The non-native freshwater fishes of Singapore: an annotated compilation

Abstract

A total of 123 species of non-native freshwater fish, including seven possible hybrids, are recorded from the inland waters in Singapore. The majority (84 species, 68.3%) are from four families: Cyprinidae (37 species, 30.1%), Cichlidae (30 species, 24.4%), Osphronemidae (9 species, 7.3%), and Poeciliidae (8 species, 6.5%). Of these, 42 species-mainly cichlids (12 species)-are established in Singapore. The likely pathways of introduction and pertinent conservation issues are briefly discussed. Notes on local distribution, species used for biological control, dubious records, early records of native fish species, ornamental fish trade and aquacultural species are also provided. An addendum is included for four more species.

Full text

150
Tan et al.: Non-native freshwater shes of Singapore
The non-native freshwater shes of Singapore: an annotated compilation
Tan Heok Hui1*, Kelvin Lim Kok Peng1, Liew Jia Huan2,3, Low Bi Wei2, Rayson Lim Bock Hing2,
Jeffrey Kwik Teik Beng2 & Darren C. J. Yeo1,2
Abstract. A total of 123 species of non-native freshwater sh, including seven possible hybrids, are recorded from
the inland waters in Singapore. The majority (84 species, 68.3%) are from four families: Cyprinidae (37 species,
30.1%), Cichlidae (30 species, 24.4%), Osphronemidae (9 species, 7.3%), and Poeciliidae (8 species, 6.5%). Of
these, 42 species—mainly cichlids (12 species)—are established in Singapore. The likely pathways of introduction
and pertinent conservation issues are briey discussed. Notes on local distribution, species used for biological
control, dubious records, early records of native sh species, ornamental sh trade and aquacultural species are
also provided. An addendum is included for four more species.
Key words. Alien species, introduction pathway, aquarium/ornamental trade, aquaculture, reservoirs, urban
waterways, forest streams, freshwater swamp, Southeast Asia
RAFFLES BULLETIN OF ZOOLOGY 68: 150–195
Date of publication: 16 April 2020
DOI: 10.26107/RBZ-2020-0016
http://zoobank.org/urn:lsid:zoobank.org:pub:8AD77FDB-B04C-4E38-AA60-FE781A01273A
© National University of Singapore
ISSN 2345-7600 (electronic) | ISSN 0217-2445 (print)
Accepted by: Zeehan Jaafar
1Lee Kong Chian Natural History Museum, National University of Singapore, 2
Conservatory Drive, Singapore 117377; Email: heokhui@nus.edu.sg (*corresponding
author)
2Freshwater and Invasion Biology Laboratory, Department of Biological Sciences,
National University of Singapore, 14 Science Drive 4, Singapore 117543
3Present address: School of Biological Sciences, The University of Hong Kong,
Hong Kong SAR, China
INTRODUCTION
The rst comprehensive listing of the freshwater sh fauna
of Singapore was by Alfred (1966a). Prior to that, only
sporadic publications covering specic taxonomic groups
were published. Alfred’s (1966a) study encompassed all
the native freshwater shes recorded from Singapore at that
juncture, and also included non-native species. The next
comprehensive coverage was by Lim & Ng (1990), which
also included non-native species. More recently, updated
fauna lists were available from Baker & Lim (2008, 2012).
In one of the rst dedicated reports of non-native sh species
in Singapore, Ng HH & Tan HH (2010) documented 54
species from 14 reservoirs (three more reservoirs have since
become operational, making a present total of 17). Ng HH &
Tan HH’s (2010) exercise did not, however, include records
from other types of freshwater habitats in Singapore.
This study presents a consolidated and updated checklist of
non-native freshwater shes from all inland aquatic habitats
in Singapore, from lentic water bodies (e.g., reservoirs,
storm-water retention ponds, landscape park ponds) to lotic
waterways (e.g., urban canals, rural streams, and to a smaller
extent natural forest streams) (Yeo & Lim, 2011). It also
draws data from more than 150 published references as well
as unpublished data and technical reports (as of 31 December
2017) from an ongoing national water agency-sanctioned
study of the aquatic biodiversity and ecology in Singapore’s
reservoirs supported by PUB Singapore’s National Water
Agency. The references cover primary data from literature
and other sources, as well as secondary data (collected or
observed by the authors in this and other related studies).
MATERIAL AND METHODS
Fish specimens were obtained using a range of methods,
such as cast/throw nets (4 to 6 m diameter, 2 cm mesh size),
long line (multiple baited hooks on line), hook-and-line
(angling), push nets (60 × 40 cm frame, 2 mm mesh size),
scoop nets (1–2 mm mesh size), gill nets (4–6 cm mesh
size), sh traps (baited and un-baited), and electroshing.
Fish species were identied using various references and
keys including Kottelat et al. (1993), Larson & Lim (2005),
Baker & Lim (2012), Kottelat (2013), Froese (2018), and
Fricke et al. (2018). The recent reclassication of the order
Cypriniformes by Tan & Armbruster (2018) was not applied
to this checklist as the proposed phylogeny is preliminary,
and would only confuse readers and conservation managers
if introduced presently. Records obtained from internet
resources are included if location is mentioned and species
can be discerned or identied properly.
Fish specimens obtained were euthanised according to the
National University of Singapore’s Institutional Animal Care
and Use Committee (IACUC) guidelines. These specimens
were then xed in 10% formalin solution, leached in water
and subsequently stored in 75% ethanol.
Taxonomy & Systematics

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RAFFLES BULLETIN OF ZOOLOGY 2020
For natural distribution ranges of the shes listed in this
paper (see Fig. 122), they are grouped into the following
categories: North America (NA), Central America (CA),
South America (SA), Africa (AF), South Asia (SAs), East
Asia (EAs), Southeast Asia (SEA), Australasia (AU).
The notation “SPC” in square parantheses below the sh
name indicates that voucher specimens had been deposited
in the Zoological Reference Collection of the Lee Kong
Chian Natural History Museum, National University of
Singapore; and/or the Natural History Museum, London,
United Kingdom.
RESULTS
A total of 123 non-native sh species from 28 families and
nine orders are recorded, covering a time period of 168
years from 1849 to 2017. Of the 123 species, seven taxa
are purported hybrids or varieties. Most of the records are
relatively recent, from 1990 onwards (100 of 123 species,
81.3%). Almost all of the earlier records (pre-1990)
pertain to imported species of aquacultural importance
(see Alfred, 1966a), or incidentals likely to have originated
from aquacultural imports, with a small number possibly
introduced for biological/vector control. Almost all of
the recent records appear to be discards or escapees from
the ornamental sh/pet trade, purposeful introduction for
recreational angling (Yeo & Chia, 2010), or unknown sources
(Yeo et al., 2010).
The majority (84 species, 68.3%) of non-native sh species
are from four families: Cyprinidae (37 species, 30.1%),
Cichlidae (30 species, 24.4%), Osphronemidae (9 species,
7.3%), and Poeciliidae (8 species, 6.5%); whilst the remaining
39 species are distributed among 24 other families. Of the 123
species, 42 species (~34%) from 15 families have established
breeding populations in Singapore, with the dominant family
being the Cichlidae (12 species) followed by Cyprinidae (nine
species). See Table 1 for an overall listing of the non-native
sh species recorded from Singapore’s inland waterways.
Colour illustrations of 119 recorded species are also provided
before each species account; all were photographed by the
rst author unless stated otherwise.
COMPILATION OF NON-NATIVE FRESHWATER
FISH SPECIES
Order Rajiformes
Family Potamotrygonidae
Potamotrygon motoro (Müller & Henle)
(Fig. 1, SA[SPC])
References. Baker & Lim, 2008, 2012; Ng et al., 2009;
Chua G, 2010; Ng et al., 2009; Ng HH & Tan HH, 2010;
Chua, 2015; Lai, 2015; Tan & Zeng, 2015; Ho et al., 2016;
Tan & Tan, 2018.
Distribution. Upper Seletar Reservoir (Ng et al., 2009; Chua
G, 2010; Ng HH & Tan HH, 2010; Ho et al., 2016); Lower
Peirce Reservoir (Lai, 2015; Ho et al., 2016); Banir stream
[direct link to Lower Seletar Reservoir] (Tan & Zeng, 2015);
Lower Seletar Reservoir (Ho et al., 2016).
Remarks. Recent observations indicate a breeding population
at Lower Peirce Reservoir, where up to six juveniles were
sighted at night in shallow water adjacent to a boardwalk
(Zhou H, pers. obs., August 2016). This stingray’s diet
includes shes and aquatic macroinvertebrates, including
freshwater gastropods, crustaceans, and insect larvae (Tan
& Tan, 2018; Kwik et al., in prep.).
Order Lepisosteiformes
Family Lepisosteidae
Atractosteus spatula (Lacepède)
(Fig. 2, NA[SPC])
Fig. 1. Potamotrygon motoro, female 163 mm disc width, Lower
Peirce Reservoir.
Fig. 2. Atractosteus spatula, ca. 1,200 mm SL, Bedok Reservoir.
References. Ng HH & Tan HH, 2010; Kwik et al., 2013;
Yeo, 2013.
Distribution. Bedok Reservoir (Ng HH & Tan HH, 2010);
Pandan Reservoir (Ng HH & Tan HH, 2010); Yishun Pond
(Kwik et al., 2013); Bishan Park (Yeo, 2013); Marina
Reservoir (Anonymous, pers. comm., 2014).
Lepisosteus oculatus Winchell
(Fig. 3, NA)
Fig. 3. Lepisosteus oculatus, 119.9 mm SL, trade material.
References. Lim, 2013a; Yeo, 2013.
Distribution. Singapore Botanic Gardens (Lim, 2013a);
Bishan Park (Yeo, 2013).

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Tan et al.: Non-native freshwater shes of Singapore
Table 1a. Non-native inland shes of Singapore. A total of 123 sh species are recorded with 43 species having established breeding
populations in Singapore.
Order and Family Species Region of Origin Remarks
ORDER RAJIFORMES
Family Potamotrygonidae Potamotrygon motoro South America established
ORDER LEPISOSTEIFORMES
Family Lepisosteidae Atractosteus spatula North America
Lepisosteus oculatus North America
ORDER OSTEOGLOSSIFORMES
Family Arapaimidae Arapaima gigas South America
Family Notopteridae Chitala ornata Southeast Asia
Notopterus notopterus Southeast Asia established
Family Osteoglossidae Osteoglossum bicirrhosum South America
Scleropages formosus Southeast Asia established
Scleropages jardini Australasia
ORDER CYPRINIFORMES
Family Cyprinidae Amblypharyngodon chulabhornae Southeast Asia established
Barbodes rhombeus Southeast Asia established
Barbodes semifasciolatus East Asia extirpated
Barbonymus altus Southeast Asia
Barbonymus gonionotus Southeast Asia
Barbonymus schwanefeldii Southeast Asia established
Carassius auratus East Asia
Cirrhinus molitorella Southeast Asia
Crossocheilus oblongus Southeast Asia
Ctenopharyngodon idella East Asia
Cyprinus carpio East Asia
Danio albolineatus Southeast Asia established
Danio albolineatus × D. rerio hybrid –
Danio rerio South Asia
Danio rerio ‘frankei’ –
Dawkinsia lamentosa South Asia
Esomus metallicus Southeast Asia
Hampala macrolepidota Southeast Asia established
Hypophthalmichthys molitrix East Asia
Hypophthalmichthys nobilis East Asia
Labeo rohita South Asia
Leptobarbus rubripinna Southeast Asia
Metzia lineata East Asia extirpated
Morulius chrysophekadion Southeast Asia
Mylopharyngodon piceus East Asia
Neolissochilus sp. Southeast Asia
Osteochilus vittatus Southeast Asia established
Pethia conchonius South Asia
Puntigrus partipentazona Southeast Asia established
Puntigrus tetrazona Southeast Asia established
Puntius sophore South Asia
Rasbora cf. notura Southeast Asia
Rasbora trilineata Southeast Asia established
Toxabramis houdemeri East Asia
Tor tambra Southeast Asia
Tor tambroides Southeast Asia
Trigonopoma gracile Southeast Asia

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Order and Family Species Region of Origin Remarks
Family Botiidae Chromobotia macracanthus Southeast Asia
Family Gyrinocheilidae Gyrinocheilus aymonieri Southeast Asia
ORDER CHARACIFORMES
Family Characidae Hemigrammus rodwayi South America established
Paracheirodon innesi South America
Thayeria boehlkei South America
Family Serrasalmidae Piaractus brachypomus South America
ORDER SILURIFORMES
Family Bagridae Hemibagrus capitulum Southeast Asia
Hemibagrus guttatus East Asia extirpated
Mystus castaneus Southeast Asia
Mystus wolfi Southeast Asia
Family Clariidae Clarias gariepinus Africa established
Clarias macrocephalus Southeast Asia
Family Callichthyidae Corydoras aeneus South America
Family Loricariidae Pterygoplichthys disjunctivus South America established
Pterygoplichthys joselimaianus South America established
Pterygoplichthys pardalis South America established
Family Mochokidae Synodontis eupterus Africa
Family Pangasiidae Pangasius sanitwongsei Southeast Asia
Pangasionodon hypophthalmus Southeast Asia
Family Pimelodidae Phractocephalus hemioliopterus South America
ORDER CYPRINODONTIFORMES
Family Aplocheilidae Aplocheilus lineatus South Asia established
Family Poeciliidae Gambusia afnis Central America established
Poecilia latipinna Central America
Poecilia reticulata Central America established
Poecilia sphenops Central America established
Poecilia velifera Central America
Xiphophorus hellerii Central America
Xiphophorus maculatus Central America
Xiphophorus variatus Central America
ORDER SYNBRANCHIFORMES
Family Mastacembelidae Macrognathus siamensis Southeast Asia
Macrognathus zebrinus Southeast Asia established
ORDER PERCIFORMES
Family Ambassidae Parambassis siamensis Southeast Asia established
Family Datnioididae Datnioides microlepis Southeast Asia
Family Monodactylidae Monodactylus sebae Africa

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Tan et al.: Non-native freshwater shes of Singapore
Order and Family Species Region of Origin Remarks
Family Cichlidae Acarichthys heckelli South America established
Amphilophus citrinellus Central America established
Amphilophus trimaculatus Central America
Apistogramma borelli South America established
Astronotus ocellatus South America
Cichla kelberi South America
Cichla orinocensis South America established
Cichla piquiti South America
Cichla temensis South America established
Coptodon zillii Africa
Etroplus suratensis South Asia established
Geophagus altifrons South America established
Herichthys carpintis Central America
Heros severus South America
Heterotilapia buttikoferi Africa established
Mayaheros urophthalmus Central America established
Mayaheros urophthalmus ×
Parachromis managuensis hybrid
–
Maylandia estherae Africa
Oreochromis aureus Africa
Oreochromis mossambicus Africa established
Oreochromis niloticus Africa established
Oreochromis mossambicus × O.
niloticus hybrid
– established
Parachromis managuensis Central America established
Pterophyllum scalare South America
Satanoperca jurupari South America
Thorichthys meeki Central America
Vieja melanura Central America established
Luohan/Flowerhorn cichlid hybrid –
Blood/Red Parrot hybrid –
Red Tilapia –
Family Gobinellidae Brachygobius sabanus Southeast Asia established
Rhinogobius similis East Asia established
Family Helostomatidae Helostoma temminckii Southeast Asia
Family Osphronemidae Betta splendens Southeast Asia established
Macropodus opercularis East Asia
Osphronemus goramy Southeast Asia established
Osphronemus laticlavius Southeast Asia
Sphaerichthys osphromenoides Southeast Asia
Trichogaster lalius South Asia
Trichopodus microlepis Southeast Asia
Trichopodus pectoralis Southeast Asia established
Trichopsis schalleri Southeast Asia established
Family Channidae Channa micropeltes Southeast Asia established

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Table 1b. Fish species listed in addendum.
Order and Family Species Region of Origin Remarks
ORDER SILURIFORMES
Family Bagridae Horabagrus brachysoma South Asia
Family Claroteidae Chrysichthys auratus Africa
ORDER CHARACIFORMES
Family Anostomidae Leporinus fasciatus South America
ORDER PERCIFORMES
Family Cichlidae Veija × Mayaheros hybrid –
Order Osteoglossiformes
Family Arapaimidae
Arapaima gigas (Schinz)
(Fig. 4, SA)
Fig. 4. Arapaima gigas, 181.2 mm SL, trade material.
References. shing_vimal, 2017.
Distribution. Marina Reservoir (shing_vimal, 2017).
Remarks. Sightings of Arapaima gigas (Arapaimidae) have
been listed in forums (e.g., http://forum.shingkaki.com/
discussion/229269/arapaima-in-singapore); however, these
were never substantiated with photographs or locations
until recently from Marina Reservoir near the steps along
Boat Quay (shing_vimal, 2017). This species is available
in the ornamental sh trade, although in limited quantity.
This species grows to be one of the largest freshwater sh
(Luna, 2020), and it is unsurprising that larger individuals
are discarded or released into articial water bodies when
they out-grow their tanks.
Family Notopteridae
Chitala ornata (Gray)
(Fig. 5, SEA[SPC])
Fig. 5. Chitala ornata, 650 mm SL, Pandan Reservoir.
References. Ng & Lim, 1997b; Baker & Lim, 2008, 2012;
Ng HH & Tan HH, 2010; Tan HTW et al., 2010.
Distribution. Jurong Lake (Ng HH & Tan HH, 2010); Kranji
Reservoir (Ng HH & Tan HH, 2010); Upper Seletar Reservoir
(Ng HH & Tan HH, 2010); Lower Seletar Reservoir (Ng &
Lim, 1997b; Ng HH & Tan HH, 2010); Pandan Reservoir
(Ng HH & Tan HH, 2010); Upper Peirce Reservoir (Ng
HH & Tan HH, 2010); MacRitchie Reservoir (unpublished
data); Punggol Reservoir (unpublished data).
Notopterus notopterus (Pallas)
(Fig. 6, SEA, SAs[SPC])
Fig. 6. Notopterus notopterus, ca. 250 mm SL, Tengeh Reservoir.
References. Baker & Lim, 2008, 2012; Ng HH & Tan HH,
2010; Baker, 2013c; Tan et al., 2016.
Distribution. Jurong Lake (Ng HH & Tan HH, 2010); Murai
Reservoir (Ng HH & Tan HH, 2010); Lower Peirce Reservoir
(Ng HH & Tan HH, 2010); Lower Seletar Reservoir (Ng
HH & Tan HH, 2010); Poyan Reservoir (Ng HH & Tan
HH, 2010); Tengeh Reservoir (Ng HH & Tan HH, 2010);
Kranji Marsh (Baker, 2013c; Tan et al., 2016); Upper Seletar
Reservoir spill gate (Tan HH, pers. obs., 2014); Pandan
reservoir (unpublished data).
Family Osteoglossidae
Osteoglossum bicirrhosum (Cuvier)
(Fig. 7, SA)
References. Anonymous, 1988a; Ng & Lim, 1997b; Ng
HH & Tan HH, 2010.

