Biological Conservation 1993, 64, 19-24
. lt. "~1
THE STATUS A N D IMPACT OF I N T R O D U C E D FRESHWATER
ANIMALS IN SINGAPORE
Peter K. L. Ng, L. M. Chou & T. J. Lam
Department of Zoology, National University of Singapore, Kent Ridge, Singapore 0511
(Received 2 December 1991; revised version received 27 March 1992; accepted 14 April 1992)
Abstract
Singapore's location at the centre of major air and shipping
routes has resulted in the introduction of numerous
species of freshwater animals from various parts of the
world. To date, some 37 introduced freshwater fishes,
turtles, molluscs and prawns have become established
here. Studies show that the introduced species have not
significantly affected the fauna in pristine forested
streams. This is probably because much of the native
freshwater fauna includes acid-water species, whereas
introduced ones prefer more neutral and harder waters.
Of the 54 native Singapore freshwater fishes, the greater
majority (over 80%) are forest species adapted to living
in acid waters. These species have suffered mainly because
of forest clearance and more than 50% of this fauna is now
extinct. A similar situation occurs for freshwater prawns.
The conservation of the native freshwater fauna in Singa-
pore should thus centre on the preservation of their natural
forest habitat. There is a potential danger posed by im-
ported fishes which originate from soft, acid waters in
Africa, South America or other parts of Southeast Asia.
Thus far, only one or two species have managed to estab-
lish themselves in Singapore. Legislation and appropriate
quarantine measures will be needed to ensure that alien
tropical acid water species do not become established here.
Keywords: Singapore, freshwater animals, introduced
species, competition, impact.
INTRODUCTION
Several factors have led to the rapid introduction of
many terrestrial and freshwater animals into Singapore
over the years. These include its status as a centre of
trade, its large immigrant population, the regular im-
port of live food fishes from neighbouring areas and
the popularity of aquatic pets. While there have been
many dramatic cases in other countries where foreign
species have displaced endemic fauna (Courtenay &
Stauffer, 1984; De Silva, 1989), none has been docu-
mented for Singapore although detailed local studies
have not been made. The degree of development in
Singapore has meant that the habitats of many areas
were drastically changed well before many of the im-
portant introductions occurred (Wee, 1964; Wee &
Biological Conservation 0006-3207/93/$06.00 © 1993 Elsevier
Science Publishers Ltd, England. Printed in Great Britain
19
Corlett, 1986; Ng, 1991a). The loss of primary and late
secondary forest has caused the reduction in numbers
or even extinction of many aquatic forest taxa, the most
notable being that of freshwater fishes. The number of
extant native freshwater fish species in Singapore now
stands at 31 (Ng & Lim, 1989a, 1990a, in press; Lim &
Ng, 1990). This means that of the 54 species recorded,
23 are now believed to be extinct. At least 14 non-
endemic fish species are known to have become
established and breeding in Singapore (Lim & Ng,
1990; Ng & Lim, in press), 16 are regularly introduced
but probably do not breed, and the status of several
others has yet to be determined (Table 1).
The most common fishes in Singapore fresh waters
are almost all introduced and dominate the drains,
canals, reservoirs and most open-water bodies--the
guppy Poecilia reticulata, mollies Poecilia sphenops and
P. latipinna, mosquito-fish Gambusia holbrookii and
tilapia Oreochromis mossambicus, etc. and hybrids)
(Lim & Ng, 1990). The introduction of tilapia by the
Japanese during the Second World War (intended as a
supplementary food source) was a 'great success'. This
highly adaptable fish has, however, become ubiquitous
in all major water bodies, though it is not popular
locally and has almost no market value. Up to the
mid-1970s, Gambusia was relatively uncommon (Alfred,
1966; Johnson, 1973), but now occurs in large numbers
in most of the reservoirs (Ng & Lim, in press). Another
recent but successful introduction is an ambassiid, the
glass fish Parambassis punctulatus (Lim & Ng, 1990 (as
Chanda sp.); Ng & Lim, in press), which now occurs in
large numbers in Peirce Reservoir.
