged 40 years ago
but has since been left for forest regeneration.
Here, more than half of the fish community are
the cyprinid fish (Fig. 3).
The diversity of cyprinids on Mindanao, Phil-
Cobitidae 7.7%
idae 7.7%
ULlarllaae /./o
46

Table 1. Fish species composition irn the Gombak River has often been cited as an example of explosive
Selangor in 1969, 1985 and 1990. Datta for 1969 is based on speciation in the last 10000 years (Myers, 1960).
Bishop (1973), and his sampling tech niques as well as sam-
pling locations were repeated for the i1985 and 1990 studies. Although there has been some doubt regarding
Presence and absence of each species is indicated by + and this pattern of speciation (Reid, 1980), there is no
- respectively. question that the fauna is endemic, but the age of
the lake, the actual size of the fauna, the exact
Species of fish 1969 1985 1990 distribution of the members, and the length of
Family Poeciliidae isolation are certainly suspect (Rainboth, 1991).
Poecilia reticulata + + + Land developments exert a tremendous pres-
Family Cyprinidae sure on biodiversity of aquatic organisms, par-
Hampala macrolepidota + - - ticularly fishes. Reduction in the number of
Mystacoleucus marginatus
Neolissochilus soroides
species has always been the result of land devel-
Osteochilus hasseltii + _ + opments. As an example, Singapore has lost over
Poropuntiussmedleyi + ++ half of its freshwater fishes by 1960 (Alfred, 1961)
Puntius binotatus + + + due to urbanization. The best example in Malay-
Rasbora sumatrana + + + sia is observed in the Gombak River, a tributary
Family Bagridae
Mystus planiceps
of the Klang river drainage. Ichthyofauna diver-
Family Sisoridae sity in the river has been well documented
Glyptothorax major + + + (Bishop, 1973) in which 27 species were recorded
Glyptothorax platypogonoides + + - in the survey. I resampled the river seven times in
Family Clariidae 1985 and five times 1990, repeating exactly
Clarias batrachus
+ + + Bishop's sites and method, and observed sub-
Clarias teijsmanni
Family Siluridae stantial changes in fish structure of the river
Silurichthys hasseltii + + + (Table 1).
Family Hemiramphidae Data for 1985 indicate that seven species of
Dermogenys pussilus + - - fish have completely vanished from the river
Hemirhamphodon pogonognathus
+ + + which include two cyprinid fish (Hampala mac-
Family Synbranchidae
Monopterus albus + + + rolepidota), a pipefish (Doryichthys deokhatoides),
Family Syngnathidae an anabantid (Anabas testudineus), a belontiid
Doryichthys deokhatoides + - - (Trichogaster trichopterus), a halfbeak (Dermog-
Family Anabantidae enys pussilus) and a mastacembelid (Mastacembe-
Anabas testudineus
+ - - lusfavus). These species represent about 25.9%
Family Belontiidae
Trichogastertrichopterus + - _ of the total fish species in the river. The 1990
Family Channidae survey indicates the disappearance of four more
Channagachua + + + species (Glyptothorax platypogonoides, Mystus
Channa lucius + + + planiceps, Macrognathus maculatus and Channa
Channa melasoma + - - melasoma) from the river system. Thus far, about
Channastriata
Family Mastacembelidae 40.7 % of the total fish fauna of the Gombak River
Macrognathusmaculatus + + _ have completely disappeared since 1969. It is
Mastacembelusfavus + - - speculated that land developments around the
- rs 1 1r 1
Uombak watershed tor logging, agriculture, in-
dustry and housing, and highway construction
ippines is worth mentioning. The island is the contributed to the substantial reduction of fish
easternmost boundary of primaary freshwater fish diversity of the Gombak River.
in Southeast Asia. Of the 23 cyprinid species The use of illegal methods to catch fish can also
known from the island, 19 speecies are endemic. become a major factor in reducing the diversity of
Fifteen of them are found in Lalke Lanao, and this the freshwater fishes of Southeast Asia (Zakaria-

Ismail, 1991). Various kinds of insecticides and
cyanide have been illegally used in many rivers
that flow through remote, logged forested areas.
Recently, there is some evidence indicating the
use of explosives in some of these rivers. Beyond
any doubt, these indiscriminate methods coupled
with habitat destructions will have a negative im-
pact on freshwater fish communities in Southeast
Asia.
Acknowledgements
I thanks R. J. Behnke, T. R. Roberts and L. R.
Parenti for valuable discussion and encourage-
ment during the course of the study. Research
leading to this paper was supported by Ministry
of Science and Environment grants R&D 04-07-
04-048 to me at the University of Malaya.
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