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Tan et al.: Non-native freshwater shes of Singapore
Fig. 7. Osteoglossum bicirrhosum, 215 mm SL, trade material.
Distribution. Singapore Botanic Gardens (Anonymous,
1988a); Lower Peirce Reservoir (Ng HH & Tan HH, 2010);
Sungei Buloh Wetland Reserve (Tan HH, pers. obs., mid-
2000s).
Scleropages formosus (Müller & Schlegel)
(Fig. 8, SEA[SPC])
Catchment Nature Reserve stream (Tan, 2014c; Ho et al.,
2016); Banir (Tan, 2015c); Jurong Lake (unpublished data);
Lower Peirce reservoir (unpublished data); Bedok reservoir
(unpublished data).
Remarks. This species is protected under the Convention
on International Trade in Endangered Species of Wild Fauna
and Flora (CITES) Appendix I and breeding populations
are established in the Central Catchment Nature Reserve
reservoirs in Singapore (Ng HH & Tan HH, 2010). One of the
documented sources is from then-PPD (Primary Productions
Department; presently divided into National Parks Board
and Singapore Food Agency) (see Ng HH & Tan HH, 2010,
for a more detailed account). The Singapore variants are
of questionable sources, and probably the consequence of
articial crosses between recognised colour variants (Andy
Yap, pers. comm.).
Scleropages jardini (Saville-Kent)
(Fig. 9, AU)
Fig. 8. Scleropages formosus, 32.0 mm SL juvenile (top), 490 mm
SL adult (bottom), Upper Peirce Reservoir.
Fig. 9. Scleropages jardini, ca. 350 mm SL, trade material.
References. None. This is a new record for Singapore.
Distribution. Singapore Botanic Gardens, Swan Lake (Lim
KKP, pers. obs., October 2010).
Order Cypriniformes
Family Cyprinidae
Amblypharyngodon chulabhornae Vidthayanon &
Kottelat
(Fig. 10, SEA[SPC])
References. Green, 1928; Anonymous, 1988a, 1989a; Lim
& Ng, 1990; Ng et al., 1993; Ng & Lim, 1997a, 1997b;
National Parks Board, 2003; Baker & Lim, 2008, 2012; Ng
HH & Tan HH, 2010; Baker, 2013b; Kwik et al., 2013; Baker,
2014; Tan, 2014b, 2014c, 2015c, 2016a; Ho et al., 2016.
Distribution. Lim & Ng (1990) listed this species from the
Central Catchment reservoirs; Lower Peirce Reservoir (Ng
& Lim, 1997a, 1997b; Baker, 2013b); Singapore Botanic
Gardens (Anonymous, 1988a); East Coast Park (Anonymous,
1988a); Woodlands Town Garden (Anonymous, 1989a);
MacRitchie Reservoir (Ng & Lim, 1997a, 1997b; Ng HH
& Tan HH, 2010; Tan, 2016a); Bedok Reservoir (Ng HH
& Tan HH, 2010); Upper Seletar Reservoir (Ng HH & Tan
HH, 2010); Lower Seletar Reservoir (Ng HH & Tan HH,
2010); Pandan Reservoir (Ng HH & Tan HH, 2010); Upper
Peirce Reservoir (Ng HH & Tan HH, 2010); Sungei Buloh
Wetland Reserve (National Parks Board, 2003; Baker, 2014);
Pangsua Pond (Kwik et al., 2013; Tan, 2014b); Central
Fig. 10. Amblypharyngodon chulabhornae, ca. 25 mm SL, Kranji
Marsh.
References. Ng et al., 1993; Ng & Lim, 1997a, 1997b;
Baker & Lim, 2012; Lim & Tan, 2012; Lim et al., 2013;
Tan et al., 2016.
Distribution. Choa Chu Kang pond (Ng & Lim, 1997a;
Lim & Tan, 2012); Kranji Marsh (Lim & Tan, 2012; Lim
et al., 2013; Tan et al., 2016).

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Fig. 14. Barbonymus gonionotus, 140.0 mm SL, trade material
(specimen with aberrant body scale pattern).
Fig. 11. Barbodes rhombeus, 57.5 mm SL, Bishan Park.
Fig. 12. Barbodes semifasciolatus, 42.2 mm SL, wild type from
Vietnam (top); ca. 30 mm SL, xanthic trade material (bottom).
Puntius semifasciolatus, as early as 1900s); Munro, 1990
(as Puntius semifasciolatus); Ng et al., 1993 (as Puntius
semifasciolatus); Lever, 1996 (as Barbus semifasciolatus);
Ng & Lim, 1997a (as Puntius semifasciolatus); Ng & Lim,
1997b (as Puntius semifasciolatus); Baker & Lim, 2008,
2012 (as Puntius semifasciolatus); Tan HH et al., 2010 (as
Puntius semifasciolatus).
Distribution. Probably extirpated in Singapore as there have
been no recent records; previously recorded from Serangoon
area (de Beaufort, 1933; Tweedie, 1936; Alfred, 1966a;
Munro, 1990); Bukit Timah area (Alfred, 1966a; Ng & Lim,
1997a); MacRitchie Reservoir (Alfred, 1966a; Anonymous,
1988a); Kallang River (Alfred, 1966a; Tan HH et al., 2010);
Sungei Seletar (Alfred, 1966a); Lower Peirce Reservoir (Ng
& Lim, 1997a, 1997b); Central Catchment Nature Reserve
— Sime Road forest (Lim, 1989; Ng & Lim, 1997a, 1997b).
Remarks. This species was an accidental introduction with
imported sh fry from China (Johnson, 1964), possibly as
early as 1912 (Alfred, 1966a).
Barbonymus altus (Günther)
(Fig. 13, SEA[SPC])
Fig. 13. Barbonymus altus, 110.8 mm SL, Pangsua Pond.
References. Subaraj et al., 1995 (as Puntius binotatus); Ng
& Lim, 1997a (as Puntius binotatus); Ng & Lim, 1997b (as
Puntius binotatus); Tan et al., 2013 (as Systomus rhombeus);
Tan & Low, 2014; Kwik & Yeo, 2015 (as Puntius binotatus);
Ho et al., 2016.
Distribution. Mandai area (Subaraj et al., 1995; Tan &
Low, 2014); Andrew Road drain (Ng & Lim, 1997a); Bukit
Brown (Tan et al., 2013); Venus Drive (Tan HH, pers. obs.,
2010); Pandan river (Tan HH, pers. obs., March 2014);
outskirts of Nee Soon Swamp Forest (Ho et al., 2016);
Tengah area (Cai YX, pers. obs., 11 January 2017); Bishan
Park (unpublished data).
Barbodes semifasciolatus (Günther) [Extirpated]
(Fig. 12, SEA, EAs[SPC])
References. de Beaufort, 1933 (as Puntius sachsi);
Tweedie, 1936 (as Puntius sachsi); Fowler, 1938 (as
Barbus sachsii); Johnson, 1964 (as Puntius semifasciolatus);
Alfred, 1966a (as Puntius semifasciolatus, as early as
1912); Johnson, 1973a (as Puntius semifasciolatus);
Anonymous, 1988a (as Capoeta semifasciolatus); Lim,
1989 (as Puntius semifasciolatus); Lim & Ng, 1990 (as
References. Ng & Lim, 1997a (as Barbodes altus); Ng HH
& Tan HH, 2010.
Distribution. Jurong Lake (Ng HH & Tan HH, 2010).
Barbonymus gonionotus (Bleeker)
(Fig. 14, SEA)
Remarks. Both juveniles and adults have been obtained
from Kranji Marsh, suggesting an established breeding
population there.
Barbodes rhombeus (Kottelat)
(Fig. 11, SEA[SPC])
References. Alfred, 1966a (as Puntius gonionotus);
Anonymous, 1989a (as Puntius gonionotus); Lim & Ng,
1990 (as Barbodes gonionotus); Ng et al., 1993 (as Barbodes

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Tan et al.: Non-native freshwater shes of Singapore
gonionotus); Ng & Lim, 1997a (as Barbodes gonionotus);
Ng & Lim, 1997b (as Barbodes gonionotus).
Distribution. Woodlands Town Garden (Anonymous,
1989a).
Remarks. This species was introduced as a food sh during
the Japanese Occupation of Singapore (1942–1945) (Alfred,
1966a; Mohsin & Ambak, 1983; Lim & Ng, 1990; Ng &
Lim, 1997a), but failed to gain popularity. There have not
been any recent records (i.e., post-1990).
Barbonymus schwanefeldii (Bleeker)
(Fig. 15, SEA[SPC])
Fig. 15. Barbonymus schwanefeldii, 95.8 mm SL, trade material.
References. Alfred, 1966a (as Puntius schwanenfeldii, rst
recorded in 1937); Baker & Lim, 2008, 2012; Ng HH &
Tan HH, 2010.
Distribution. Jurong area (Alfred, 1966a); MacRitchie
Reservoir (Ng HH & Tan HH, 2010); Marina Reservoir
(unpublished data); Jurong Lake (unpublished data).
Remarks. Recorded as extinct in Singapore (Alfred, 1966a);
recent records probably represent non-native introductions
(Ng HH & Tan HH, 2010).
Carassius auratus (Linnaeus)
(Fig. 16, EAs[SPC])
Fig. 16. Carassius auratus, ca. 35 mm SL, Yishun Pond.
Distribution. Bedok Reservoir (Ng HH & Tan HH, 2010);
Kallang River (now part of Marina Reservoir; Tan HH et
al., 2010).
Remarks. This species was most probably introduced from
Java for aquaculture (Mohsin & Ambak, 1983). Present
records are mainly from escapees or releases, as this species
is a very popular ornamental sh and frequently used as
feeder sh (Lim & Ng, 1990; Ng & Lim, 1997a).
Cirrhinus molitorella (Valenciennes, in Cuvier &
Valenciennes)
(Fig. 17, SEA)
Fig. 17. Cirrhinus molitorella, 99.5 mm SL, trade material.
References. Herre & Myers, 1937; Le Mare, 1949 (as
Cirrhina molitorella); Tham, 1973 (as Cirrhina molitorella);
Munro, 1990; Ng & Lim, 1997a, 1997b.
Distribution. Not known.
Remarks. Herre & Myers’ (1937) record was an escapee
from cultivation, and this may not have established in
Singapore. Tham (1973) commented that this is a rarely
used sh for aquaculture, with fry imported from China.
Presently, the juveniles of this sh are used quite extensively
as feed for predatory sh in the ornamental sh trade (Tan
HH, pers. obs.).
Crossocheilus oblongus Kuhl & van Hasselt, in van
Hasselt
(Fig. 18, SEA[SPC])
Fig. 18. Crossocheilus oblongus, 112.7 mm SL, Punggol Reservoir.
References. None. This is a new record for Singapore.
Distribution. Punggol Reservoir (unpublished data).
Remarks. This is probably a discarded pet, as this species
is actively promoted in the ornamental sh trade as an algae
eater and for algae control (Tan HH, pers. obs.).
References. Lim & Ng, 1990; Ng et al., 1993; Ng & Lim,
1997a, 1997b; Ng HH & Tan HH, 2010; Tan HH et al., 2010.

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Ctenopharyngodon idella (Valenciennes, in Cuvier &
Valenciennes)
(Fig. 19, EAs)
Fig. 19. Ctenopharyngodon idella, 83.7 mm SL, trade material.
References. Herre & Myers, 1937; Le Mare, 1949
(as Ctenopharyngodon idellus); Tham, 1973 (as
Ctenopharyngodon idellus); Mohsin & Ambak, 1983; Yang,
1984; Lim & Ng, 1990; Munro, 1990; Public Utilities Board,
1991; Ng et al., 1993; Ng & Lim, 1997a, 1997b.
Distribution. Upper Seletar Reservoir (previously known
as Seletar Reservoir; Yang, 1984).
Remarks. Herre & Myers’ (1937) record came from an
experimental sh pond, and this may not have established
in Singapore. Tham (1973) commented that it was the most
popular sh cultured in sh ponds and the fry were imported
from China. This species was reared in experimental oating
cages in reservoirs as early as 1972, and subsequently used
primarily for control of submerged macrophytes (Yang,
1984).
Cyprinus carpio Linnaeus
(Fig. 20, EAs[SPC])
Remarks. Herre & Myers’ (1937) record came from an
experimental fish pond, and this species may not have
established in Singapore then and presently (Lim & Ng,
1990). Tham (1973) commented that this was a popular
cultured sh, which was bred locally. Many of the recent
records refer to the colourful variants, indicative of discarded
pets or mercy/religious release.
Danio albolineatus (Blyth)
(Fig. 21, SEA[SPC])
Fig. 20. Cyprinus carpio, 220 mm SL, Pangsua Pond.
Fig. 21. Danio albolineatus, 26.0 mm SL, Mandai area.
Fig. 22. Danio albolineatus × D. rerio hybrid, 27.2 mm SL,
Mount Pleasant.
Fig. 23. Danio rerio, 35.6 mm SL, Mount Pleasant.
References. Tan et al., 2013.
Distribution. Bukit Brown (Tan et al., 2013).
References. Lim & Tan, 2011; Baker & Lim, 2012; Tan et
al., 2013; Tan & Low, 2014.
Distribution. Mandai area (Lim & Tan, 2011; Tan & Low,
2014); Bukit Brown (Tan et al., 2013); Bukit Timah area
(Tan HH, pers. obs., February 2015); Tengah area (Cai YX,
pers. obs., 11 January 2017).
Danio albolineatus × D. rerio
(Fig. 22 [SPC])
References. Herre & Myers, 1937; Le Mare, 1949; Tham,
1973; Mohsin & Ambak, 1983; Yang, 1984; Lim & Ng,
1990; Munro, 1990; Ng et al., 1993; Ng & Lim, 1997a,
1997b; National Parks Board, 2003; Ng HH & Tan HH,
2010; Tan HTW et al., 2010; Yeo & Chia, 2010; Davison
et al., 2012 (as carp); Kwik et al., 2013; Tan, 2016b; Tan
HH et al., 2017b.
Distribution. Upper Seletar Reservoir (previously known
as Seletar Reservoir; Yang, 1984); Sungei Buloh Wetland
Reserve (National Parks Board, 2003); Catchment Pond (Tan
HH, pers. obs., 2000s); Bedok Reservoir (Ng HH & Tan
HH, 2010); Pandan Reservoir (Ng HH & Tan HH, 2010);
Kent Ridge Park pond (Tan HTW et al., 2010; Tan, 2016b);
Pangsua pond (Kwik et al., 2013); Singapore Botanic Gardens
(Tan HH, pers. obs., 2015); Singapore Quarry (Tan HH et
al., 2017b); Bishan Park (unpublished data).
References. Tan et al., 2013.
Distribution. Bukit Brown (Tan et al., 2013).
Remarks. The presence of this hybrid was reported by Tan
et al. (2013).
Danio rerio (Hamilton)
(Fig. 23, SEA[SPC])

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Tan et al.: Non-native freshwater shes of Singapore
Fig. 24. Danio rerio ‘frankei’, 37.6 mm SL, Mount Pleasant.
References. Tan et al., 2013.
Distribution. Bukit Brown (Tan et al., 2013).
Dawkinsia lamentosa (Valenciennes, in Cuvier &
Valenciennes)
(Fig. 25, SAs[SPC])
Fig. 25. Dawkinsia lamentosa, 77.9 mm SL, Woodlands Pond
(Kwik JTB).
Fig. 26. Esomus metallicus, ca. 40 mm SL, trade material (note
the diagnostic character of long barbels).
Remarks. This species is occasionally sold as live feed in
the ornamental sh trade (Tan HH, pers. obs.).
Hampala macrolepidota Kuhl & van Hasselt, in van
Hasselt
(Fig. 27, SEA[SPC])
Fig. 27. Hampala macrolepidota, ca. 250 mm SL female, Upper
Peirce Reservoir.
References. Herre & Myers, 1937; Alfred, 1966a; Ng &
Lim, 1997b; Baker & Lim, 2008, 2012; Ng HH & Tan HH,
2010; Tan HTW et al., 2010; Yeo & Lim, 2010.
Distribution. Upper Peirce Reservoir (Ng & Lim, 1997b;
Ng HH & Tan HH, 2010); Lower Peirce Reservoir (Ng
HH & Tan HH, 2010); Singapore Botanic Gardens (Yeo
& Lim, 2010).
Remarks. Alfred (1966a) considered this species extinct
in Singapore. Recent populations are probably intentional
releases or from Johor, Peninsular Malaysia, via raw water
transfer (Ng & Lim, 1997b; Ng HH & Tan HH, 2010).
Hypophthalmichthys molitrix (Valenciennes, in Cuvier
& Valenciennes)
(EAs)
References. Le Mare, 1949; Tham, 1973; Mohsin & Ambak,
1983; Yang, 1984; Munro, 1990; Ng et al., 1993; Ng &
Lim, 1997b.
Distribution. Upper Seletar Reservoir (previously known
as Seletar Reservoir; Yang, 1984).
Remarks. This species is cultivated in ponds for food,
and the fry stock is imported from China (Le Mare, 1949;
Tham, 1973; Munro, 1990; Ng et al., 1993). This species
was reared in experimental oating cages in reservoirs as
early as 1972 (Yang, 1984).
Hypophthalmichthys nobilis (Richardson)
(Fig. 28, EAs)
References. Le Mare, 1949 (as Aristichthys nobilis); Tham,
1973 (as Aristichthys nobilis); Mohsin & Ambak, 1983 (as
Aristichthys nobilis); Yang, 1984 (as Aristichthys nobilis);
Lim & Ng, 1990; Munro, 1990; Public Utilities Board,
1991; Ng et al., 1993; Ng & Lim, 1997a, 1997b; Ng HH
& Tan HH, 2010.
Distribution. Pandan Reservoir (Lim & Ng, 1990; Ng HH &
Tan HH, 2010); Upper Seletar Reservoir (Yang, 1984; Lim
References. Kwik, 2013; Kwik et al., 2013.
Distribution. Woodlands Town Park pond (Kwik, 2013;
Kwik et al., 2013).
Esomus metallicus Ahl
(Fig. 26, SEA)
Danio rerio ‘frankei’
(Fig. 24 [SPC])
References. Anonymous, 1988a; Lim & Ng, 1990; Munro,
1990; Ng et al., 1993; Ng & Lim, 1997a, 1997b; Baker &
Lim, 2008, 2012.
Distribution. Sungei Sembawang — Senoko (Anonymous,
1988a; Ng & Lim, 1997a); Mandai Road area (Anonymous,
1998a; Ng & Lim, 1997a); Lower Seletar Reservoir (Tan
HH, pers. obs., early 1990s).