HISTORY OF FISH INTRODUCTIONS
Our knowledge of freshwater fish introductions prior to
the 1970s is largely due to the efforts of Alfred (1961,
1966) and Johnson (1973) (see also Johnson & Soong,
1963). Between them, they identified only eight species
as being introduced, although the indigenous status of
some species remains debatable. Alfred (1966) believed
the giant snakehead Channa micropeltes was introduced
as there were no early records of the fish despite
numerous collections. However, it is native to this part
of the world, and many earlier collectors sampled
selectively and sporadically, so it might have been
'neglected' or missed. A similar situation prevails for
 20 Peter K. L. Ng, L. M. Chou, T. J. Lam

Table 1. List and status of introduced freshwater organisms

Taxon Status a Taxon Status a

Class Teleostei Family Ambassidae

Family Osteoglossidae Parambassis punctulatus (Fraser-Brunner) ES
Scleropagesformosus (MOller & Schlegel) FE (ES ?) Amblypharyngodon chulabhornae Vidthyanan
& Kottelat FE.~
Family Cyprinidae
Rasbora borapetensis Smith ES Family Cichlidae
Esomus metallicus Ahl ES Oreochromis mossambicus (Peters) ES
Rasborinus lineatus takakii Oshima ES b Oreochromis aureus (Steindachner) ES?
Puntius partipentazona Fowler ES Oreochromis niloticus (Hasselquist) ES?
Puntius tetrazona (Bleeker) ES Tilapia zillii (Gervais) ES?e
Astronotus ocellatus FE
Puntius semifasciolatus (GOnther) ES
Leptobarbus hoeveni (Bleeker) FO Colossoma cf. macropomum FO
Cichlasoma meeki FE
Barbodes gonionotus (Bleeker) FO
Paracheirodon innesi FE
Ctenopharyngodon idella (Valenciennes) FO
Puntius conchonius FE
Cyprinus carpio Linnaeus FO/FE
Carasius auratus (Linnaeus) FE Class Amphibia
Hypophthalmichthys nobilis (Richardson) FO
Hypopthalmichthys molitrix (Valenciennes) FO Family Ranidae
Rana catesbiana (Shaw) FO
Family Pangasiidae
Pangasius sutchi Fowler FO/FE Class Reptilia
Family Loricariidae Family Emydidae
Pterygoplichthys sp. ES Trachemys scripta elegans (Wied) ES
Siebenrockiella crassicollis (Gray) FE
Family Poeciliidae Manouria emys FE
Poecilia reticulata Peters ES
Poecilia latipinna (Lesueur) ES? Family Trionycidae
Poecilia velifera ES? Pelodiscus sinensis (Wiegmann) ES
Poecilia sphenops Valenciennes ES Lissemys punctata Lacrprde FE
Gambusia holbrookii (Girard) ES Class Bivalvia
Xiphophorus variatus (Meek) ES?
Xiphophorus maculatus (G~inther) ES? Family Unionidae
Xiphophorus helleri Heckel ES? Pseudodon vondembuschianus (Lea) ES
Family Belontiidae Class Gastropoda
Betta imbellis Ladiges ES c Family Ampullaridae
Trichogaster pectoralis Regan ES Pomacea cf. canaliculata ES
Trichogaster microlepis ES?
Colisa lalia FE Family Vivipariidae
Viviparus polyzonata ES
Family Osphronemidae
Osphronemus goramy Lac6p~de FO/FE Phylum Arthropoda
Family Helostomatidae Subphylum Crustacea
Helostoma temminckii Cuvier FE
Family Palaemonidae
Family Channidae Macrobrachium lanchesteri (De Man) ES
Channa micropeltes (Cuvier) ES Macrobrachium nipponense (De Haan) ES

a FE, feral; ES, established; FO, food animal.

b Bhnhrescu (1967) recognised two subspecies of Rasborinus lineatus and regarded the Singapore population as belonging to R.
lineatus takakii.
c The status of Betta imbellis in Singapore today is uncertain. Tweedie (1952), Alfred (1966) and Johnson (1973) (as B. splendens)
all comment that is has become established, but feral populations of the species appear to have become very rare in recent years.
The change in the species name is recent. Schaller and Kottelat (1990) commented that specimens commonly known as B.
splendens in this part of the world actually belong to two very similar species--B, splendens Regan, 1910 and B. imbellis Ladiges,
1975. The true splendens is Thai whereas imbellis is Malaysian. They can also be separated by their body form (slimmer in B.
imbellis), coloration of their fins and behaviour. Schaller and Kottelat (1990) could recognise both species from Regan's (1910)
preserved type material. The specimens from Thailand were made the lectotype of B. splendens s. str.; with all Malaysian material
then referrable to B. imbellis. It is uncertain if B. imbellis was ever native to Singapore (see Tweedie, 1952). The origin of
Singapore's original stocks (assuming they were introduced) is not known, and they could well have come from both Thailand
and Malaysia. Examination of wild specimens collected from Singapore in the past (all rather small) show that they agree quite
well with what is known as B. imbellis. No large Singapore specimens were, however, available. Comparisons of this material and
Alfred's (1966: PI. 8, Fig. 3) excellent figure of B. splendens with a large series of fresh specimens of B. imbellis recently collected
from Johor further support our belief that the Singapore 'splendens' are most probably B. imbellis.
d Specimens collected by K. K. P. Lim from rural areas in Singapore have been identified to Amblypharyngodon chulabhornae,
recently described from northern Thailand (Vidthayanan & Kottelat, 1991). This fish probably reached Singapore accidently via
the aquarium trade as it is not a colourful or exotic species.
e Alfred (1968) lists Tilapia zillii as introduced to Singapore but we have not been able to find this species in recent years. It might

well have being replaced by O. mossambicus or its hybrids.