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Fig. 28. Hypophthalmichthys nobilis, 104.3 mm SL, trade material.
& Ng, 1990; Ng HH & Tan HH, 2010); Kranji Reservoir
(Ng HH & Tan HH, 2010); Jurong Lake (Lim KKP, pers.
obs., 2011).
Remarks. This species is popular as food sh and fry were
imported from China (Le Mare, 1949; Tham, 1973). This
species was used extensively by the Public Utilities Board
(PUB) to control plankton levels in reservoirs as early as
1972 (Yang, 1984; Lim & Ng, 1990; Ng & Lim, 1997a);
however, this species is not known to breed in the tropics
and all stock for aquaculture is imported (Ng & Lim, 1997b).
Labeo rohita (Hamilton)
(Fig. 29, SAs)
Fig. 29. Labeo rohita, 240 mm SL, trade material.
References. Ng HH & Tan HH, 2010.
Distribution. MacRitchie Reservoir (Ng HH & Tan HH,
2010).
Remarks. This sh species is sometimes sold in wet markets
and supermarkets in Singapore (Tan HH, pers. obs.).
Leptobarbus rubripinna (Fowler)
(Fig. 30, SEA[SPC])
Fig. 30. Leptobarbus rubripinna, ca. 300 mm SL, Jurong Lake.
Distribution. Jurong Lake (Ng HH & Tan HH, 2010); Lower
Peirce Reservoir (Ng HH & Tan HH, 2010); Lower Seletar
Reservoir (Ng HH & Tan HH, 2010); Kent Ridge Park pond
(Tan, 2016b); Punggol Reservoir (unpublished data).
Remarks. This species is not established in Singapore as
no breeding populations are known (Lim & Ng, 1990; Ng
& Lim, 1997a).
Metzia lineata (Pellegrin) [Extirpated]
(Fig. 31. EAs[SPC])
Fig. 31. Metzia lineata, 41.9 mm SL, Vietnam.
References. Regan, 1913 (described as Rasborichthys altior);
Weber & de Beaufort, 1916 (as Rasborichthys altior);
Tweedie, 1936 (as Rasborichthys altior); Herre & Myers, 1937
(as Rasborichthys altior); Blythe, 1951 (as Rasborichthys
altior); Johnson, 1964 (as Rasborichthys altior); Alfred,
1966a (as Rasborichthys altior); Bănărescu, 1971 (as
Rasborinus lineatus); Johnson, 1973a (as Rasborichthys
altior); Lim & Ng, 1990 (as Rasborinus lineatus, as early
as late 1800s or early 1900s); Munro, 1990 (as Rasborinus
lineatus); Ng et al., 1993 (as Rasborinus lineatus takakii);
Ng & Lim, 1997a (as Rasborinus lineatus takakii); Ng &
Lim, 1997b (as Rasborinus lineatus); Tan HH et al., 2010;
Tan HTW et al., 2010 (as Rasborinus lineatus takakii);
Baker & Lim, 2008, 2012.
Distribution. Probably extirpated in Singapore as there
have been no recent records (Ng & Lim, 1997a, 1997b);
previously recorded from Singapore Botanic Gardens
(Tweedie, 1936; Alfred, 1966a); Serangoon area (Tweedie,
1936; Alfred, 1966a); Mandai Road area (Herre & Myers,
1937); MacRitchie Reservoir (Alfred, 1966a); Sungei Seletar
(Alfred, 1966a); Sungei Whampoe (Alfred, 1966a); Kallang
River (Alfred, 1966a; Tan HTW et al., 2010); Bukit Timah
campus (Alfred ER, pers. comm., 2007); Lower Peirce
Reservoir (Ng & Lim, 1997a); Upper Seletar Reservoir (Ng
& Lim, 1997a).
Remarks. Regan (1913) described this sh species on the
presumption that it was endemic to Singapore. Weber & de
Beaufort (1916) followed suit in regarding the species as
native and endemic, as did subsequent workers (Tweedie,
1936; Herre & Myers, 1937; Alfred, 1966a; Johnson,
1973a). The taxonomy was only resolved by Bănărescu in
1971, and its endemic/native status debunked. This species
is now understood to be an accidental import, along with
ngerlings of food sh species, imported from Hong Kong
and southern China (Lim & Ng, 1990).
References. Lim & Ng, 1990 (as L. hoeveni); Ng et al.,
1993 (as L. hoeveni); Ng & Lim, 1997a (as L. hoeveni); Ng
& Lim, 1997b (as L. hoeveni); Ng HH & Tan HH, 2010;
Tan, 2016b.

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Tan et al.: Non-native freshwater shes of Singapore
Morulius chrysophekadion (Bleeker)
(Fig. 32, SEA[SPC])
Remarks. This species could not be identied to species
level as the taxonomic status is complicated and remains
unresolved.
Osteochilus vittatus (Valenciennes, in Cuvier &
Valenciennes)
(Fig. 35, SEA[SPC])
Fig. 32. Morulius chrysophekadion, 375 mm SL, Punggol Reservoir.
References. None. This is a new record for Singapore.
Distribution. Punggol Reservoir (unpublished data).
Mylopharyngodon piceus (Richardson)
(Fig. 33, EAs)
Fig. 33. Mylopharyngodon piceus, ca. 500 mm SL, Tengah ponds
(Yeo DCJ).
References. Herre & Myers, 1937 (as Mylopharyngodon
aethiops).
Distribution. Not known.
Remarks. Herre & Myers’ (1937) record came from an
experimental fish pond, and this species may not have
established in Singapore. Presently, this species can be
occasionally found in ponds stocked for angling purposes
(Yeo DCJ, pers. obs.).
Neolissochilus sp.
(Fig. 34, SEA[SPC])
Fig. 34. Neolissochilus sp., ca. 250 mm SL, Pandan Reservoir.
Fig. 35. Osteochilus vittatus, ca. 150 mm SL, Sime forest.
References. Ng & Lim, 1997a (as O. hasselti); Ng & Lim,
1997b (as O. hasselti); Baker & Lim, 2008, 2012 (as O.
hasselti); Ng HH & Tan HH, 2010; Baker, 2013a; Tan, 2013c,
2013d; Kwik & Yeo, 2015; Ho et al., 2016; Li et al., 2016.
Distribution. Upper Seletar Reservoir (Ng & Lim, 1997a,
1997b); Lower Seletar Reservoir (Ng HH & Tan HH, 2010);
Upper Peirce Reservoir (Ng HH & Tan HH, 2010); Lower
Peirce Reservoir (Ng HH & Tan HH, 2010); Banir (Baker,
2013a); Central Catchment Nature Reserve — Sime forest
(Tan, 2013c, 2013d; Ho et al., 2016); Central Catchment
Nature Reserve — Nee Soon Swamp Forest (Ho et al., 2016;
Li et al., 2016); Bishan Park (unpublished data); MacRitchie
reservoir (unpublished data).
Pethia conchonius (Hamilton)
(Fig. 36, SAs[SPC])
Fig. 36. Pethia conchonius, 42.8 mm SL, trade material.
References. Anonymous, 1990; Ng et al., 1993 (as Puntius
conchonius); Ng & Lim, 1997a (as Puntius conchonius); Ng
& Lim, 1997b (as Puntius conchonius).
Distribution. Reported as an escapee in Sime Road forest
(Anonymous, 1990) in Central Catchment Nature Reserve;
this species is not an established species (Ng & Lim, 1997a).
References. Ng HH & Tan HH, 2010.
Distribution. MacRitchie Reservoir (Ng HH & Tan HH,
2010).

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Puntigrus partipentazona (Fowler)
(Fig. 37, SEA[SPC])
tetrazona); Lim et al., 1995 (as Puntius tetrazona); Ng &
Lim, 1997a, 1997b (as Puntius tetrazona); National Parks
Board, 2003 (as Systomus tetrazona); Baker & Lim, 2008,
2012 (as Systomus tetrazona); Ng HH & Tan HH, 2010 (as
Puntius tetrazona); Ho et al., 2016; Li et al., 2016; Lim et
al., 2016; Tan, 2016b.
Distribution. Herre & Myers (1937) obtained a single
specimen from Singapore and commented that it had an
aberrant barring pattern, but provided no location details;
Central Catchment Nature Reserve — Nee Soon Swamp
Forest (Anonymous, 1988a; Munro, 1990; Ng & Lim, 1997a;
Ho et al., 2016; Li et al., 2016); Bukit Timah — Catchment
Pond (Anonymous, 1988a); Upper Seletar Reservoir (Munro,
1990; Lim et al., 1995; Ng HH & Tan HH, 2010); Seletar
Reservoir Park (Munro, 1990); Seletar area (Ng et al., 1993);
Sungei Buloh Wetland Reserve (National Parks Board, 2003);
Upper Peirce Reservoir (Ng HH & Tan HH, 2010); Lower
Peirce Reservoir (Ng HH & Tan HH, 2010); Lower Seletar
Reservoir (Ng HH & Tan HH, 2010); MacRitchie Reservoir
(Ng HH & Tan HH, 2010); Murai Reservoir (Ng HH & Tan
HH, 2010); Tekong Reservoir (Ng HH & Tan HH, 2010;
Lim et al., 2016); Tengah Reservoir (Ng HH & Tan HH,
2010); Kent Ridge Park ponds (Tan, 2016b).
Remarks. Ng HH & Tan HH (2010) only recorded this
species as they listed species located from reservoirs.
This species can be distinguished from its congener (P.
partipentazona) that is also present and non-native to
Singapore, on basis of the width of black bars (vs. slimmer),
depth of body (vs. less deep), and larger overall adult size
(vs. smaller).
Puntius sophore (Hamilton)
(Fig. 39, SAs[SPC])
References. Lim & Tan, 2011; Baker & Lim, 2012; Lim et
Fig. 37. Puntigrus partipentazona, 31.0 mm SL, Banir stream.
References. Boeseman, 1957 (as Puntius partipentazona);
Anonymous, 1990 (as Puntius partipentazona); Lim & Ng,
1990 (as Puntius partipentazona); Munro, 1990 (as Puntius
partipentazona); Ng, 1991 (as Puntius partipentazona); Ng et
al., 1993 (as Puntius partipentazona); Lim, 1995 (as Puntius
partipentazona); Lim et al., 1995 (as Puntius partipentazona);
Subaraj et al., 1995 (as Puntius partipentazona); Ng & Lim,
1997a (as Puntius partipentazona); Ng & Lim, 1997b (as
Puntius partipentazona); Baker & Lim, 2008, 2012 (as
Systomus partipentazona).
Distribution. MacRitchie Reservoir (Lim & Ng, 1990; Ng
& Lim, 1997b); Central Catchment Nature Reserve — Nee
Soon Swamp Forest (Munro, 1990); MacRitchie Reservoir
(Subaraj et al., 1995); Sime Road forest (Ng & Lim, 1997a,
1997b); Bukit Timah Nature Reserve — eastern boundary,
Catchment Pond (Lim, 1995; Ng & Lim, 1997b); Upper
Seletar Reservoir (Munro, 1990); Lower Peirce Reservoir
(Anonymous, 1990; Lim et al., 1995; Ng & Lim, 1997a,
1997b); Sungei Seletar (Banir conuence; Tan HH, pers.
obs?, March 2017).
Remarks. This species is occasionally sold as live feed in
the ornamental sh trade (Tan HH, pers. obs.).
Puntigrus tetrazona (Bleeker)
(Fig. 38, SEA[SPC])
Fig. 38. Puntigrus tetrazona, 46.6 mm SL, Bukit Batok Quarry.
Fig. 39. Puntius sophore, 60.0 mm SL, Sungei Buloh.
al., 2013; Kwik & Yeo, 2015; Tan et al., 2016.
Distribution. Sungei Seletar (Lim & Tan, 2011); Kranji
Marsh (Lim & Tan, 2011; Tan et al., 2016); Sungei Buloh
Wetland Reserve (Lim & Tan, 2011; Lim et al., 2013).
Rasbora borapetensis Smith
(Fig. 40, SEA[SPC])
References. Anonymous, 1988a, 1988b, 1990; Lim & Ng,
1990; Munro, 1990; Ng, 1991; Lim & Ng, 1992; Ng et al.,
1993; Subaraj et al., 1995; Ng & Lim, 1997a, 1997b; Baker
& Lim, 2008, 2012; Ng HH & Tan HH, 2010; Yeo & Lim,
2010; Lim et al., 2013; Ng HH & Tan HH, 2013; Tan HH
References. Herre & Myers, 1937 (as Puntius tetrazona);
Anonymous, 1988a (as Capoeta tetrazona); Lim & Ng,
1990 (as Puntius tetrazona); Munro, 1990 (as Barbus
tetrazona and Puntius tetrazona); Ng et al., 1993 (as Puntius

164
Tan et al.: Non-native freshwater shes of Singapore
Fig. 40. Rasbora borapetensis, 25.0 mm SL, Kranji Marsh.
et al., 2015; Li et al., 2016; Tan, 2016a, 2016b; Ho et al.,
2016; Li et al., 2016; Tan et al., 2016.
Distribution. Singapore Botanic Gardens (Anonymous,
1988a; Lim & Ng, 1990; Munro, 1990; Ng & Lim, 1997a;
Yeo & Lim, 2010); Bukit Timah — Institute of Education
pond (Anonymous, 1988b); MacRitchie Reservoir (Subaraj
et al., 1995); Sungei Sembawang — Senoko (Ng & Lim,
1997a); Choa Chu Kang (Ng & Lim, 1997a); Cluny Road
drain (Ng & Lim, 1997a); Mandai Road area (Ng & Lim,
1997a); Upper Seletar Reservoir streams (Anonymous, 1988a,
1990; Ng & Lim, 1997a, 1997b); Lower Seletar Reservoir
(Ng HH & Tan HH, 2010); MacRitchie Reservoir (Ng HH
& Tan HH, 2010; Tan, 2016a); Kranji Marsh (Lim et al.,
2013; Tan et al., 2016); Punggol Reservoir (Ng HH & Tan
HH, 2013); Serangoon Reservoir (Ng HH & Tan HH, 2013);
Pasir Panjang canal (Tan HH et al., 2015); Central Catchment
Nature Reserve — Nee Soon Swamp Forest (Lim & Ng,
1992; Ho et al., 2016; Li et al., 2016); Kent Ridge Park
ponds (Tan, 2016b); Yew Tee canal (Tan HH, pers. obs.,
2016); Marina South (Tan HH, pers. obs., 2016); Bishan
Park (Liew JH, pers. obs., 2016); Sungei Seletar (Banir
conuence; Tan HH, pers. obs., March 2017).
Remarks. This species is occasionally sold as live feed in
the ornamental sh trade (Tan HH, pers. obs.).
Rasbora cf. notura Kottelat
(Fig. 41, SEA[SPC])
Fig. 41. Rasbora cf. notura, 57.8 mm SL, Banir stream.
References. Kwik, 2014; Kwik & Yeo, 2015.
Distribution. Banir (Kwik, 2014; Kwik & Yeo, 2015).
Remarks. Originally thought to be a non-native species,
it could potentially represent a new taxon or hybrid (Tan
HH, in prep.).
Fig. 42. Rasbora trilineata, 33.0 mm SL, Catchment Pond.
Fig. 43. Tor tambra, 80.0 mm SL, Sabah.
References. Ng HH & Tan HH, 2010.
Distribution. Pandan Reservoir (Ng HH & Tan HH, 2010).
Tor tambroides (Bleeker)
(Fig. 44, SEA)
References. Herre & Myers, 1937; Alfred, 1966a.
Distribution. Not known.
Remarks. Herre & Myers’ (1937) record came from an
unnamed stream in Singapore, and no further details were
available. No records of this taxon recently. Species from
the genus Tor are unlikely to be native to Singapore, due
to lack of suitable habitat. Tor species are found typically
in large river habitats, which geologically never existed in
Singapore.
Fig. 44. Tor tambroides, 168.0 mm SL, Sarawak.
Rasbora trilineata Steindachner
(Fig. 42, SEA[SPC])
References. Ng & Lim, 1997a, 1997b; Lim & Tan, 2011;
Baker & Lim, 2012; Tan & Zeng, 2014.
Distribution. Old Mandai Road area (Ng & Lim, 1997a);
Bukit Timah Nature Reserve — Catchment Pond (Lim &
Tan, 2011; Tan & Zeng, 2014).
Remarks. This species appears to be established at one
location although many individuals from this population
appear to be inbred and deformed (see Tan & Zeng, 2014).
Tor tambra (Valenciennes, in Cuvier & Valenciennes)
(Fig. 43, SEA[SPC])

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References. Ng HH & Tan HH, 2010.
References. Admin, 2006.
Distribution. Sungei Buloh Wetland Reserve (Chim CK,
pers. comm., 2006); Kranji Reservoir (Admin, 2006,
identied as Chanda sp.; Lim RHB, pers. comm., 2018);
Murai Reservoir (Lim RHB, pers. comm., 2018).
Fig. 45. Toxabramis houdemeri, 75.7 mm SL, Yishun Pond.
Toxabramis houdemeri Pellegrin
(Fig. 45, EAs[SPC])
Fig. 46. Trigonopoma gracile, 32.0 mm SL, Sumatra.
References. Ng & Lim, 1997b (as Rasbora gracilis).
Distribution. Not known.
Remarks. Probably an escapee from the ornamental sh
trade (Ng & Lim, 1997b). Earlier workers (pre-1990) used
the name Rasbora taeniata (Kottelat, 1991).
Family Botiidae
Chromobotia macracanthus (Bleeker)
(Fig. 47, SEA[SPC])
Fig. 47. Chromobotia macracanthus, ca. 250 mm SL, Kranji
Reservoir.
Fig. 48. Gyrinocheilus aymonieri, 136.0 mm SL, trade material
(top); 117.4 mm SL, xanthic variety, Bedok Reservoir (bottom).
References. None. This is a new record for Singapore.
Distribution. Bedok Reservoir (Anonymous, pers. comm.,
2015).
Remarks. This is probably a discarded pet, as the specimen
obtained was a xanthic variant with a bright yellow orange
body but with normal pigmented eyes. A normal coloured
individual is illustrated in Fig. 48 for reference.
Order Characiformes
Family Characidae
Hemigrammus rodwayi Durbin
(Fig. 49, SA[SPC])
Fig. 49. Hemigrammus rodwayi, 26.5 mm SL, Kranji Reservoir.
Distribution. Kranji Reservoir (Ng HH & Tan HH, 2010).
Remarks. This record is probably an escapee, as this species
is very popular in the ornamental sh trade, being seasonally
available after monsoon season (Tan HH, pers. obs.).
Family Gyrinocheilidae
Gyrinocheilus aymonieri (Tirant)
(Fig. 48A, B, SEA[SPC])
References. Lim & Kwik, 2012 (as Toxobramis houdemeri);
Kwik et al., 2013 (as Toxobramis houdemeri).
Distribution. Yishun Pond (Lim & Kwik, 2012; Kwik et
al., 2013).
Remarks. This species is sold as live feed for in the
ornamental sh trade (Tan HH, pers. obs.).
Trigonopoma gracile (Kottelat)
(Fig. 46, SEA)

166
Tan et al.: Non-native freshwater shes of Singapore
Fig. 50. Paracheirodon innesi, 26.8 mm SL, trade material.
References. Ng et al., 1993; Ng & Lim, 1997a, 1997b.
Distribution. Recorded as an escapee in the Mandai area
and not an established species (Ng & Lim, 1997a).
Thayeria boehlkei Weitzman
(Fig. 51, SA)
Fig. 51. Thayeria boehlkei, 25.5 mm SL, trade material.
References. None. This is a new record for Singapore.
Distribution. Based on a sighting at Bukit Batok Nature
Park (Lim KKP, pers. obs., 2010).
Family Serrasalmidae
Piaractus brachypomus (Cuvier)
(Fig. 52, SA)
References. Lim & Ng, 1990 (as Colossoma sp.); Ng et al.,
1993 (as Colossoma macropomum); Ng & Lim, 1997a (as
Colossoma macropomum); Ng & Lim, 1997b (as Colossoma
macropomum); Ng HH & Tan HH, 2010; Baker, 2013e; Tan
& Leong, 2015; Tan, 2016b.
Distribution. Jurong Lake (Ng HH & Tan HH, 2010);
MacRitchie Reservoir (Ng HH & Tan HH, 2010); Bukit
Timah Nature Reserve — Catchment Pond (Baker, 2013e);
Punggol Reservoir (Tan & Leong, 2015); Kent Ridge Park
pond (Tan, 2016b); Serangoon reservoir (unpublished data).
Remarks. This species is not known to breed in Singapore
(Ng & Lim, 1997a). All recent records are released/discarded
pets (Ng HH & Tan HH, 2010).
Order Siluriformes
Family Bagridae
Hemibagrus capitulum (Popta)
(Fig. 53, SEA[SPC])
Fig. 53. Hemibagrus capitulum, ca. 350 mm SL, Jurong Lake.
References. Alfred, 1966a (as Mystus nemurus); Ng HH &
Tan HH, 2010 (as Hemibagrus nemurus).
Distribution. Jurong Lake (Ng HH & Tan HH, 2010).
Remarks. Ng & Kottelat (2013b) recently revised the
taxonomy of the Asian Hemibagrus, and resolved many issues
with existing species. The common lowland species found
throughout most of the lower Sundaic drainages has been
re-identied as H. capitulum; whereas H. nemurus, which
was the name erroneously used for the species in Singapore
previously, is restricted to the island of Java.
Hemibagrus guttatus (Lacepède) [Extirpated]
(EAs)
References. Günther, 1864 (as Macrones elongatus);
Fowler, 1938 (as Mystus elongatus); Alfred, 1966a (as
Mystus elongatus); Ng et al., 1993; Ng & Lim, 1997b; Ng
& Kottelat, 1998.
Distribution. Not known.
Remarks. The rst record of this species (deposited in
ZRC in 2006) was from a brackish water pond at Sungei
Buloh Wetland Reserve and could have been ushed out
from one of the sh farms adjacent to the reserve (Tan HH,
pers. obs.). In 2006, and more recently in 2018, this species
has been observed in large schools in Kranji and Murai
Reservoir (Admin, 2006; Lim RHB, pers. obs.). This sh
has been occasionally found in waters with varying degrees
of salinity (Froese, 2018). The persistence of this sh in
Kranji Reservoir from 2006 to present suggests that it has
established a viable breeding population.
Paracheirodon innesi (Myers)
(Fig. 50, SA)
Fig. 52. Piaractus brachypomus, 220 mm SL, Serangoon Reservoir.