 Introduced freshwater animals in Singapore
the red-eyed barb Cyclocheilichthys apogon (Cyprinidae).
Johnson (1973) harboured doubts about its endemicity
although Alfred (1966) believed it was native. As recently
as 1990, however, taxa such as Channa melasoma and
Leiocassis aft. poecilopterus (Bagridae) were still
reported as new records from Singapore (Ng & Lim,
1990a, b in press).
Alfred (1966) systematically analysed all earlier
records of introduced and doubtful species. He noted
that other than the species cultured for food--Javan
barb Barbodes (Puntius s. lato aut.) gonionotus, grass
carp Ctenopharyngodon idella, common carp Cyprinus
carpio, bighead carp Hypophthalmichthys (= Aristich-
lhys aut.) nobilis and mud carp H. molitrix--only the
snakeskin gouramy Trichogaster pectoralis, Siamese
fighting-fish Betta imbellis, tilapia, molly, guppy and
mosquito-fish had become established. Of these, the snake-
skin gouramy and Siamese fighting-fish are Asiatic
species brought to Singapore from northern Malaysia.
The others are native to Africa or South America.
Most of the earliest fish species introduced into
Singapore by the Chinese immigrants in the early part of
this century have not established themselves in the wild.
These include Cyprinus carpio, Hypophthalmichthys nobilis
and Ctenopharyngodon idella, current stocks of which
are almost all cultured from imported fry. Breeding of
l:hese fishes is not carried out on any scale locally and
feral populations have not yet been known to breed.
Two cyprinids which did establish themselves, probably
brought to Singapore accidentally with the imported
carp-fry from China, are the green barb Puntius semi-
fasciolatus and Chinese minnow Rasborinus lineatus.
The case of P. semifasciolatus has been documented by
Alfred (1966) and this species is still extant in some
secondary forest areas. The history of Rasborinus is
very convoluted. Regan (1913) described it as a new
species, Rasborichthys altior, from Singapore specimens,
and for a long time it was believed to be Singapore's
only endemic fish (Alfred, 1966; Johnson, 1973). It was
only relatively recently that Bhnhrescu (1967) showed
that R. altior was actually a known Chinese species--
Rasborinus lineatus (see also Ng & Lim, in press). We
have not found Rasborinus in Singapore in recent years,
although Alfred recorded it as being common up to
1966. It is not unlikely that it has been displaced by
other introduced species such as Gambusia and Param-
bassis, which are far more common today than in 1966
(see Lim & Ng, 1990).
While Alfred (1966) and Johnson (1973) noted that
the local fighting-fish Betta imbellis (as Betta splendens,
~ee Table 1) was still common, it is certainly not the case
today. The reduction in area of rural land might be a
factor. The only recent record of this species was from
the Sime Road area in Singapore (Ng & Lim, in press).
It is uncertain when the tiger barbs Puntius partipenta-
zona and P. tetrazona became feral, although it was
possibly in the late 1970s or early 1980s. Certainly, by
1983, P. partipentazona had become well-established in
Seletar and other reservoirs and ponds, often in large
schools. P. tetrazona may have been more recent as it
21
has been found in some numbers in the Seletar area
only in the last five years. Both species have been
regarded by some authors as subspecies separated by
geography, but in Singapore inter-breeding can be
expected. We have encountered specimens, possibly
hybrids, whose precise identification is not possible.
Two northern Malaysian/Thai species--the red-tailed
rasbora Rasbora borapetensis and flying barb Esomus
metallicus--have become remarkably successful. The
time of introduction is not known, probably in the early
or mid-1980s, and almost certainly via the aquarium
trade. They are today very common in drains, ponds,
lakes and rural streams (Lim & Ng, 1990; Ng & Lim,
in press).
The status of two other cyprinids is also uncertain.
The river barb Barbodes schwanenfeldii and Hassett's
barb Osteochilus hasselti were reported by early
workers, but Alfred (1966) regarded both as being
extinct in Singapore. In recent years, specimens of both
have been seen or collected from or near reservoirs (Ng
& Lim, in press), and are likely to be reintroductions
via the aquarium trade. As both species previously
occurred in Singapore, they are not discussed with the
other introduced species here.