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Remarks. This species was imported accidentally through
the food sh route (Alfred, 1966a; Ng et al., 1993; Ng &
Lim, 1997b), and appears to be extirpated in Singapore,
as there had been no recent records. Ng & Kottelat (1998)
claried the taxonomy of the species, and H. elongatus,
the name used previously, is a subjective junior synonym
to H. guttatus.
Mystus castaneus Ng
(Fig. 54, SEA[SPC])
Fig. 54. Mystus castaneus, ca. 65 mm SL, Upper Peirce Reservoir
inlet.
References. Ng HH & Tan HH, 2010.
Distribution. Upper Peirce Reservoir stream (Ng HH &
Tan HH, 2010).
Mystus wolfi (Bleeker)
(Fig. 55, SEA[SPC])
Fig. 55. Mystus wolfi, ca. 200 mm SL, Upper Seletar Reservoir.
References. Ng HH & Tan HH, 2010; Ng, 2012; Ng HH
& Tan HH, 2013.
Distribution. Lower Peirce Reservoir (Ng HH & Tan HH,
2010); Upper Seletar Reservoir (Ng HH & Tan HH, 2010).
Remarks. Ng (2012) has claried the status of Mystus wolfi
in Singapore. The estuarine and dammed river-reservoir
records probably represent native populations (i.e., Murai
and Lower Seletar Reservoirs in Ng HH & Tan HH [2010];
Punggol and Serangoon Reservoirs in Ng PX & Tan HH
[2013]), but the records from the inland reservoirs are
probably non-native stock from raw water transferred from
Johor, Peninsular Malaysia.
Family Clariidae
Clarias gariepinus (Burchell)
(Fig. 56, AF[SPC])
References. Baker & Lim, 2008, 2012; Ng HH & Tan HH,
2010; Tan HH et al., 2010; Tan HTW et al., 2010; Yeo &
Fig. 56. Clarias gariepinus, 240 mm SL, Upper Seletar Reservoir.
Lim, 2010; Baker, 2013d; Baker & Lim, 2013; Kwik et al.,
2013; Ng et al., 2013; Yeo, 2013; Tan, 2014a; Kwik & Yeo,
2015; Tan & Koh, 2015; Ho et al., 2016; Lim et al., 2016;
Tan et al., 2016; Tan et al., 2017b.
Distribution. Bedok Reservoir (Ng HH & Tan HH, 2010);
Jurong Lake (Ng HH & Tan HH, 2010); Kranji Reservoir
(Ng HH & Tan HH, 2010); Upper Seletar Reservoir (Ng
HH & Tan HH, 2010; Baker & Lim, 2013); Lower Seletar
Reservoir (Ng HH & Tan HH, 2010); Pandan Reservoir
(Ng HH & Tan HH, 2010); Upper Peirce Reservoir (Ng
HH & Tan HH, 2010); Rochor canal (now part of Marina
Reservoir; Tan HH et al., 2010); Singapore Botanic Gardens
(Yeo & Lim, 2010); Lower Peirce Reservoir (Baker, 2013d);
Pangsua pond (Kwik et al., 2013); Woodlands Town Park
pond (Kwik et al., 2013); Yishun pond (Kwik et al., 2013);
Bishan Park (Yeo, 2013; Tan & Koh, 2015); West Coast
Park marsh pond (Tan, 2014a); Singapore Botanic Gardens
(Tan HH, pers. obs., 2015); Pulau Tekong (Lim et al., 2016);
Kranji Marsh (Tan et al., 2016); Singapore Quarry (Tan et
al., 2017b).
Remarks. This African species appears to have displaced
the native congener, Clarias batrachus in many of its known
habitats in Singapore, and is the subject of present research
by Low BW (Ng et al., 2013). The juveniles of this species
range from 1–2 cm to 15 cm length, and are commonly sold
as live feed in the ornamental sh trade (Tan HH, pers. obs.).
The species has been observed to be the subject of mercy/
religious release (Yeo DCJ, pers. obs.).
Clarias macrocephalus Günther
(SEA)
References. Tan HTW et al., 2010.
Distribution. Not known.
Remarks. Hybrids between C. macrocephalus and C.
gariepinus have been used for aquaculture (Tan HTW et
al., 2010).
Family Callichthyidae
Corydoras aeneus (Gill)
(Fig. 57, SA)
References. Anonymous, 1988b; Ng & Lim, 1997a, 1997b.
Distribution. Recorded as an escapee in Mandai area and
not an established species (Anonymous, 1988b; Ng & Lim,
1997a).

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Tan et al.: Non-native freshwater shes of Singapore
Fig. 57. Corydoras aeneus, 33.6 mm SL, trade material.
Remarks. This species is occasionally sold as live feed in
the ornamental sh trade (Tan HH, pers. obs.).
Family Loricariidae
Pterygoplichthys disjunctivus (Weber)
(Fig. 58, SA[SPC])
Fig. 58. Pterygoplichthys disjunctivus, 355 mm SL, Woodlands
Pond (dorsal, lateral, and ventral views).
Central Catchment Nature Reserve — Nee Soon Swamp
Forest (Ng & Lim, 1989); Sungei Buloh Wetland Reserve
(National Parks Board, 2003); Choa Chu Kang ponds (Page
& Robins, 2006); Mandai area (Ng & Lim, 1989; Page &
Robins, 2006); Bedok Reservoir (Ng HH & Tan HH, 2010);
Jurong Lake (Ng & Lim, 1989; Ng HH & Tan HH, 2010);
Kranji Reservoir (Ng & Lim, 1989; Ng HH & Tan HH,
2010); Upper Peirce Reservoir (Ng & Lim, 1989; Ng HH
& Tan HH, 2010); Lower Peirce Reservoir (Ng HH & Tan
HH, 2010); MacRitchie Reservoir (Ng & Lim, 1989; Ng
HH & Tan HH, 2010); Pandan canal (Ng & Lim, 1989);
Woodlands Town Park pond (Kwik et al., 2013); Bishan
Park (Tan, 2016c); Marina Reservoir (unpublished data);
Serangoon Reservoir (unpublished data).
Remarks. Ng & Lim (1989) reported the presence of this
species in the early 1980s although it is uncertain if there
was more than one taxon as both P. disjunctivus and P.
pardalis were extensively available in the ornamental sh
trade (Page & Robins, 2006).
Pterygoplichthys joselimaianus (Weber)
(Fig. 59, SA[SPC])
Fig. 59. Pterygoplichthys joselimaianus, 270 mm SL, Pangsua
Pond (dorsal, lateral, and ventral views).
References. Ng & Lim, 1989 (as Pterygoplichthys sp.);
Munro, 1990 (as Pterygoplichthys sp.); Ng et al., 1993
(as Pterygoplichthys sp.); National Parks Board, 2003 (as
Liposarcus spp.); Page & Robins, 2006; Baker & Lim,
2008, 2012; Ng HH & Tan HH, 2010; Kwik et al., 2013;
Kwik & Yeo, 2015 (as Pterygoplichthys sp.); Ho et al.,
2016; Tan, 2016c.
Distribution. Sungei Sembawang (Ng & Lim, 1989); Hong
Kah (Ng & Lim, 1989); Lim Chu Kang (Ng & Lim, 1989);
Stamford Canal (Ng & Lim, 1989); Pandan Reservoir (Ng
& Lim, 1989); Upper Seletar Reservoir (Ng & Lim, 1989);
References. Ng HH & Tan HH, 2010; Tan, 2014g, 2016c;
Ho et al., 2016.
Distribution. Pandan Reservoir (Ng HH & Tan HH,
2010); Lower Seletar Reservoir (Ng HH & Tan HH, 2010);
Banir (Tan, 2014g; Ho et al., 2016); Bishan Park (Tan,
2016c); Jurong Lake (unpublished data); Marina Reservoir
(unpublished data); Serangoon Reservoir (unpublished data).

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Fig. 60. Pterygoplichthys pardalis, 320 mm SL, Bishan Park (dorsal,
lateral, and ventral views).
References. Anonymous, 1988a (as Hypostomus sp.);
Anonymous, 1988b (as Hypostomus ?plecostomus); Ng &
Lim, 1989 (as Pterygoplichthys sp.); Lim & Ng, 1990 (as
Hypostomus sp.); Ng, 1991 (as Pterygoplichthys sp.); Lim
& Ng, 1992 (as Liposarcus pardalis); Ng & Lim, 1997a
(as Liposarcus pardalis); Ng & Lim, 1997b (as Liposarcus
pardalis); Goh et al., 2002 (as Liposarcus sp.); National Parks
Board, 2003 (as Liposarcus spp.); Page & Robins, 2006;
Baker & Lim, 2008, 2012; Ng HH & Tan HH, 2010; Tan
HTW et al., 2010; Kwik et al., 2013 (as Pterygoplichthys
pardaris); Ho et al., 2016.
Distribution. Mandai area rural streams (Anonymous,
1988b); Sungei Sembawang (Ng & Lim, 1989; Lim & Ng,
1992; Page & Robins, 2006); Hong Kah (Ng & Lim, 1989);
Lim Chu Kang (Ng & Lim, 1989); Stamford Canal (Ng &
Lim, 1989); Pandan Reservoir (Ng & Lim, 1989; Ng HH
& Tan HH, 2010); Pandan Canal (Ng & Lim, 1989); Upper
Seletar Reservoir (Ng & Lim, 1989); Central Catchment
Nature Reserve — Nee Soon Swamp Forest (Ng & Lim,
1989); Kranji Reservoir (Ng & Lim, 1989, 1997a, 1997b; Ng
HH & Tan HH, 2010); Mandai area (Ng & Lim, 1989; Page
& Robins, 2006); Jurong Lake (Ng & Lim, 1989); Upper
Peirce Reservoir (Anonymous, 1988a; Ng & Lim, 1989);
Lower Peirce Reservoir (Ng & Lim, 1997a, 1997b); Upper
Seletar Reservoir (Ng & Lim, 1989, 1997a, 1997b); Sungei
Buloh Wetland Reserve (National Parks Board, 2003); Choa
Chu Kang ponds (Page & Robins, 2006); Lower Seletar
Reservoir (Ng HH & Tan HH, 2010); MacRitchie Reservoir
(Ng & Lim, 1989; Ng HH & Tan HH, 2010); Pangsua pond
References. Ng HH & Tan HH, 2010.
Distribution. Pandan Reservoir (Ng HH & Tan HH, 2010).
Remarks. This individual appears to be a discarded pet.
Family Pangasiidae
Pangasionodon hypophthalmus (Sauvage)
(Fig. 62, SEA)
Fig. 62. Pangasionodon hypophthalmus, 131.0 mm SL, trade
material.
References. Lim & Ng, 1990 (as Pangasianodon sutchi);
Ng et al., 1993 (as Pangasius sutchi); Ng & Lim, 1997a (as
Pangasius hypophthalmus); Ng & Lim, 1997b (as Pangasius
hypophthalmus).
Distribution. Jurong Lake (Lim KKP, pers. obs., 2011).
Remarks. This species is commonly stocked in ponds for
aquacultural and angling purposes (Ng & Lim, 1997a).
Pterygoplichthys pardalis (Castelnau)
(Fig. 60, SA[SPC])
Fig. 61. Synodontis eupterus, 183 mm SL, Pandan Reservoir.
(Kwik et al., 2013); Singapore Botanic Gardens (Tan HH,
pers. obs., 2015); Jurong Lake (unpublished data); Bishan
Park (unpublished data); Bedok Reservoir (unpublished data);
Marina Reservoir (unpublished data); Serangoon Reservoir
(unpublished data); Punggol Reservoir (unpublished data).
Remarks. See earlier remarks on P. disjunctivus.
Family Mochokidae
Synodontis eupterus Boulenger
(Fig. 61, AF[SPC])

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Tan et al.: Non-native freshwater shes of Singapore
Fig. 63. Pangasius sanitwongsei, 102.7 mm SL, trade material.
Fig. 64. Phractocephalus hemioliopterus, 700 mm SL, Pandan
Reservoir.
References. Ng & Lim, 1997b; Ng HH & Tan HH, 2010.
Distribution. Pandan River (Ng & Lim, 1997b, based on a
1996 newspaper report); Pandan Reservoir (Ng HH & Tan
HH, 2010); Bedok Reservoir (Anonymous, 2014).
Remarks. These reported individuals are likely discarded
pets. This species is common in the ornamental sh trade as
juveniles (less than 10 cm). Once shes reach sub-adulthood,
they are typically too large to be accommodated in home
aquaria and are conveniently discarded in public ponds
and reservoirs (Ng HH & Tan HH, 2010). Recently, an
articial hybrid between Phractocephalus hemioliopterus
and Pseudoplatystoma fasciatum became available through
the ornamental sh trade (Tan HH, pers. obs.). This articial
hybrid can grow to a large size as both parental species
exceed 90 cm in size, and introductions can be expected in
the near future in the articial water bodies in Singapore.
Order Cyprinodontiformes
Family Aplocheilidae
Aplocheilus lineatus (Valenciennes, in Cuvier &
Valenciennes)
(Fig. 65, SAs[SPC])
Fig. 65. Aplocheilus lineatus, 47.4 mm SL male (top), 38.9 mm SL
female (centre), 20.1 mm SL juvenile (bottom); Pandan.
Fig. 66. Gambusia afnis, 23.8 mm SL female (top), 19.5 mm SL
male (bottom); Upper Seletar Reservoir inlet.
References. Johnson, 1973a; Johnson, 1973b (as Gambusia
spp.); Anonymous, 1988a; Lim & Ng, 1990 (as Gambusia
holbrooki); Munro, 1990 (as G. holbrooki); Ng et al., 1993
(as G. holbrookii); Fernando & Phang, 1994; Lim et al., 1995
(as G. holbrooki); Subaraj et al., 1995 (as G. holbrooki);
Lever, 1996; Ng & Lim, 1997a (as G. holbrookii); Ng &
Lim, 1997b (as G. holbrookii); Baker & Lim, 2008, 2012;
Ng HH & Tan HH, 2010; Tan HTW et al., 2010; Yeo &
Lim, 2010; Davison et al., 2012; Ng HH & Tan HH, 2013;
Tan, 2014d, 2016a; Kwik & Yeo, 2015; Ho et al., 2016;
Lim et al., 2016; Tan et al., 2016; Tan et al., 2017b.
Distribution. Seletar Reservoir Park (Munro, 1990); Sungei
Whampoa (Munro, 1990); Lornie Forest stream (Subaraj
et al., 1995); Ulu Sembawang/Lorong Gambas (Lim et al.,
Pangasius sanitwongsei Smith
(Fig. 63, SEA)
References. Tan & Yusandi, 2015.
Distribution. Marina Reservoir (Tan & Yusandi, 2015).
Family Pimelodidae
Phractocephalus hemioliopterus (Bloch & Schneider)
(Fig. 64, SA[SPC])
References. Yeo & Lim, 2010; Baker & Lim, 2012; Kwik
& Yeo, 2015; Tan et al., 2017a.
Distribution. Bukit Batok Nature Park (Yeo & Lim, 2010;
Kwik & Yeo, 2015); Singapore Botanic Gardens (Yeo &
Lim, 2010); Sungei Pandan feeder stream (Tan et al., 2017a).
Remarks. This species has established breeding populations
in Bukit Batok Nature Park (Yeo & Lim, 2010), and Sungei
Pandan feeder stream (Tan et al., 2017a).
Family Poeciliidae
Gambusia afnis (Baird & Girard)
(Fig. 66, CA[SPC])

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RAFFLES BULLETIN OF ZOOLOGY 2020
1995); Bedok Reservoir (Ng HH & Tan HH, 2010); Jurong
Lake (Ng HH & Tan HH, 2010); Kranji Reservoir (Ng HH
& Tan HH, 2010); Lower Peirce Reservoir (Anonymous,
1988a; Lim et al., 1995; Ng & Lim, 1997a, 1997b; Ng HH &
Tan HH, 2010); Kent Ridge Park pond (Ng & Lim, 1997a);
Lower Seletar Reservoir (Ng HH & Tan HH, 2010); Murai
Reservoir (Ng HH & Tan HH, 2010); Tengah Reservoir (Ng
HH & Tan HH, 2010); Upper Peirce Reservoir (Ng HH &
Tan HH, 2010); Upper Seletar Reservoir (Ng HH & Tan
HH, 2010; Tan, 2014d); Singapore Botanic Gardens (Yeo &
Lim, 2010); Punggol Reservoir (Ng HH & Tan HH, 2013);
Serangoon Reservoir (Ng HH & Tan HH, 2013); MacRitchie
Reservoir (Subaraj et al., 1995; Tan, 2016a); outskirts of Nee
Soon Swamp Forest (Ho et al., 2016); Pulau Tekong (Lim
et al., 2016); Kranji Marsh (Tan et al., 2016); Singapore
Quarry (Tan et al., 2017b).
Remarks. This species had often been introduced as a
mosquito control agent but its efficacy is questionable
(Lever, 1996). This species is commonly sold with either
Poecilia reticulata or Poecilia sphenops as live feed in the
ornamental sh trade (Tan HH, pers. obs.).
Poecilia latipinna (Lesueur)
(CA)
References. Welcomme, 1988; Lim & Ng, 1990; Munro,
1990; Ng et al., 1993; Fernando & Phang, 1994; Lever,
1996; Ng & Lim, 1997a (as Poecilia cf. latipinna), 1997b
(as P. latippina).
Distribution. Widespread in canals and brackish water
habitats (Lim & Ng, 1990). It may have hybridised with P.
sphenops (Lim & Ng, 1990).
Poecilia reticulata Peters
(Fig. 67, CA[SPC])
& Lim, 2008, 2012; Ng HH & Tan HH, 2010; Tan HTW et
al., 2010; Yeo & Chia, 2010; Yeo & Lim, 2010; Ng PX &
Tan HH, 2013; Tan et al., 2013; Tan & Low, 2014; Kwik
& Yeo, 2015; Ho et al., 2016; Tan, 2016b.
Distribution. Widespread in Singapore, except in forest
streams (Maxwell, 1921; Alfred, 1966a; Lim & Ng, 1990);
MacRitchie Reservoir (Alfred, 1966a); Upper Seletar
Reservoir (Alfred, 1966a; Ng & Lim, 1997a, 1997b); Upper
Peirce Reservoir (Anonymous, 1989b); Central Catchment
Nature Reserve — Nee Soon Swamp Forest (Munro, 1990);
Napier Road and Bukit Timah Road (Munro, 1990); Seletar
Reservoir Park (Munro, 1990); Kent Ridge Park (Ng & Lim,
1997a); Pandan Canal (Ng & Lim, 1997a); Cluny Road
drain (Ng & Lim, 1997a); Bukit Timah Nature Reserve
peripheral streams (Lim, 1995; Ng & Lim, 1997b); Sungei
Buloh Wetland Reserve (National Parks Board, 2003);
Lower Seletar Reservoir (Ng HH & Tan HH, 2010); Bukit
Batok Nature Park (Yeo & Lim, 2010); Punggol Reservoir
(Ng PX & Tan HH, 2013); Serangoon Reservoir (Ng PX &
Tan HH, 2013); Bukit Brown (Tan et al., 2013); Greenbank
Park (Tan HH, pers. obs., 1990s); Kranji Reservoir (Tan HH,
pers. obs., 2010); Commonwealth area (Tan HH, pers. obs.,
2010); Mandai area (Tan & Low, 2014); Tai Seng area (Tan
HH, pers. obs., 2016); outskirts of Nee Soon Swamp Forest
(Ho et al., 2016); Kent Ridge Park ponds (Tan, 2016b).
Remarks. Tham (1973) commented that Poecilia reticulata
was used for mosquito control in sh ponds. This species
is commonly sold as live feed in the ornamental sh trade
(Tan HH, pers. obs.). See later account under “Species for
biological control” for more details.
Poecilia sphenops Valenciennes, in Cuvier &
Valenciennes
(Fig. 68, CA[SPC])
Fig. 67. Poecilia reticulata, 14.8 mm SL male (top), 20.0 mm SL
female (bottom); Mandai area. Fig. 68. Poecilia sphenops, 50.2 mm SL male (top), 57.0 mm SL
female (bottom); Pandan canal.
References. Hanitsch, 1912 (as Girardinus poeciloides);
Herre, 1940 (as Lebistes reticulatus); Johnson, 1964 (as
Lebistes reticulatus); Alfred, 1966a (as Lebistes reticulatus);
Johnson, 1973b; Tham, 1973 (as Lebistes reticulatus);
Anonymous, 1989b; Lim & Ng, 1990; Munro, 1990; Ng,
1991; Ng et al., 1993; Fernando & Phang, 1994 (introduced
as early as 1937); Lim, 1995; Lever, 1996; Ng & Lim, 1997a,
1997b; Goh et al., 2002; National Parks Board, 2003; Baker
References. Johnson & Soong, 1963 (as Mollienisia
sphenops); Alfred, 1966a (as Mollienisia sphenops); Johnson,
1973a (as Mollienisia sphenops); Johnson, 1973b (as
Mollienisia spp.); Lim & Ng, 1990; Munro, 1990; Ng, 1991;
Ng et al., 1993; Fernando & Phang, 1994; Subaraj, 1995;
Lever, 1996; Ng & Lim, 1997a, 1997b; Ng & Sivasothi,
1999; Goh et al., 2002; National Parks Board, 2003; Ng