The South American loricarid catfish Pterygo-
plichthys (as Hypostomus in Lim & Ng, 1990) is also
now well-established in Singapore waters, adapting to a
wide variety of habitats (Ng & Lim, 1989b). It has also
become established in other parts of Southeast Asia
(Peninsular Malaysia and Java, unpublished data).
HISTORY OF OTHER VERTEBRATE AND
INVERTEBRATE INTRODUCTIONS
The Chinese softshell turtle Pelodiscus sinensis might
well have been here for several decades (Chou & Lam,
1989), and Lissemys punctata might have been recently
introduced (Yong, 1990). The same is true for terra-
pins, although Trachemys scripta is probably the only
truly established species (Lira & Chou, 1990). No
amphibian species is known to have established itself
although feral Rana catesbiana, the edible American
bullfrog (Lim & Chou, 1990) have been reported. Two
freshwater prawns (Macrobrachium lanchesteri and M.
nipponense) are introductions, probably entering with
the food fish or aquarium trade (Chong et al., 1987;
Ng, 1990). Two snails (Pomacea cf. canaliculata and
Viviparus polyzonata) have become residents, again
probably by the same channels as the prawns (Johnson,
1973; Ng, 1991a). Even a freshwater bivalve (Pseudo-
don vondembuschianus), hitherto unknown from Singa-
pore, has recently become established (Yang, 1990).
In summary, some 30 aquatic species have become
established in our fresh waters (Table 1).
FISH TRADES AS AGENCIES OF
INTRODUCTIONS
The past two decades have also seen the growth of the
aquarium industry, which grossed over US$ 35 million
22 Peter K. L. Ng, L. M. Chou, T. J. Lam
in 1989. While much of this was due to ornamental
varieties of goldfish, guppies and mollies, a substantial
sum (not always reflected in the official figures) is for
fishes brought into Singapore for re-export. These
come from all around Southeast Asia as well as
Australia, South America and Africa. Some are sold
locally. Released specimens and escapes have contributed
to introductions.
The aquarium/pet trade and culture of aquarium
species are responsible for most of the aquatic intro-
ductions into Singapore, where there is minimal control
of imported species. In many cases, animals escape or
are released for various reasons. While most fail to
establish themselves, some inevitably do. One must at
least be wary of species potentially capable of com-
peting or ousting local acid water species. Apart from
the loricariid catfish Pterygoplichthys, none of the
South American/Amazonian species of characins, tetra
and catfishes, many of which thrive in acid-waters,
have yet to establish themselves in Singapore, although
they are imported and sold in large numbers.
The Singapore government is a signatory of the
CITES treaty on endangered species, and it is illegal to
trade in fish which are on the list, e.g. Scleropages and
Arapaima. Nevertheless, a large number of endangered
species are still smuggled into Singapore and sold
illegally. A large number of other potentially threatened
species, especially Southeast Asian taxa, however, are
not on the list (Ng, 1991b), although there is an
attempt to revise the fish species under the CITES
treaty (M. Kottelat, pers. comm.).
Another source of introductions is via the food- and
sport-fish trade and the culture of food fishes, which
accounts for about 17% of Singapore's introduced
species. Not all feral species and introductions are the
result of escapes. Quite often, imported fish fry may
contain individuals of other species. Puntius semifascio-
latus and Rasborinus lineatus most probably entered
Singapore this way. Bivalve species such as Pseudodon
vondembuschianus probably entered Singapore as
parasitic glochidia larvae. Gastropods such as Viviparus
could also have entered Singapore similarly.
THE IMPACT OF INTRODUCTIONS ON THE
LOCAL FISH FAUNA
Despite the fears of Alfred (1966) and Johnson (1973),
there is no report of any introduced fish species in
Singapore having displaced a native one. This contrasts
with the experiences of some countries, notably Australia,
where introduced fishes have caused the decline or even
extinction of some native species (McKay, 1984, 1989;
Barlow et al., 1987; Arthington, 1989). Nevertheless,
Lim and Ng (1990) and Munro (1990) warn of the
potential dangers to local ichthyofauna if the pace of
introductions persists.
In undisturbed forest streams, where the waters are
peaty, acidic, well-shaded and cool, the fish present are
almost all native species. In open-country waters, over
50% of the species are introduced feral taxa. Even in
intermediate habitats (e.g. forest streams which join
open-country waters), up to 30% are feral. Non-
endemic species thus do not appear to migrate into
forest habitats.