172
Tan et al.: Non-native freshwater shes of Singapore
HH & Tan HH, 2010; Tan HH et al., 2010; Tan HTW et
al., 2010; Jaafar et al., 2012; Leong, 2012; Ng HH & Tan
HH, 2013; Tan, 2014d, 2016a; Ng et al., 2015; Ho et al.,
2016; Li et al., 2016.
Distribution. MacRitchie Reservoir (Alfred, 1966a; Tan,
2016a); Pulau Ubin (Subaraj, 1995); Cluny Road drain (Ng
& Lim, 1997a); Pandan Canal (Ng & Lim, 1997a); Sungei
Sembawang — Senoko (Ng & Lim, 1997a); Mandai coastal
streams (Tan HH, pers. obs., 2004); Sungei Buloh Wetland
Reserve (National Parks Board, 2003); Bedok Reservoir
(Ng HH & Tan HH, 2010); Lower Seletar Reservoir (Ng
HH & Tan HH, 2010); Pandan canal (Tan HH, pers. obs.,
2010); Jurong River (Ng HH, pers. obs., 2010); Kranji
Reservoir (Tan HH, 2010); Marina Reservoir (comprising
of Singapore River, Rochor canal, Kallang River, Geylang
River and Marina Channel; Tan HH et al., 2010); Siglap
canal (Leong, 2012); Punggol Reservoir (Ng HH & Tan
HH, 2013); Serangoon Reservoir (Ng HH & Tan HH, 2013);
Upper Seletar Reservoir (Tan, 2014d); Singapore Botanic
Gardens (Tan HH, pers. obs., 2015); Central Catchment
Nature Reserve — Nee Soon Swamp Forest (Ho et al.,
2016; Li et al., 2016); Yew Tee canal (Tan HH, pers. obs.,
2016); West Coast Park marsh pond (Tan HH, pers. obs.,
November 2016); Bishan Park (Liew JH, pers. obs., 2016).
Remarks. Fernando & Phang (1994) stated that this species
had been very likely introduced as an escaped pet sh in
the 1940s, and they easily hybridise with P. latipinna. This
species is commonly sold as live feed in the ornamental sh
trade (Tan HH, pers. obs.).
Poecilia velifera (Regan)
(Fig. 69, CA)
Distribution. Recorded by Ng & Lim (1997a) from canals in
Singapore, but no details provided; Woodlands Town Garden
(Anonymous, 1989a); Pulau Tekong (Lim et al., 2016).
Remarks. Fernando & Phang (1994) noted that the
introductions of X. hellerii, X. maculatus and X. variatus
into Singapore for ornamental aquaculture probably started
in the mid-1940s. This species is occasionally sold as live
feed in the ornamental sh trade (Tan HH, pers. obs.).
Xiphophorus maculatus (Günther)
(Fig. 71, CA[SPC])
Fig. 69. Poecilia velifera, 60 mm SL male, trade material.
Fig. 70. Xiphophorus hellerii, 41.8 mm SL male (top), 39.1 mm
SL female (bottom); trade material (wild type colour form).
Fig. 71. Xiphophorus maculatus, 27.3 mm SL, Mount Pleasant.
References. Lim & Ng, 1990; Ng et al., 1993; Fernando &
Phang, 1994; Ng & Lim, 1997a, 1997b; Tan HH et al., 2010;
Yeo & Lim, 2010; Tan et al., 2013; Kwik & Yeo, 2015.
Distribution. Cluny Road drain (Ng & Lim, 1997a); Kallang
River (now part of Marina Reservoir; Tan HH et al., 2010);
Bukit Batok Nature Park (Yeo & Lim, 2010); Bukit Brown
(Tan et al., 2013).
Remarks. This species is commonly sold as live feed in the
ornamental sh trade (Tan HH, pers. obs.).
Xiphophorus variatus (Meek)
(Fig. 72, CA[SPC])
References. Munro, 1990; Ng et al., 1993.
Distribution. Not known.
Remarks. The records of this species are probably escapees
from the ornamental sh trade (Munro, 1990).
Xiphophorus hellerii Heckel
(Fig. 70, CA[SPC])
References. Anonymous, 1989a; Ng et al., 1993; Fernando
& Phang, 1994; Ng & Lim, 1997a, 1997b; Lim et al., 2016.
Fig. 72. Xiphophorus variatus, 28.3 mm SL, Bukit Brown (note
deformed dorsal n).

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Fig. 73. Macrognathus siamensis, 120.4 mm SL, trade material.
References. Ng & Lim, 1997a, 1997b.
Distribution. Recorded as an escapee in Seletar Reservoir
(within Central Catchment Nature Reserve) and not an
established species (Ng & Lim, 1997a).
Macrognathus zebrinus (Blyth)
(Fig. 74, SEA[SPC])
Fig. 74. Macrognathus zebrinus, 195.9 mm SL, Lower Seletar
Reservoir.
References. Baker & Lim, 2008, 2012; Ng, 2010; Ng HH
& Tan HH, 2010; Baker & Leong, 2014; Ho et al., 2016;
Tan et al., 2016; Tan & Teo, 2018.
Distribution. Kranji Reservoir (Ng, 2010; Ng HH & Tan
HH, 2010); Lower Peirce Reservoir (Ng, 2010; Ng HH &
Tan HH, 2010); Lower Seletar Reservoir (Ng, 2010; Ng HH
& Tan HH, 2010); Tengah Reservoir (Ng, 2010; Ng HH &
Tan HH, 2010); Banir (Baker & Leong, 2014; Ho et al.,
2016); Kranji Marsh (Tan et al., 2016); Marina Reservoir
(Tan & Teo, 2018).
Distribution. Originally found in the northern half of
Singapore, recent records show it is now at Marina Reservoir
(see Tan & Teo, 2018). The mode of transfer or spread is
unknown.
Order Perciformes
Family Ambassidae
Parambassis siamensis (Fowler)
(Fig. 75, SEA[SPC])
References. Anonymous, 1989b (as ?Chanda nama);
Anonymous, 1990 (as Parambassis punctulatus); Lim &
Ng, 1990 (as Chanda sp.); Munro, 1990 (as Parambassis
punctulatus); Ng et al., 1993 (as Parambassis punctulatus);
Fig. 75. Parambassis siamensis, 39.9 mm SL, Serangoon Reservoir.
References. Ng et al., 1993; Fernando & Phang, 1994; Ng
& Lim, 1997a; Tan et al., 2013.
Distribution. Bukit Brown (Tan et al., 2013).
Order Synbranchiformes
Family Mastacembelidae
Macrognathus siamensis (Günther)
(Fig. 73, SEA)
Lim et al., 1995; Subaraj et al., 1995; Ng & Lim, 1997a,
1997b; Baker & Lim, 2008, 2012; Ng HH & Tan HH, 2010;
Kwik et al., 2013; Ng HH & Tan HH, 2013; Ho et al., 2016;
Li et al., 2016; Tan, 2018.
Distribution. Lim & Ng (1990) observed this species in
Central Catchment reservoirs; Kranji Reservoir (Ng &
Lim, 1997a, 1997b; Ng HH & Tan HH, 2010); Upper
Peirce Reservoir (Anonymous, 1989b; Ng & Lim, 1997a,
1997b; Ng HH & Tan HH, 2010); Lower Peirce Reservoir
(Anonymous, 1990; Munro, 1990; Lim et al., 1995; Ng &
Lim, 1997a, 1997b; Ng HH & Tan HH, 2010); Bukit Batok
Nature Park (Tan, 2018); Jurong Lake (Ng HH & Tan
HH, 2010); Upper Seletar Reservoir (Ng HH & Tan HH,
2010); Lower Seletar Reservoir (Ng HH & Tan HH, 2010);
MacRitchie Reservoir (Subaraj et al., 1995; Ng HH & Tan
HH, 2010); Murai Reservoir (Ng HH & Tan HH, 2010);
Singapore Botanic Gardens, Swan Lake (Lim KKP, pers.
obs., 2010); Pangsua pond (Kwik et al., 2013); Woodlands
Town Park pond (Kwik et al., 2013); Punggol Reservoir (Ng
HH & Tan HH, 2013); Serangoon Reservoir (Ng HH & Tan
HH, 2013); Central Catchment Nature Reserve—Nee Soon
Swamp Forest (Ho et al., 2016; Li et al., 2016); Sungei Seletar
(Banir conuence; Tan HH, pers. obs., March 2017); Jurong
Lake (unpublished data); Bishan Park (unpublished data);
Pandan Reservoir (unpublished data); Serangoon Reservoir
(unpublished data); Bedok Reservoir (unpublished data).
Remarks. This species was probably introduced into
Singapore’s reservoirs along with raw water transferred from
Johor, Peninsular Malaysia (Ng HH & Tan HH, 2010). This
species consumes zooplankton and insect larvae (Okutsu et
al., 2011; Lim RHB, pers. obs.).
Family Datnioididae
Datnioides microlepis Bleeker
(Fig. 76, SEA[SPC])
References. Ng HH & Tan HH, 2010.
Distribution. Pandan Reservoir (Ng HH & Tan HH, 2010);
Bedok Reservoir (unpublished data).
Remarks. This species is probably from discarded pet sh,
as this is a popular species in the ornamental sh trade. As

174
Tan et al.: Non-native freshwater shes of Singapore
Fig. 76. Datnioides microlepis, 290 mm SL, Pandan Reservoir.
with other taxa commonly discarded, this species also grows
to a large size not suitable for most home aquarists.
Family Monodactylidae
Monodactylus sebae (Cuvier)
(Fig. 77, AF)
Fig. 77. Monodactylus sebae, 59.5 mm SL, trade material.
Family Cichlidae
Acarichthys heckelli (Müller & Troschel)
(Fig. 78, SA[SPC])
Fig. 78. Acarichthys heckelli, 96.0 mm SL male, Lower Peirce
Reservoir.
Fig. 79A. Amphilophus citrinellus, 80.6 mm SL juvenile barred
form, Pangsua Pond.
References. Tan & Lim, 2008; Baker & Lim, 2008, 2012;
Ng HH & Tan HH, 2010; Kwik et al., 2013; Liew et al.,
2013; Tan, 2014d, 2016a; Ho et al., 2016.
Distribution. Central Catchment Nature Reserve — Nee
Soon Swamp Forest (Tan & Lim, 2008; Ho et al., 2016);
Upper Peirce Reservoir (Ng HH & Tan HH, 2010; Liew
et al., 2013); Lower Peirce Reservoir (Ng HH & Tan HH,
2010; Liew et al., 2013); Lower Seletar Reservoir (Ng HH
& Tan HH, 2010; Liew et al., 2013); Woodlands Town
Park pond (Kwik et al., 2013); Sungei Seletar (Liew et al.,
2013); Banir (Liew et al., 2013); Upper Seletar Reservoir
(Tan, 2014d); MacRitchie Reservoir (Tan, 2016a); Bishan
Park (unpublished data); Marina Reservoir (unpublished
data); Punggol Reservoir (unpublished data).
Remarks. Autecology of this recently introduced species
was documented by Liew et al. (2013).
Amphilophus citrinellus (Günther)
(Fig. 79A, B, CA[SPC])
References. Ng HH & Tan HH, 2010; Kwik et al., 2013;
Lim, 2013c; Yeo, 2013; Tan, 2014f, 2016c.
Distribution. Jurong Lake (Ng HH & Tan HH, 2010);
Pandan Reservoir (Ng HH & Tan HH, 2010); Pangsua Pond
(Kwik et al., 2013; Tan, 2014f); Woodlands Town Park
References. Tan & Soo, 2016.
Distribution. Marina Reservoir (Tan & Soo, 2016).
Remarks. This individual is probably a discarded pet sh,
as this species is a popular sh species in the ornamental
sh trade.

175
RAFFLES BULLETIN OF ZOOLOGY 2020
Pond (Kwik et al., 2013); Yishun Pond (Kwik et al., 2013;
Lim, 2013c); Bishan Park (Yeo, 2013; Tan, 2016c); Marina
South (Tan, 2014f); Pandan canal (Tan, 2014f); Sungei Buloh
Wetland Reserve (Tan, 2014f); West Coast Park marsh pond
(Tan, 2015a); Singapore Botanic Gardens (Tan HH, pers.
obs., 2015); Marina Reservoir (Tan HH, pers. obs., 2016);
Bedok Reservoir (unpublished data); Serangoon Reservoir
(unpublished data).
Remarks. This is one of the parent species for the Louhan
cichlid and Blood parrot hybrids.
Amphilophus trimaculatus (Günther)
(Fig. 80, CA)
Remarks. An established population of this species exists
in a stream along the outskirts of Nee Soon Swamp Forest
(Liew et al., 2012; Ho et al., 2016). Its presence has been
known since the 1990s.
Astronotus ocellatus (Agassiz, in Spix & Agassiz)
(Fig. 82, SA)
Fig. 79B. Amphilophus citrinellus, 219.3 mm SL adult male,
Pandan Reservoir.
Fig. 80. Amphilophus trimaculatus, West Coast marsh pond.
References. Tan, 2014a (as Cichlasoma trimaculatum).
Distribution. West Coast Park marsh pond (Tan, 2014a).
Remarks. This is one of the parent species for the Louhan
cichlid.
Apistogramma borelli (Regan)
(Fig. 81, SA[SPC])
References. Tan HTW et al., 2010 (as Apistogramma sp.);
Liew et al., 2012; Ho et al., 2016.
Distribution. Upper Seletar Reservoir Park stream (Liew
et al., 2012); outskirts of Nee Soon Swamp Forest (Ho et
al., 2016).
Fig. 81. Apistogramma borelli, 34.3 mm SL, Upper Seletar area
(Lim KKP).
Fig. 82. Astronotus ocellatus, 59.1 mm SL, trade material.
References. Lim & Ng, 1990; Ng et al., 1993; Ng & Lim,
1997b; Liew et al., 2012.
Distribution. Singapore Botanic Gardens (Liew et al., 2012),
Bishan Park (Tan HH, pers. obs., March 2017).
Remarks. This is a popular species in the ornamental sh
trade, available in several colour varieties.
Cichla kelberi Kullander & Ferreira
(Fig. 83, SA[SPC])
Fig. 83. Cichla kelberi, 232 mm SL, Bedok Reservoir.
References. None. This is a new record for Singapore.
Distribution. Bedok Reservoir (unpublished data).

176
Tan et al.: Non-native freshwater shes of Singapore
Remarks. Species of Cichla are large (more than 40 cm in
length), highly territorial predatory shes. Their presence in
the water bodies will impact existing sh communities as
this genus is highly piscivorous (Kovalenko et al., 2010).
The four Cichla species could have been introduced by
recreational anglers or/and released as unwanted ornamental
pets (Ng HH & Tan HH, 2010).
Cichla orinocensis Humboldt, in Humboldt &
Valenciennes
(Fig. 84, SA[SPC])
References. None. This is a new record for Singapore.
Distribution. Marina Reservoir (unpublished data).
Cichla temensis Humboldt, in Humboldt &
Valenciennes
(Fig. 86, SA[SPC])
Fig. 84. Cichla orinocensis, 217.5 mm SL, Pandan Reservoir.
References. Ng & Lim, 1997a (as Cichla cf. monoculus);
Ng & Lim, 1997b (as Cichla ocellaris); Baker & Lim, 2008,
2012; Ng HH & Tan HH, 2010; Yeo & Chia, 2010; Kwik
et al., 2013; Ng HH & Tan HH, 2013; Tan, 2013b, 2014d,
2014g; Ho et al., 2016.
Distribution. Lower Peirce Reservoir (Ng & Lim, 1997a;
Ng HH & Tan HH, 2010); Bedok Reservoir (Ng HH &
Tan HH, 2010); Jurong Lake (Ng HH & Tan HH, 2010);
Kranji Reservoir (Ng HH & Tan HH, 2010); Upper Seletar
Reservoir (Tan, 2013b, 2014d); Lower Seletar Reservoir (Ng
HH & Tan HH, 2010); MacRitchie Reservoir (Ng HH & Tan
HH, 2010); Pandan Reservoir (Ng HH & Tan HH, 2010);
Woodlands Town Park pond (Kwik et al., 2013); Punggol
Reservoir (Ng HH & Tan HH, 2013); Serangoon Reservoir
(Ng HH & Tan HH, 2013); Banir (Tan, 2014g; Ho et al.,
2016); Marina Reservoir (near Marina Bay Sands; Wozniak
S, pers. obs., October 2016); Bishan Park (unpublished data).
Remarks. Juvenile individuals were recently obtained from
a stream feeding into Lower Seletar Reservoir (Tan, 2014g),
while adults were observed to be breeding in Upper Seletar
Reservoir (Tan, 2013b); this is indicative that this species
has established in Singapore.
Cichla piquiti Kullander & Ferreira
(Fig. 85, SA[SPC])
Fig. 85. Cichla piquiti, 251.7 mm SL, Marina Reservoir.
Fig. 86. Cichla temensis, 245 mm SL, Bedok Reservoir.
References. Liew et al., 2012; Lai, 2015; Ho et al., 2016;
Tan, 2016a.
Distribution. Upper Seletar Reservoir (Liew et al., 2012);
Lower Peirce Reservoir (Lai, 2015); MacRitchie Reservoir
(Tan, 2016a); Bedok Reservoir (unpublished data); Upper
Peirce Reservoir (unpublished data); Marina Reservoir
(unpublished data).
Remarks. Juvenile individuals were obtained from Upper
Seletar Reservoir (Liew et al., 2012), while adult pairs have
been observed at MacRitchie Reservoir (Tan, 2016a); this is
indicative of this species establishing breeding populations
in Singapore.
Coptodon zillii (Gervais)
(Fig. 87, AF)
Fig. 87. Coptodon zillii, 36.5 mm SL, Sumatra.
References. Burkill, 1963 (as Tilapia zillii); Alfred, 1968
(as Tilapia zillii); Munro, 1990 (as Tilapia zillii); Ng et al.,
1993 (as Tilapia zillii); Ng & Lim, 1997a (as Tilapia zillii),
1997b (as Tilapia zillii).
Distribution. Singapore Botanic Gardens (Burkill, 1963;
Alfred, 1968; Ng & Lim, 1997a, 1997b).
Remarks. There have been no recent records.

177
RAFFLES BULLETIN OF ZOOLOGY 2020
Etroplus suratensis (Bloch)
(Fig. 88, SAs[SPC])
Upper Peirce Reservoir (Ng HH & Tan HH, 2010); Lower
Peirce Reservoir (Ng HH & Tan HH, 2010); Upper Seletar
Reservoir (Ng HH & Tan HH, 2010; Tan, 2014d); Lower
Seletar Reservoir (Ng HH & Tan HH, 2010); MacRitchie
Reservoir (Ng HH & Tan HH, 2010; Tan, 2016a); Tekong
Reservoir (Ng HH & Tan HH, 2010); Woodlands Town Park
pond (Kwik et al., 2013); Yishun pond (Kwik et al., 2013);
outskirts of Nee Soon Swamp Forest (Ho et al., 2016); Pulau
Tekong (Lim et al., 2016); Jurong Lake (unpublished data);
Pandan Reservoir (unpublished data); Serangoon Reservoir
(unpublished data).
Remarks. This is a common species in the ornamental
sh trade.
Herichthys carpintis (Jordan & Snyder)
(Fig. 90, CA[SPC])
Fig. 88. Etroplus suratensis, 185 mm SL, Marina Reservoir
(Kallang).
References. Lim et al., 1995; Ng & Sivasothi, 1999; Goh et
al., 2002; Chua EK, 2010; Ng TH & Tan, 2010; Ng HH &
Tan HH, 2010; Tan HH et al., 2010; Tan HTW et al., 2010;
Jaafar et al., 2012; Leong, 2012; Kwik et al., 2013; Ng PX
& Tan HH, 2013; Tan, 2013a; Ng et al., 2015; Tan, 2016c,
2017a; Tan & Ng, 2016; Theng et al., 2016; Toh et al., 2016.
Distribution. Sungei Buloh Wetland Reserve (Lim et al.,
1995; Ng TH & Tan, 2010; Tan, 2013a; Theng et al., 2016);
Kallang (now part of Marina Reservoir; Ng TH & Tan, 2010);
Kranji Reservoir (Ng HH & Tan HH, 2010); Lower Seletar
Reservoir (Ng HH & Tan HH, 2010); Punggol Reservoir
(Ng TH & Tan HH, 2010; Ng PX & Tan, 2013; Theng et
al., 2016); Serangoon Reservoir (Ng TH & Tan HH, 2010;
Ng PX & Tan HH, 2013; Theng et al., 2016); Pulau Ubin
(Ng TH & Tan, 2010); Marina Reservoir — encompassing
Singapore River, Rochor canal, Kallang River, Geylang River
and Marina channel (Tan HH et al., 2010); Siglap canal
(Leong, 2012); Yishun pond (Kwik et al., 2013); Tuas —
Rafes Marina (Toh et al., 2016); Bishan Park (Tan, 2016c);
Sentosa (Tan & Ng, 2016); West Coast Park (Tan, 2017a);
Jurong Lake (unpublished data); Lower Peirce Reservoir
(unpublished data); MacRitchie Reservoir (unpublished data);
Bedok Reservoir (unpublished data); Upper Peirce Reservoir
(unpublished data); Serangoon Reservoir (unpublished data).
Remarks. This euryhaline South Asian cichlid is well-
established in Singapore (Ng TH & Tan, 2010) and besides
being found in freshwater reservoirs, has also adapted to
saline conditions of estuarine and coastal waters (Tan &
Ng, 2016; Tan, 2017a).
Geophagus altifrons Heckel
(Fig. 89, SA[SPC])
References. Baker & Lim, 2008, 2012; Ng HH & Tan HH,
2010; Tan HH et al., 2010 (as G. surinamensis); Tan HTW
et al., 2010 (as Geophagus sp.); Kwik et al., 2013; Tan,
2014d, 2016a; Ho et al., 2016; Lim et al., 2016.
Distribution. Kallang River (now part of Marina Reservoir;
Tan HH et al., 2010); Bedok Reservoir (Ng HH & Tan
HH, 2010); Kranji Reservoir (Ng HH & Tan HH, 2010);
Fig. 89. Geophagus altifrons, 145.6 mm SL, Bedok Reservoir.
Fig. 90. Herichthys carpintis, 210 mm SL, Sungei Buloh.
References. Jaafar et al., 2012; Liew et al., 2012.
Distribution. Sungei Buloh Wetland Reserve (Jaafar et al.,
2012; Liew et al., 2012).
Heros severus Heckel
(Fig. 91, SA[SPC])
References. Ng HH & Tan HH, 2010.
Distribution. Bedok Reservoir (Ng HH & Tan HH, 2010).
Remarks. Not established as this is regarded as a discarded
aquarium sh.

178
Tan et al.: Non-native freshwater shes of Singapore
Heterotilapia buttikoferi (Hubrecht)
(Fig. 92, AF[SPC])
Fig. 91. Heros severus, 70.2 mm SL, trade material.
Fig. 92. Heterotilapia buttikoferi, 205 mm SL, Yishun Pond.
References. Baker & Lim, 2008, 2012 (as Tilapia buttikoferi);
Ng HH & Tan HH, 2010 (as Tilapia buttikoferi); Tan
HTW et al., 2010 (as Tilapia buttikoferi); Kwik et al.,
2013 (as Tilapia buttikoferi); Tan, 2013d, 2014a (as Tilapia
buttikoferi); Tan, 2016c.
Distribution. Bedok Reservoir (Ng HH & Tan HH, 2010);
Jurong Lake (Ng HH & Tan HH, 2010); Upper Peirce
Reservoir (Ng HH & Tan HH, 2010); Lower Peirce Reservoir
(Ng HH & Tan HH, 2010); Upper Seletar Reservoir (Ng
HH & Tan HH, 2010); Lower Seletar Reservoir (Ng HH
& Tan HH, 2010); Pandan Reservoir (Ng HH & Tan HH,
2010); Pangsua pond (Kwik et al., 2013); Yishun pond (Kwik
et al., 2013); Central Catchment Nature Reserve — Sime
forest (Tan, 2013d); West Coast Park marsh pond (Tan,
2014a); Bishan Park (Tan, 2016c); Jurong Lake (Tan HH,
pers. obs., 2018); Marina Reservoir (unpublished data);
Pandan Reservoir (unpublished data); Serangoon Reservoir
(unpublished data).
Remarks. This is a popular sh in the ornamental sh trade.
Its introduction into Singapore is probably through pet shes
that are released into the water bodies.
Fig. 93. Mayaheros urophthalmus, 151.1 mm SL, Japanese Garden.
References. National Parks Board, 2003 (as Nandopsis
festae or Cichlasoma urophthalmus); Nico et al., 2007
(as “Cichlasoma” urophthalmus); Baker & Lim, 2008,
2012 (as Cichlasoma urophthalmum); Ng HH & Tan
HH, 2010 (as Cichlasoma urophthalmum); Tan HTW et
al., 2010 (as Cichlasoma urophthalmus); Yeo & Lim,
2010 (as Cichlasoma urophthalmum); Jaafar et al., 2012
(as Cichlasoma urophthalmum); Kwik et al., 2013 (as
Cichlasoma urophthalmus); Ng HH & Tan HH, 2013 (as
Cichlasoma urophthalmum); Tan, 2013a (as Cichlasoma
urophthalmum); Yeo, 2013 (as Cichlasoma urophthalmum);
Tan, 2014a, 2014f (as Cichlasoma urophthalmum); Ng et
al., 2015 (as Cichlasoma urophthalmum); Tan, 2016c; Tan
& Koh, 2016 (as Cichalsoma uropthalmum); Theng et al.,
2016 (as Cichlasoma urophthalmus); Tan et al., 2017b.
Distribution. Sungei Buloh Wetland Reserve (National Parks
Board, 2003; Tan, 2013a; Theng et al., 2016); Punggol River
(now dammed as Punggol Reservoir; Nico et al., 2007);
Bedok Reservoir (Ng HH & Tan HH, 2010); Sarimbun
Reservoir (Ng HH & Tan HH, 2010); Upper Peirce Reservoir
(Ng HH & Tan HH, 2010); Singapore Botanic Gardens (Yeo
& Lim, 2010); Punggol Reservoir (Ng PX & Tan, 2013;
Theng et al., 2016); Serangoon Reservoir (Ng PX & Tan,
2013; Theng et al., 2016); Bishan Park (Yeo, 2013; Tan,
2016c); West Coast Park marsh pond (Tan, 2014a); Pandan
canal (Tan 2014f); Yishun Pond (Kwik et al., 2013; Tan,
2014f); Jurong Lake (Tan, 2014f); Marina South (Tan &
Koh, 2016); Singapore Quarry (Tan et al., 2017b); Bishan
Park (unpublished data).
Remarks. This species has adapted to brackish water and
can be sighted in the waterways at Sungei Buloh Wetland
Reserve (Tan, 2013a).
Mayaheros urophthalmus × Parachromis managuensis
(Fig. 94)
References. Jaafar et al., 2012.
Distribution. Sungei Buloh Wetland Reserve (Jaafar et
al., 2012).
Mayaheros urophthalmus (Günther)
(Fig. 93, CA[SPC])

179
RAFFLES BULLETIN OF ZOOLOGY 2020
2012; Leong, 2012; Kwik et al., 2013; Ng PX & Tan, 2013;
Yeo, 2013; Tan, 2014a; Ng et al., 2015; Tan YK et al., 2015
(as Oreochromis sp.); Lim et al., 2016; Tan, 2016b, 2016c
(as Oreochromis sp.); Tan & Koh, 2016; Theng et al., 2016;
Tan et al., 2017b.
Distribution. Upper Seletar Reservoir (previously known
as Seletar Reservoir; Johnson, 1973a; Yang, 1984; Munro,
1990); Pulau Ubin (Subaraj, 1995); MacRitchie Reservoir
(Subaraj et al., 1995); Lower Peirce Reservoir (Lim et al.,
1995; Ng & Lim, 1997a); Central Catchment Nature Reserve
— Nee Soon Swamp Forest (Ng & Lim, 1997a); Bedok
Reservoir (Ng HH & Tan HH, 2010); Tekong Reservoir (Ng
HH & Tan HH, 2010); Marina Reservoir — encompassing
Singapore River, Rochor canal, Kallang River and Geylang
River (Tan HH et al., 2010); Singapore Botanic Gardens
(Yeo & Lim, 2010); Siglap canal (Leong, 2012); Pangsua
pond (Kwik et al., 2013); Woodlands Town Park pond (Kwik
et al., 2013); Yishun pond (Kwik et al., 2013); Punggol
Reservoir (Ng PX & Tan HH, 2013; Theng et al., 2016);
Serangoon Reservoir (Ng PX & Tan, 2013; Theng et al.,
2016); Bishan Park (Yeo, 2013; Tan, 2016c); West Coast
Park marsh pond (Tan, 2014a); Hindhede Quarry (Tan HH,
pers. obs., July 2014); Marina Reservoir (Tan YK et al.,
2015); Marina South (Tan & Koh, 2016); Sungei Buloh
Wetland Reserve (National Parks Board, 2003; Theng et
al., 2016); Kent Ridge Park ponds (Tan, 2016b); Tai Seng
area (Tan HH, pers. obs., 2016); Pulau Tekong (Lim et al.,
2016); Singapore Quarry (Tan et al., 2017b); Jurong Lake
(unpublished data).
Remarks. This species was introduced as a source of food
by the Japanese Army from Javan population during the
Japanese Occupation of Singapore (1942–1945) (Alfred,
1966a; Tham, 1973; Lim & Ng, 1990; Yeo & Chia, 2010).
It is now widespread in all Central Catchment reservoirs and
other articial water bodies and brackish water habitats (Ng
& Lim, 1997b; Baker & Lim, 2008, 2012). The juveniles of
this species are sometimes sold as live feed in the ornamental
sh trade (Tan HH, pers. obs.).
Fig. 94. Mayaheros urophthalmus × Parachromis managuensis
hybrid, ca. 200 mm SL, Sungei Buloh.
Fig. 95. Maylandia estherae, 115.0 mm SL, Pandan canal.
References. Liew et al., 2012.
Distribution. Pandan canal (Liew et al., 2012).
Remarks. This individual is probably a discarded pet sh.
Oreochromis aureus (Steindachner)
(AF)
References. Welcomme, 1988; Lim & Ng, 1990; Ng et al.,
1993; Lever, 1996; Ng & Lim, 1997b; Jaafar et al., 2012.
Distribution. Not known.
Remarks. This species is regarded as an escapee from
aquaculture (Ng & Lim, 1997b).
Oreochromis mossambicus (Peters)
(Fig. 96, AF[SPC])
References. Le Mare, 1949 (as Tilapia mossambica); Burkill,
1961 (as Tilapia); Johnson, 1964 (as Tilapia mossambica);
Alfred, 1966a (as Tilapia mossambicus, as early as 1944);
Johnson, 1973a, 1973b (as Tilapia); Tham, 1973 (as Tilapia
mossambica); Yang, 1984 (as Tilapia mossambicus); Lim &
Ng, 1990; Munro, 1990; Ng, 1991; Public Utilities Board,
1991 (as Tilapia mossambicus); Ng et al., 1993; Fernando
& Phang, 1994; Lim et al., 1995; Subaraj, 1995; Subaraj
et al., 1995; Lever, 1996; Ng & Lim, 1997a, 1997b; Ng &
Sivasothi, 1999; Goh et al., 2002; National Parks Board,
2003; Baker & Lim, 2008, 2012; Ng HH & Tan HH, 2010;
Tan HH et al., 2010; Tan HTW et al., 2010; Yeo & Chia,
2010; Yeo & Lim, 2010; Davison et al., 2012; Jaafar et al.,
Fig. 96. Oreochromis mossambicus, 150 mm SL male (top), 100
mm SL female (bottom); Pandan Reservoir.
Maylandia estherae (Konings)
(Fig. 95, AF[SPC])

180
Tan et al.: Non-native freshwater shes of Singapore
Distribution. Punggol Reservoir (Ng PX & Tan, 2013);
Serangoon Reservoir (Ng PX & Tan, 2013).
Remarks. This hybrid has morphological characters of
both parent species. Mature males tend to not exhibit the
blackened body, but still having the multiple black bars and
red margin over the caudal n (Tan HH, pers. obs.).
Parachromis managuensis (Günther)
(Fig. 99, CA[SPC])
Fig. 97. Oreochromis niloticus, 72.1 mm SL juvenile, Kranji Marsh.
Oreochromis niloticus (Linnaeus)
(Fig. 97, AF[SPC])
Fig. 98. Oreochromis mossambicus × O. niloticus hybrid, 195 mm
SL male (top), 170 mm SL female (bottom); Serangoon Reservoir.
Fig. 99. Parachromis managuensis, 130.0 mm SL, Pandan Reservoir.
References. Baker & Lim, 2008, 2012; Ng HH & Tan HH,
2010; Tan HTW et al., 2010; Kwik et al., 2013.
Distribution. Jurong Lake (Ng HH & Tan HH, 2010); Pandan
canal (Tan HH, pers. obs., 2010); Pangsua pond (Kwik et
al., 2013); Yishun pond (Kwik et al., 2013); Bishan Park
(unpublished data), Pandan Reservoir (unpublished data).
Remarks. This species is common in the ornamental sh
trade.
Pterophyllum scalare (Schultze, in Lichtenstein)
(Fig. 100, SA[SPC])
References. Ng PX & Tan, 2013.
Fig. 100. Pterophyllum scalare, 55.0 mm SL, Pangsua Pond.
References. Liew et al., 2012.
Distribution. Pangsua Pond (Liew et al., 2012).
References. Lim & Ng, 1990; Ng et al., 1993; Ng & Lim,
1997b; National Parks Board, 2003; Ng HH & Tan HH,
2010; Jaafar et al., 2012; Kwik et al., 2013; Ng PX & Tan
HH, 2013; Tan, 2013a; Tan et al., 2016.
Distribution. Sungei Buloh Wetland Reserve (National Parks
Board, 2003; Tan, 2013a); Jurong Lake (Ng HH & Tan HH,
2010); Murai Reservoir (Ng HH & Tan HH, 2010); Sarimbun
Reservoir (Ng HH & Tan HH, 2010); Tengah Reservoir (Ng
HH & Tan HH, 2010); Upper Peirce Reservoir (Ng HH &
Tan HH, 2010); Lower Peirce Reservoir (Ng & Lim, 1997b);
Yishun pond (Kwik et al., 2013); Punggol Reservoir (Ng PX
& Tan, 2013); Serangoon Reservoir (Ng PX & Tan, 2013);
Kranji Marsh (Tan et al., 2016); Bishan Park (unpublished
data); Marina Reservoir (unpublished data).
Remarks. Both O. niloticus and O. mossambicus can adapt
to saline conditions.
Oreochromis mossambicus × O. niloticus
(Fig. 98 [SPC])

181
RAFFLES BULLETIN OF ZOOLOGY 2020
Remarks. The specimens obtained from the listed site
probably originated as discarded pets from the ornamental
sh trade.
Satanoperca jurupari (Heckel)
(Fig. 101, SA[SPC])
References. Baker & Lim, 2008, 2012 (as Paratheraps
synspilus); Ng HH & Tan HH, 2010 (as Veija synspila);
Tan HTW et al., 2010 (as Paratheraps synspila); Kwik et
al., 2013 (as Paratheraps synspilum); Ng et al., 2015 (as
Paraneetroplus melanurus); Tan, 2015b, 2016c.
Distribution. Jurong Lake (Ng HH & Tan HH, 2010);
Tengah Reservoir (Ng HH & Tan HH, 2010); Pandan canal
(Tan HH, pers. obs., 2010); Pangsua Pond (Kwik et al.,
2013); Yishun Pond (Kwik et al., 2013); Bishan Park (Tan,
2015b, 2016c); Marina Reservoir (near Marina Bay Sands;
Wozniak S, pers. obs., October 2016); Singapore Botanic
Gardens (Tan HH, pers. obs., 2002).
Remarks. This is likely one of the parent species for the
Luohan cichlid.
Luohan/Flowerhorn cichlid
(Fig. 104 [SPC])
Fig. 101. Satanoperca jurupari, ca. 150 mm SL, Kranji Reservoir.
References. Ng HH & Tan HH, 2010; Baker & Lim, 2012.
Distribution. Kranji Reservoir (Ng HH & Tan HH, 2010).
Thorichthys meeki Brind
(Fig. 102, CA)
Fig. 102. Thorichthys meeki, 63.9 mm SL, trade material.
References. Ng et al., 1993 (as Cichlasoma meeki); Ng &
Lim, 1997a, 1997b.
Distribution. Cluny Road drain (Ng & Lim, 1997a, 1997b).
Remarks. This species is not established, and it is regarded
as an escapee.
Vieja melanura (Günther)
(Fig. 103, CA[SPC])
Fig. 103. Vieja melanura, 180 mm SL, Pandan canal.
Fig. 104. Luohan/Flowerhorn cichlid hybrid, ca. 200 mm SL,
Pandan Reservoir.
References. Baker & Lim, 2008, 2012 (as Cichlasoma
hybrid [Luohan]); Ng HH & Tan HH, 2010 (as Cichlasoma
hybrid); Jaafar et al., 2012 (as Veija spp.); Kwik et al., 2013
(as Amphilophus × Paratheraps); Ng PX & Tan, 2013 (as
Cichlasoma sp. [Luohan]).
Distribution. Khatib Bongsu (Tan HH, pers. obs., 2004);
Pandan river (Tan HH, pers. obs., 2004); Jurong Lake (Ng
HH & Tan HH, 2010); Yishun Pond (Kwik et al., 2013);
Punggol Reservoir (Ng PX & Tan, 2013); Serangoon
Reservoir (Ng PX & Tan, 2013); Hindhede Quarry (Tan
HH, pers. obs., 2014); Singapore Botanic Gardens (Tan
HH, pers. obs., 2015).
Remarks. This multiple-genera hybrid sh was extremely
popular in the mid-2000s but it has now fallen out of
favour. At the height of the craze, lucrative sales prompted
the opening of many boutique shops that sold only Luohan
cichlids, as well as aquarium shops that opened 24 hours
(Tan HH, pers. obs.).