The effects of introduced species on the local fishes
of open-country habitats are probably significant but
difficult to assess because most of these areas are so
disturbed or developed that no comparisons or studies
are possible. However, the open-country aquatic fauna
of Singapore is naturally probably quite poor. Of
the 54 native freshwater fish species known from
Singapore, only about 10 can be regarded as open-
country taxa (cf. Mohsin & Ambak, 1983; Roberts,
1989). Of these, only one is now believed to be extinct.
This is in sharp contrast to the extinction rate of the
obligate forest species, of which 50% are now believed
extinct. Most of the surviving 50% are highly threatened,
with very restricted distributions. The surviving open-
country species on the other hand have not declined
or disappeared in recent years. Species such as the
common snakehead Channa striata, swamp eel
Monopterus albus, climbing perch Anabas testudineus,
walking cat fish Clarias batrachus, common goby
Stigmatogobius sp., halfbeak Dermogenys pusillus and
whitespot Aplocheilus panchax are still quite common.
Most are very adaptable and can also survive in tree-
country and forest habitats. Only in more urbanised
areas where concrete canals and drains predominate
are they perhaps affected by introduced species. Even
then, species such as Channa striata, Monopterus albus
and Clarias batrachus are still present, although not
always as conspicuous as guppies and tilapia.
The fact that most introduced species in Singapore
are lacustrine (e.g cichlid) or hard water (e.g. poeciliid)
species makes it easier to understand why our native
fishes have apparently not suffered greatly at the
expense of the introductions (Fernando, 1990).
The recent establishment of the tiger barb Puntius
tetrazona is a matter of serious concern as it is capable
of thriving in the acid-water habitats of the native
species. Another possible cause for concern is that the
success of the immigrant fauna in disturbed areas may
prevent some of the native fauna from establishing
themselves in man-made environments. Not all our
native organisms are obligate forest-dwellers. Several
species, e.g. the common barb Puntius binotatus, possibly
the two spot rasbora Rasbora elegans, Dermogenys
pusillus, are highly adaptable, and may well survive in
more urban and built-up areas if immigrant species are
absent. Others such as Anabas testudineus, Monopterus
albus, Clarias batrachus, Aplocheilus panchax and
Channa striata may do even better.
THE IMPACT OF INTRODUCTIONS ON THE
NATIVE INVERTEBRATE FAUNA
While the emphasis has been on fish, it is worthwhile
also comparing the situation for the aquatic inverte-
brates. The apple snail Pomacea cf. canaliculata does
not appear to compete with the native Pila scutata.
 Introduced freshwater animals in Singapore
Pomacea is more adaptable to poor water conditions,
will graze out of water, feed on tougher plant material,
and seems to prefer open areas with hard substrates
(e.g. rocks). Its bright pink eggs are laid on bare rock
or plant surfaces without shade, sometimes several
metres from the water, whereas the white eggs of Pila
cling close to the water surface, usually among clumps
of grass and vegetation. The continued development
and the creation of open-country habitats, concrete
drains, etc. in Singapore will favour the survival and
proliferation of Pomacea.
As regards the prawns, neither Macrobrachium lan-
chesteri or M. nipponense appear to challenge any na-
tive species. Most o f the native species such as M.
malayanum and M. trompii prefer more acidic waters
and some degree o f shelter. The two introduced
prawns, on the other hand, prefer more disturbed and
open areas with harder and less acidic waters (see also
Ng, 1990).
The bivalve Pseudodon vondembuschianus fills an
apparently 'empty niche' in Singapore. It has long been
noted that Singapore has no native freshwater bivalves
(Johnson, 1973). As these filter-feeding bivalves have
parasitic glochidia larvae, they may conceivably have a
detrimental effect on the open water fishes. Lim (1990)
recently described new helminth gill parasites from
imported cultured catfish Pangasius sutchi in Peninsular
Malaysia, and it is likely that these parasites are also
present in Singapore as these fish are widely kept as
pets, food and game-fish.
A measure of government control and regulation
such as appropriate quarantine is needed to control the
growing trade in aquatic organisms in Singapore. We
have been fortunate so far that no indigenous fauna
has been eliminated by introduced species or drastically
reduced by exotic disease and parasites.
ACKNOWLEDGEMENTS
Thanks are due especially to M r Kelvin Lim for
assistance with the field work, and his help with the fish
and reptiles. Mr Richard Yeong, Joseph Tan and Ng
Hock Ping kindly assisted the first author during field
studies on numerous occasions over the years. The
National Parks Board kindly granted us permission to
conduct surveys in the Central Catchment Area. The
many interesting discussions with Drs Maurice Kottelat
and Tyson Roberts on numerous aspects o f fish
taxonomy are very much appreciated.
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