182
Tan et al.: Non-native freshwater shes of Singapore
Blood/Red Parrot hybrid
(Fig. 105 [SPC])
Family Gobiidae
Brachygobius sabanus Inger
(Fig. 107, SEA[SPC])
Fig. 105. Blood/Red Parrot hybrid, 88.0 mm SL, Pangsua Pond.
References. Liew et al., 2012 (as Amphilophus citrinellus ×
Paraneetroplus melanurus); Kwik et al., 2013 (as Cichlasoma
hybrid).
Distribution. Pangsua Pond (Liew et al., 2012); Yishun
Pond (Kwik et al., 2013).
Remarks. This mutated hybrid continues to be an extremely
popular aquarium sh despite its articially-induced physical
defects (Tan HH, pers. obs.).
Red Tilapia
(Fig. 106 [SPC])
Fig. 106. Red Tilapia, 132.0 mm SL, Pangsua Pond.
References. Lim & Ng, 1990; Munro, 1990; Lim & Ng,
1992 (as Oreochromis spp. Hybrid); Goh et al., 2002 (as
Tilapia hybrid); Tan, 2014a (as Oreochromis hybrid); Tan
& Koh, 2017 (as Oreochromis sp.).
Distribution. Pulau Ubin HDB quarry (Lim & Ng, 1992);
West Coast Park marsh pond (Tan, 2014a), Sungei Buloh
Wetland Reserve (Tan & Koh, 2017); Yew Tee canal (Tan
HH, pers. obs., 2016); Bishan Park (Andre, pers. obs.,
October 2016).
Remarks. This supposedly sterile hybrid of three species is
brightly coloured (pink, orange or platinum) and frequently
used for aquaculture (Lim & Ng, 1990; Tan & Koh, 2017).
The juvenile individuals of this species are commonly sold as
live feed in the ornamental sh trade (Tan HH, pers. obs.).
References. Baker & Lim, 2012; Lim & Tan, 2012; Tan et
al., 2014; Kwik & Yeo, 2015 (as Brachygobius doriae); Ho
et al., 2016; Larson et al., 2016; Tan et al., 2016.
Distribution. Poyan Reservoir (Lim & Tan, 2012); Tengah
Reservoir (Lim & Tan, 2012); Kranji Reservoir (Lim & Tan,
2012); Kranji Marsh (Lim & Tan, 2012; Tan et al., 2016);
Sungei Buloh Wetland Reserve (Lim & Tan, 2012); Banir
(Lim & Tan, 2012); Upper Seletar Reservoir (Tan et al.,
2014); Sungei Seletar (Ho et al., 2016).
Remarks. This species is well-established in Singapore
(Larson et al., 2016).
Rhinogobius similis Gill
(Fig. 108, EAs[SPC])
Fig. 107. Brachygobius sabanus, 12.7 mm SL, Upper Seletar
Reservoir inlet.
Fig. 108. Rhinogobius similis, 34.8 mm SL male, Upper Seletar
Reservoir spillway.
References. Alfred, 1966a (as Stigmatogobius poicilosoma);
Johnson, 1973a, 1973b (as Stigmatogobius poicilosoma);
Lim & Ng, 1990 (as Stigmatogobius poecilosoma);
Munro, 1990 (as Stigmatogobius poicilosoma); Ng,
1991 (as Stigmatogobius poecilosoma); Lim et al., 1995
(as Rhinogobius cf. giurinus); Subaraj et al., 1995 (as
Rhinogobius cf. giurinus); Ng & Lim, 1997a, 1997b (as
Rhinogobius giurinus); Larson & Lim, 2005 (as Rhinogobius
giurinus); Larson et al., 2008 (as Rhinogobius giurinus);
Baker & Lim, 2008, 2012 (as Rhinogobius giurinus); Ng
HH & Tan HH, 2010 (as Rhinogobius giurinus); Tan &
Lim, 2011 (as Rhinogobius giurinus); Lim et al., 2013 (as
Rhinogobius giurinus); Tan, 2014e, 2014g (as Rhinogobius
giurinus); Kwik & Yeo, 2015 (as Rhinogobius giurinus); Ho
et al., 2016 (as Rhinogobius giurinus); Larson et al., 2016
(as Rhinogobius giurinus); Li et al., 2016 (as Rhinogobius
giurinus).
Distribution. MacRitchie Reservoir (Munro, 1990); Upper
Seletar Reservoir (Munro, 1990; Larson & Lim, 2005; Ng &

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RAFFLES BULLETIN OF ZOOLOGY 2020
Lim, 1997b; Ng HH & Tan HH, 2010; Tan, 2014e, 2014g);
Banir (Munro, 1990; Tan, 2014g); MacRitchie Reservoir
(Subaraj et al., 1995); Central Catchment Nature Reserve
(Ng & Lim, 1997a); Jurong Road area (Ng & Lim, 1997a);
Singapore Botanic Gardens (Ng & Lim, 1997a); Jurong
Lake (Ng HH & Tan HH, 2010); Upper Peirce Reservoir
(Ng HH & Tan HH, 2010); Lower Peirce Reservoir (Lim
et al., 1995; Ng & Lim, 1997b; Ng HH & Tan HH, 2010);
Lower Seletar Reservoir (Ng HH & Tan HH, 2010); Murai
Reservoir (Ng HH & Tan HH, 2010); Poyan Reservoir (Ng
HH & Tan HH, 2010); Tengah Reservoir (Ng HH & Tan
HH, 2010); Banir (Tan & Lim, 2011); Kranji Marsh (Lim et
al., 2013); Central Catchment Nature Reserve — Nee Soon
Swamp Forest (Ho et al., 2016; Li et al., 2016); Bishan Park
(unpublished data).
Remarks. Alfred’s (1966a) material of Stigmatogobius
poicilosoma consisted of three taxa, viz., Eugnathogobius
siamensis, Pseudogobiopsis oligactis (recently rediscovered
in Singapore; Tan & Lim, 2011) and Rhinogobius similis
(Ng & Lim, 1997b; Larson et al., 2008, 2016). It is a
well-established non-native species in Singapore that was
accidentally introduced with sh fry imported from China
(Larson et al., 2016).
The taxonomic status of R. giurinus was resolved recently
and is considered junior to R. similis (Suzuki et al., 2015).
Family Helostomatidae
Helostoma temminckii Cuvier
(Fig. 109, SEA)
Fig. 109. Helostoma temminckii, ca. 80 mm SL, trade material.
References. Ng et al., 1993
Distribution. No detailed location provided.
Family Osphronemidae
Betta splendens Regan
(Fig. 110, SEA[SPC])
References. Cantor, 1849 (as Macropodus pugnax); Herre,
1940; Tweedie, 1952; Johnson, 1964; Johnson, 1973a; Lim
& Ng, 1990; Munro, 1990; Ng et al., 1993 (as B. imbellis);
Lever, 1996; Ng & Lim, 1997b; National Parks Board, 2003;
Tan HH et al., 2010; Tan, 2016b.
Fig. 110. Betta splendens, ca. 30 mm SL male, trade material.
Fig. 111. Macropodus opercularis, 45.5 mm SL male, trade material.
References. Tan et al., 2013.
Distribution. Bukit Brown (Tan et al., 2013).
Remarks. This species is sometimes sold as live feed in the
ornamental sh trade (Tan HH, pers. obs.).
Osphronemus goramy Lacepède
(Fig. 112, SEA[SPC])
References. Green, 1928 (as Osphronemus olfax); Smith,
1933 (introduced as early as 1898); Herre & Myers, 1937;
Tweedie, 1940; Alfred, 1966a; Johnson, 1973a; Anonymous,
1988a; Lim & Ng, 1990; Munro, 1990; Ng et al., 1993; Ng
& Lim, 1997a, 1997b; National Parks Board, 2003; Ng HH
& Tan HH, 2010; Chua, 2015; Tan, 2017b.
Distribution. This species had been sighted by Cantor (1849)
and subsequently obtained by Herre in 1940 from Mandai
area; Sembawang-Nee Soon area (Johnson, 1964, 1973a);
Jurong Lake (Johnson, 1973a); Sungei Buloh Wetland
Reserve (National Parks Board, 2003); Kallang River (now
part of Marina Reservoir; Munro, 1990; Tan HH et al.,
2010); Kent Ridge Park ponds (Tan, 2016b); Bukit Batok
(Ng DJJ, pers. obs., 2018).
Remarks. This is a very popular ornamental sh species
ubiquitous in aquarium shops, thus its presence in Singapore’s
water bodies is not surprising. Recently, this species is
commonly sold as live feed in the ornamental sh trade,
the culled sh being a by-product from selective breeding
(Tan HH, pers. obs.).
Macropodus opercularis (Linnaeus)
(Fig. 111, EAs[SPC])

184
Tan et al.: Non-native freshwater shes of Singapore
Distribution. Herre & Myers (1937) collected this species
from Singapore but did not provide location details.
MacRitchie Reservoir (Alfred, 1966a; Anonymous, 1988a;
Ng & Lim, 1997a, 1997b; Ng HH & Tan HH, 2010); Sungei
Buloh Wetland Reserve (National Parks Board, 2003); Bedok
Reservoir (Ng HH & Tan HH, 2010); Kranji Reservoir (Ng
HH & Tan HH, 2010); Upper Peirce Reservoir (Ng HH &
Tan HH, 2010); Lower Peirce Reservoir (Ng HH & Tan HH,
2010); Lower Seletar Reservoir (Ng HH & Tan HH, 2010);
Singapore Botanic Gardens (Tan HH, pers. obs., 2015);
Sungei Seletar (Tan, 2017b); Jurong Lake (unpublished data);
Bedok Reservoir (unpublished data); Serangoon Reservoir
(unpublished data).
Remarks. This species is of economic importance as food sh
in the Southeast Asian region, but not popular in Singapore.
Xanthic and albinio varieties are popular in the ornamental
sh trade; some individuals are even tattooed with numbers
or coloured patterns on the bodies (Tan HH, pers. obs.).
Osphronemus laticlavius Roberts
(Fig. 113, SEA)
Sphaerichthys osphromenoides Canestrini
(Fig. 114, SEA)
Fig. 112. Osphronemus goramy, 142 mm SL, Serangoon Reservoir.
Fig. 113. Osphronemus laticlavius, 72.9 mm SL, trade material.
References. Lim, 2013b.
Distribution. Singapore Quarry (Lim, 2013b).
Remarks. This is a popular species in the ornamental sh
trade as it is considered lucky or good for “feng shui” (Lim
& Ng, 1990).
Fig. 114. Sphaerichthys osphromenoides, 32.0 mm SL, Malaysia.
References. Boeseman, 1957; Ng & Lim, 1997b.
Distribution. Not known.
Remarks. These records are of individuals that are likely
from the ornamental sh trade, not established (Lim & Ng,
1997b).
Trichogaster lalius (Hamilton)
(Fig. 115, SAs)
Fig. 115. Trichogaster lalius, 43.1 mm SL male, trade material.
References. Lim & Ng, 1990 (as Colisa lalia); Ng et al.,
1993 (as Colisa lalia).
Distribution. Not known.
Remarks. This species is probably from ornamental sh
trade, and not established (Lim & Ng, 1990). The females of
this species are sometimes sold as live feed in the ornamental
sh trade (Tan HH, pers. obs.).
Trichopodus microlepis (Günther)
(Fig. 116, SEA)
References. Anonymous, 1988a (as Trichogaster microlepis);
Lim & Ng, 1990 (as Trichogaster microlepis); Ng et al.,
1993 (as Trichogaster microlepis); Ng & Lim, 1997a (as
Trichogaster microlepis); Ng & Lim, 1997b (as Trichogaster
microlepis); Low & Lim, 2012.
Distribution. Mandai area (Anonymous, 1988a; Ng & Lim,
1997a); Jurong area (Ng & Lim, 1997a).

185
RAFFLES BULLETIN OF ZOOLOGY 2020
Trichopodus pectoralis Regan
(Fig. 117, SEA[SPC])
Ng & Lim, 1992; Ng & Lim, 1997a, 1997b); Upper Seletar
Reservoir (Anonymous, 1988a; Ng & Lim, 1997a, 1997b);
Mandai area (Anonymous, 1988b; Ng & Lim, 1997a);
Sungei Buloh Wetland Reserve (Lim & Ng, 1992; Ng &
Lim, 1997a; National Parks Board, 2003); Kallang River
(now part of Marina Reservoir; Tan HH et al., 2010); Kranji
Marsh (Tan et al., 2016).
Remarks. Regan (1910) described Trichopodus pectoralis
based on a series of six specimens from Thailand (Siam) and
Singapore, but did not provide more precise locations. This
species had apparently been introduced into Singapore in
1898 for aquaculture (Smith, 1933), and is presently sold in
local markets, salted and without heads (Tan HH, pers. obs.).
Trichopsis schalleri Ladiges
(Fig. 118, SEA[SPC])
Fig. 116. Trichopodus microlepis, 92.6 mm SL, trade material.
Fig. 117. Trichopodus pectoralis, 96.6 mm SL, Sungei Buloh.
References. Hanitsch, 1901, 1904 (as Osphromenus
siamensis); Regan, 1910 (described as Trichogaster
pectoralis); Smith, 1933 (as Trichogaster pectoralis); Herre
& Myers, 1937 (as Trichogaster pectoralis); Fowler, 1938;
Tweedie, 1952 (as Trichogaster pectoralis); Alfred, 1961,
1966a (as Trichogaster pectoralis, introduced in 1898);
Johnson, 1964 (as Trichogaster pectoralis); Johnson, 1973a
(as Trichogaster pectoralis); Tan et al., 1987 (as Trichogaster
pectoralis); Anonymous, 1988a, 1988b (as Trichogaster
pectoralis); Chou & Lam, 1989 (as Trichogaster pectoralis);
Lim & Ng, 1990, 1991, 1992 (as Trichogaster pectoralis);
Munro, 1990 (as Trichogaster pectoralis); Lim, 1991 (as
Trichogaster pectoralis); Ng & Lim, 1992 (as Trichogaster
pectoralis); Ng et al., 1993 (as Trichogaster pectoralis);
Lever, 1996 (as Trichogaster pectoralis); Ng & Lim,
1997a (as Trichogaster pectoralis); Ng & Lim, 1997b (as
Trichogaster pectoralis); National Parks Board, 2003; Yeo
& Chia, 2010 (as Trichogaster pectoralis); Tan HH et al.,
2010; Low & Lim, 2012; Tan et al., 2016.
Distribution. Herre & Myers (1937) obtained 14 specimens
from Singapore but did not provide location details; Jurong
and Kranji Rivers (Alfred, 1961); Lower Peirce Reservoir
(Alfred, 1966a); Singapore Botanic Gardens (Anonymous,
1988a); Seletar area (Munro, 1990); Central Catchment
Nature Reserve — Nee Soon Swamp Forest (Lim, 1991;
Fig. 118. Trichopsis schalleri, ca. 35.0 mm SL, Thailand.
References. Ng HH & Tan HH, 2010.
Distribution. Lower Seletar Reservoir (Ng HH & Tan HH,
2010).
Remarks. The source of introduction is not known as this
is a rarely-encountered species in the ornamental sh trade
(Tan HH, pers. obs.).
Family Channidae
Channa micropeltes (Cuvier, in Cuvier &
Valenciennes)
(Fig. 119A, B, SEA[SPC])
Fig. 119. Channa micropeltes, ca. 30 mm SL juvenile (top), trade
material; 435 mm SL adult (bottom), Bedok Reservoir.

186
Tan et al.: Non-native freshwater shes of Singapore
References. Burkill, 1961, 1963; Alfred, 1966a; Yang, 1984;
Anonymous, 1988c, 1989a, 1989b; Lim & Ng, 1990; Munro,
1990; Lim, 1991; Ng, 1991; Public Utilities Board, 1991;
Lim & Ng, 1992; Ng et al., 1993; Lim et al., 1995; Subaraj
et al., 1995; Ng & Lim, 1997a, 1997b; National Parks Board,
2003; Baker & Lim, 2008, 2012; Ng HH & Tan HH, 2010;
Ho et al., 2016; Tan, 2016b; Tan & Herder, 2016.
Distribution. Present in all reservoirs (Ng & Lim, 1997b)
and widespread in ponds (Baker & Lim, 2008, 2012).
Singapore Botanic Gardens (Burkill, 1963; Alfred, 1966a;
Ng & Lim, 1997a); Sime Road forest stream (Anonymous,
1988c); Woodlands Town Garden (Anonymous, 1989a);
Chestnut forest (Subaraj et al., 1995); Bedok Reservoir (Ng
HH & Tan HH, 2010); Jurong Lake (Ng HH & Tan HH,
2010); Kranji Reservoir (Lim, 1991; Ng & Lim, 1997a,
1997b; Ng HH & Tan HH, 2010); Upper Peirce Reservoir
(Anonymous, 1989b; Ng & Lim, 1997a, 1997b; Ng HH &
Tan HH, 2010); Lower Peirce Reservoir (Lim et al., 1995;
Ng & Lim, 1997a, 1997b; Ng HH & Tan HH, 2010); Upper
Seletar Reservoir (Yang, 1984; Ng & Lim, 1997a, 1997b;
Ng HH & Tan HH, 2010); Lower Seletar Reservoir (Ng &
Lim, 1997a, 1997b; Ng HH & Tan HH, 2010); MacRitchie
Reservoir (Subaraj et al., 1995; Ng & Lim, 1997a, 1997b;
Ng HH & Tan HH, 2010); Kranji Reservoir (Ng & Lim,
1997a); Murai Reservoir (Ng HH & Tan HH, 2010); Pandan
Reservoir (Ng HH & Tan HH, 2010); Poyan Reservoir (Ng
HH & Tan HH, 2010); Sarimbun Reservoir (Ng HH & Tan
HH, 2010); Tengah Reservoir (Ng & Lim, 1997a, 1997b;
Ng HH & Tan HH, 2010); Sungei Buloh Wetland Reserve
(Ng & Lim, 1997a; National Parks Board, 2003; Chua, 2015;
Tan & Herder, 2016; Nee Soon Swamp Forest (Lim & Ng,
1992; Ho et al., 2016); Kent Ridge Park ponds (Tan, 2016b);
Yew Tee canal (Tan HH, pers. obs., 2016).
Remarks. Channa micropeltes was rst released into Seletar
Reservoir (now known as Upper Seletar Reservoir) to
control the abundant tilapia (Oreochromis sp.) population,
and subsequently introduced into other reservoirs with
large populations of tilapia (Yang, 1984). The juvenile
individuals of this species are commonly sold as live feed
in the ornamental sh trade (Tan HH, pers. obs.).
DISCUSSION
Non-native freshwater sh records over time. The rst
published records of non-native sh species in Singapore
was in 1849. Since then, the number of non-native sh
species recorded has increased exponentially in recent years
(Fig. 120).
One of the main reasons for the recent increase in reports of
non-native shes is the increase in sh survey reports, and the
growing popularity of citizen science, and readily-accessible
online journals (e.g., Nature in Singapore: http://lkcnhm.
nus.edu.sg/nus/index.php/nhmpublications/nis; Singapore
Biodiversity Records: http://lkcnhm.nus.edu.sg/nus/index.
php/nhmpublications/singapore-biodiversity-records), and
nature blogs (e.g., Bird Ecology Study Group).
Reports of non-native sh species (established as well as
non-established) were from throughout Singapore (Fig. 121).
Notably, reservoirs were focal points of species reports. This
is consistent with known associations between modied
habitats and non-native fish establishment (Liew et al.,
2016a; Liew et al., 2018a), especially where native species
diversity is poor (Liew et al., 2016b). Recent ndings suggest
that non-native species form predator-prey interactions with
native habitat generalists (e.g., Channa striata) in complex
food webs (Liew et al., 2018b). In addition, some non-native
species (e.g., cichlids, the dominant group of established
freshwater sh) appear to be particularly well-suited or
adaptable ecologically to Singapore’s freshwater habitats
(Kwik et al., in press).
Unsurprisingly, sh species from tropical regions form the
majority of introductions (see Fig. 122; Central America
[15] + South America [23] + Africa [9] + South Asia [8]
+ Southeast Asia [46] + Australasia [1] = 102 species;
82.9%). Of the regions, Southeast Asia is the main source of
non-native sh (46 species, 37.3%). This somewhat reects
the primary sources of ornamental sh species traded in
Singapore.
Species for biological control. Hanitsch (1912) reported
on the feasibility of introducing Poecilia reticulata (listed
as Girardinus poeciloides) as a vector control agent against
the mosquito-borne disease malaria, which was prevalent
in Singapore at that time. He also surveyed for potential
local aquatic agents of vector control, and listed several
species of freshwater shes with potential — Aplocheilus
armatus (listed as Haplocheilus panchax; presently the
Sundaic population has been re-identied by Katwate et
al., 2018), Dermogenys collettei (listed as Heiramphus
uviatilis), Betta pugnax, Trichopodus trichopterus (listed
as Osphronemus trichopterus), Anabas testudineus (listed as
Anabas scandens), Channa striata (listed as Ophiocephalus
striatus) and Clarias batrachus (listed as Clarias magur).
He cautioned about the risk of accidental introduction of
other undesirable organisms. However, Poecilia reticulata
was still introduced shortly after as a vector control against
malaria (Lim & Ng, 1990: 75).
Fig. 120. Number of rst records of non-native sh species from
1849 to 2016.

187
RAFFLES BULLETIN OF ZOOLOGY 2020
This diminutive and very prolic species has since colonised
many of Singapore’s disturbed freshwater bodies (including
at forest edges) since its introduction as a biological control;
populations are present even in back mangrove habitats
(Tan HH, pers. obs.). This species is very tolerant and can
survive in polluted canals in industrial estates, and even
sewerage tanks (Lim & Ng, 1990). This species has been
an integral part of Singapore’s ornamental sh trade, being
part of the multi-million dollar aquarium/ornamental sh
industry (Fernando & Phang, 1994). Fishes are subject
to stringent selective breeding with numerous fancy and
colourful variants. The rejected or culled shes become a
source of live feed (Lim & Ng, 1990) for large carnivorous
ornamental sh species.
Gambusia afnis is another species introduced for biological
control of mosquitoes (Lim & Ng, 1990). This small species
is also ubiquitous and can also tolerate brackish conditions
(Tan HH, pers. obs.).
Channa micropeltes was introduced into reservoirs to control
the large populations of Oreochromis spp. present in the
1970s–1980s (Yang SL, pers. comm.; Ng HH & Tan HH,
2010). This species quickly established and became widely
distributed in all articial water bodies in Singapore. This
is a highly coveted food sh, and is a sought-after target for
recreational angling, including illegal angling in prohibited
areas.
Dubious records. One of the earliest lists of ornamental
shes imported from Singapore into Europe was documented
by Boeseman (1957). In his list, he included species
previously not recorded from Singapore, viz., Kryptopterus
vitreolus (previously known as K. bicirrhus; a species
known from southern and eastern Thailand; see Ng &
Kottelat, 2013a), Chromobotia macracanthus (identied as
Botia macaracanthus; a species known only from Sumatra
and Borneo; see Tan & Kottelat, 2009, and Kottelat et al.,
1993), Trigonopoma pauciperforatum (identied as Rasbora
pauciperforata; a species known from from Peninsular
Malaysia, Borneo and Sumatra; see Kottelat et al., 1993),
Trigonopoma gracile (identified as Rasbora taeniata; a
species known from Malaysia, Borneo and Sumatra; see
Kottelat et al., 1993), Puntigrus partipentazona (identied
as Puntius partipentazona; a species known from Peninsular
Malaysia and Thailand; see Mohsin & Ambak, 1983),
Sphaerichthys osphromenoides (a species known from
peat swamps in Peninsular Malaysia, Sumatra and western
Borneo; see Kottelat et al., 1993, and Tan & Ng, 2005) and
Gymnochanda lamentosa (described as a new genus and
new species, but later found to be a homonym of an earlier
identical name given by Fraser-Brunner in 1955; apparently
Fig. 121. Distribution of non-native sh species (established as well as non-established) records from 1849 to 2016 (n = 648). Intensity
of orange hues represents species record frequency, in which darker shades correspond to higher occurrence records. Species records
were location- and species-specic, which is to say that multiple reports of the same species from a single location is represented only
once in the heat map.

188
Tan et al.: Non-native freshwater shes of Singapore
found only in Peninsular Malaysia and Sumatra; see Tan &
Lim, 2014). In 1957, the ornamental sh trade was already
an important industry in Singapore, and many of these
species were imported from neighbouring countries and
territories, including Peninsular Malaysia, southern Thailand
and Sumatra. Alfred (1966a) commented that Boeseman’s
sh list is almost entirely based upon sh imports for the
ornamental sh trade and thus was not used in his own
checklist.
Native sh species. The number of extant native freshwater
sh species recorded from Singapore is 42 (see Table 2).
Of these, 22 species (52.4%) are restricted to forest streams
within the Central Catchment and Bukit Timah Nature
Reserves.
At least 11 native sh species are at present known to be
extirpated; viz., Cyprinidae — Barbodes dunckeri (with
known specimen deposition), Osteochilus spilurus, Rasbora
cephalotaenia, Rasbora paucisqualis; Cobitidae — Pangio
semicincta (with known specimen deposition); Siluridae —
Ompok leiacanthus; Sisoridae — Gyptothorax callopterus
(with known specimen deposition); Mastacembelidae
— Mastacembelus favus; Pristolepididae — Pristolepis
fasciatus; and Osphronemidae — Belontia hasseltii, Betta
tomi (with known specimen deposition) (Alfred, 1966a;
Lim & Ng, 1990). Some of these species are listed based
on personal opinion (of the rst author) and many years of
ichthyological eld work in Singapore and the Southeast
Asian region.
Prior to the comprehensive list of freshwater fishes of
Singapore by Alfred (1966a), there were very few published
records from the island city state. Earlier records were based
mainly on the works of Cantor (1849), Duncker (1904),
Regan (1910), Tweedie (1936, 1940, 1950, 1952), Herre
& Myers (1937), Herre (1937, 1940), Fowler (1938), and
Fraser-Brunner (1940). These works generally encompassed
a larger geographic area which included sporadic collections
made in Singapore.
Alfred’s (1966a) publication was based on his earlier works
conducted from 1960. However, even by then, much of
Singapore’s natural landscape and habitats had been altered,
due to urbanisation and cultivation of cash crops (e.g.,
gambier). Corner (1978) documented the demise of many
swamp forest habitats in southern Johor and Singapore
in pre-Second World War era, although the account was
only published much later. For Singapore, Corner (1978)
documented the clear felling of swamp forest habitats
mainly from Mandai (present day Upper and Lower Seletar
Reservoirs and residential estates), Jurong (present day
Jurong Lake, Chinese and Japanese Gardens, industrial and
residential estates), and Ang Mo Kio (present day industrial
and residential areas). No documentation of the then aquatic
inhabitants was made. Thus, many indigenous species, and
knowledge of these species, were lost. Some of the earlier
records by Herre & Myers (1937) and Herre (1940) included
some of these localities, however these reports were not
comprehensive but gave positive correlations with acid-water
swamp forest habitats (e.g., Silurichthys hasseltii).
Ornamental sh trade. Prior to 1950s, Singapore was
already a hub for the ornamental sh trade (Boeseman,
1957). To date, more than 760 species from 89 families
have been recorded from the ornamental sh trade based in
Singapore; these are imported from all over the world (Lee,
2008, unpublished thesis). Many of the tropical shes are
potentially invasive species. Many species in cultivation were
inadvertently introduced into the water bodies of Singapore,
either as escapees or intentional releases (Fernando & Phang,
1994).There is an ongoing trend for keeping monster or
Fig. 122. World map showing region of origin for the 116 non-native freshwater sh species recorded in Singapore (excluding seven hybrids).

189
RAFFLES BULLETIN OF ZOOLOGY 2020
Table 2. Extant native inland freshwater shes of Singapore. This is a current listing, which comprises 42 species from 19 families and 8
orders (data from Baker & Lim, 2012; and unpublished data). Names in bold represent species restricted to forest streams within protected
areas like the Central Catchment Nature Reserve.
Family Species
MEGALOPIDAE Megalops cyprinoides
CYPRINIDAE Barbodes banksi
Barbodes lateristriga
Boraras maculatus
Cyclocheilichthys apogon
Desmopuntius hexazona
Rasbora einthovenii
Rasbora elegans
Trigonostigma heteromorpha
COBITIDAE Pangio muraeniformis
NEMACHEILIDAE Nemacheilus selangoricus
BAGRIDAE Mystus gulio
Pseudomystus leiacanthus
SILURIDAE Silurichthys hasseltii
AKYSIDAE Parakysis longirostris
CLARIIDAE Clarias batrachus
Clarias leiacanthus
Clarias nieuhoi
SYNBRANCHIDAE Monopterus javanensis
MASTACEMBELIDAE Macrognathus maculatus
tank-buster sh species (INJAF, 2018); these species can
potentially grow to large adult sizes, some even larger than
humans. These species are usually obtained during the
juvenile stage, but rapidly outgrow hobbyists’ limited tank
sizes. This trend is a cause for concern as these monster
species are high-potential candidates for release. Disposal
of adult or sub-adult shes will eventually become an issue
because there are limited options for adoption or public
displays due to their large adult size. Some of these shes have
been recorded in this present study and include Potamotrygon
motoro, Atractosteus spatula, Chitala ornata, Arapaima
gigas, Osteoglossum bicirrhosum, Scleropages spp.,
Hampala macrolepidota, Leptobarbus rubripinna, Morulius
chrysophekadion, Chromobotia macracanthus, Piaractus
brachypomus, Hemibagrus capitulum, Clarias gariepinus,
Pterygoplichthys spp., Pangasionodon hypophthalmus,
Pangasius sanitwongsei, Phractocephalus hemioliopterus,
Datnioides microlepis, Cichla spp., Osphronemus goramy,
and Channa micropeltes. Some of these species are known
to be harmful to humans due to venomous spines, e.g.,
Potamotrygon motoro (Ng et al., 2009); or aggressive
nature, and large adult sizes, e.g., Channa micropeltes (Ng
& Lim, 1990).
Currently, not many freshwater sh species are commercially
bred in Singapore due to space constraints and cost. The few
which are mainly consisting of fast growing and easily bred
species, e.g., Poeciliidae (see Fernando & Phang, 1994).
Food fish aquaculture. Alfred (1966b) provided a
bibliography of 291 entries pertaining to Malayan freshwater
fisheries; this work provided insights to the range of
publications and resources available during Singapore’s
colonial history (1849–1964).
Birtwistle (1931a) provided a detailed account on the
cultivation of five species of Chinese carps (Cirrhinus
molitorella, Ctenopharyngodon idella, Cyprinus carpio,
Hypophthalmichthys molitrix and H. nobilis) in mud ponds
in Malaya (which then included Singapore). The fry of the
abovementioned species was imported from overseas (China,
Hong Kong and Taiwan) and subsequently placed into mud
ponds for the growing-out phase. Maxwell (1921), Stead
(1923) and Birtwistle (1931b) gave interesting accounts on
how fertilised ova and sh fry were shipped from China in
large jars of freshwater, and how the water and contents were
vigorously stirred for aeration. Le Mare (1949) provided an
additional mode of transportation of fry in tin cans during
the North-East monsoon by air freight. Tham (1973) also
provided a brief account of the above practice, while noting
that H. molitrix and H. nobilis were subsequently articially
bred in Malaysia and the fry then exported to Singapore.
Family Species
ADRIANICHTHYIDAE Oryzias javanicus
APLOCHEILIDAE Aplocheilus armatus
ZENARCHOPTERIDAE Dermogenys collettei
Hemirhamphodon pogonognathus
ELEOTRIDAE Oxyeleotris marmorata
Oxyeleotris urophthalmus
GOBINELLIDAE Eugnathogobius siamensis
Glossogobius aureus
Gobiopterus brachypterus
Mugilogobius chulae
Pseudogobiopsis oligactis
NANDIDAE Nandus nebulosus
ANABANTIDAE Anabas testudineus
OSPHRONEMIDAE Betta imbellis
Betta pugnax
Luciocephalus pulcher
Trichopodus trichopterus
Trichopsis vittata
CHANNIDAE Channa gachua
Channa lucius
Channa melasoma
Channa striata

190
Tan et al.: Non-native freshwater shes of Singapore
Barbonymus gonionotus (Lampam jawa) was also introduced
into Malaya from Indonesia during the Japanese Occupation
(1942–1945) (Mohsin & Ambak, 1983). It was a very popular
food sh with the local Malay community and was widely
cultivated.
Tilapia (Oreochromis sp.) was introduced into Malaya from
Indonesia by the Japanese army during the latter part of the
Japanese Occupation, ca. 1941–1945 (Le Mare, 1949; Tham,
1973; Mohsin & Ambak, 1983). However, Costa-Pierce
(2003) indicates that the introduction could date earlier to
1938. During the 1950s, it was a very popular food sh
cultured in ponds and mining pools throughout Malaya.
At present, tilapia remains a staple food sh in Singapore;
available live mainly from Peninsular Malaysia, as fresh
whole sh, or as pre-packed llets.
A recent import for the live food sh trade is Monopterus
cuchia (Synbranchidae) from India, which is usually
sold in some supermarkets (Tan HH, pers. obs.). Each
individual can weigh over a kilogram and is an esteemed
food sh in northern Chinese cuisine restaurants. There is
a possibility that this species has already been introduced
into Singapore’s water bodies, but this suspicion remains
unveried. Monopterus cuchia is supercially similar to
the native species Monopterus javanensis, and can only be
distinguished with the aid of the microscope; M. cuchia
possesses body scales which are absent in the native species;
and morphology of the gill opening and buccal chambers
(Nico et al., 2019).
At present, the bulk of sh aquaculture in Singapore focuses
on marine species, usually set in netted cages in the open
system along the Straits of Johor and Straits of Singapore.
Very few freshwater fish species are actively used in
aquaculture. The aquacultured freshwater sh species are
mainly imported from Malaysia.
ACKNOWLEDGEMENTS
This is part of the contribution for the Aquatic Invasive
Alien Species in Southeast Asia Symposium, held at the
National University of Singapore, 26 to 27 July 2017. The
Lee Foundation and the National University of Singapore
(Faculty of Science, Department of Biological Sciences,
and Lee Kong Chian Natural History Museum) had kindly
co-funded the Symposium.
We would like to acknowledge the following:
Peter Ng KL, the late Navjot Sodhi, and Hugh Tan TW,
for providing the initial impetus and framework to gather
baseline aquatic data from the impounded reservoirs in
Singapore; Alvin Lok SL, Tan Kai-xin, Ng Ting Hui, Ng
Heok Hee, Norman Lim T-L, Zeng Yiwen, Kenny Chua
WJ, Daniel Ng JJ, Jonathan Ho IK, Martyn Low EY, Ng
Pei Xin, Mary Rose Posa, Zeehan Jaafar, Lim Swee Cheng,
Hwang Wei Song, Lim Haw Chuan, Esther Clews, Jeremy
Yeo, for assistance with eldwork and companionship in
the eld; Yang SL (retired), Low E-wen and Quek Boon
Shan, for assistance and logistical support from PUB; Andy
Yap (Qian Hu), for verifying the Asian Arowana identity;
Tan Soo Ha, Nicole Chin, and Patrick Yap, for assistance
with ornamental sh trade aspects and specimens; Looh
CW, for providing Fig. 120; Martyn Low EY, for assistance
with locating obscure references; Rajeev Raghavan, Cai
Yixiong, Darren Sim ZH, Meryl Theng, Zhou Hang, and
Steve Wozniak, for providing recent sh records; HH Ng
and Z Jaafar, for providing critical comments and reviews.
We would also like to thank all the PUB site staff at all the
reservoirs for all their support. Funding from LKCNHM,
an MOE ARF Tier 1 grant (NUS grant no. R-154-000-
633-112), an Ah Meng Memorial Conservation Fund grant
(R-154-000-617-720), and PUB Singapore’s National Water
Agency (R-154-000-619-490, R-154-000-A20-490), to carry
out the research work is acknowledged.
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ADDENDUM
This section includes three new records post-2017 and one overlooked record. We feel strongly these should be included
to illustrate the complexity and uidity of monitoring the presence of non-native sh situation in Singapore; and the ease
of overlooking elusive records.
Order Siluriformes
Family Bagridae
Horabagrus brachysoma (Günther)
(Fig. 123, SAs)
Order Characiformes
Family Anostomidae
Leporinus fasciatus (Bloch)
(Fig. 125, SA)
References. None. This is a new record for Singapore.
Distribution. Kranji Reservoir.
Remarks. This is an overlooked record which was brought
to our attention recently (Raghavan R, pers. comm., February
2020). This species is common in the ornamental sh trade
(Tan HH, pers. obs.).
Family Claroteidae
Chrysichthys auratus (Geoffroy St. Hilaire)
(Fig. 124, AF[SPC])
References. Tan, 2019.
Distribution. Kallang River (Tan, 2019); Bishan Park
(unpublished record).
Remarks. This is based on observations (Tan, 2019; Tan &
Tay, 2019) and actual specimens. The source of introduction
of the specimens is a mystery (Tan, 2019).
References. Tan & Tay, 2019.
Distribution. Kallang River (Tan & Tay, 2019).
Remarks. This is a common ornamental sh, and most
likely a discarded pet sh.
Order Perciformes
Family Cichlidae
Veija × Mayaheros (?) hybrid
(Fig. 126, [SPC])
References. None. This is a new record for Singapore.
Distribution. Jurong Lake area (Tan HH, pers. obs.).
Remarks. This is most likely a hybrid, as all Veija specimens
lack the blue iridescent spots on the head.
Fig. 123. Horabagrus brachysoma, length not recorded, 700 g,
Kranji Reservoir (Looh CW).
Fig. 124. Chrysichthys auratus, 242.0 mm SL, Bishan (Lim RHB).
Fig. 125. Leporinus fasciatus, 67.0 mm SL, trade material.
Fig. 126. Veija × Mayaheros (?) hybrid, 115.9 mm SL, Jurong Lake.