ASIAN MYRMECOLOGY — Volume 15, e015006, 2022
ISSN 1985-1944 | eISSN: 2462-2362

DOI: 10.20362/am.015006

Remarkable diversity in a little red dot: a comprehensive checklistof known ant species in Singapore (Hymenoptera: Formicidae)
with notes on ecology and taxonomy
Wendy Y. Wang1*, Eunice J.Y. Soh2, Gordon W.J. Yong2, Mark K.L. Wong3,
Benoit Guénard4, Evan P. Economo5,6 and Seiki Yamane7
Lee Kong Chian Natural History Museum, National University of Singapore, 2 Conservatory Drive,
Singapore 117377
2
Department of Biological Sciences, National University of Singapore, 16 Science Drive 4, Singapore
1175583School of Biological Sciences, The University of Western Australia, Crawley WA 6009, Australia
4
School of Biological Sciences, The University of Hong Kong, Kadoorie Biological Sciences Building,
Pok Fu Lam Road, Hong Kong SAR, China
5
Biodiversity and Biocomplexity Unit, Okinawa Institute of Science and Technology Graduate University,
1919-1 Tancha, Onna-son, Kunigami-gun, Okinawa, 904-0495, Japan
6
Radcliffe Institute for Advanced Study, Harvard University, Cambridge, MA, USA 02138
7
Haruyama-cho, Kagoshima, 899-2704, Japan
1

Corresponding author: wywang24@gmail.com*

ABSTRACT. Despite a legacy of extensive deforestation, the 720 km2 city state of Singapore still
harbours impressively diverse flora and fauna. Given increasing evidence of global insect declines, weurgently need to better document and protect local insect diversity. Numerous species of ants (Hymenoptera, Formicidae) have been recorded or described from Singapore since its founding in 1819. However,it has been over a century since Hugo Viehmeyer (1916) documented a total of 159 species found in the
country. Here, we present an updated comprehensive checklist of all named species and subspecies ofants found in Singapore, with specimen collection data, and notes on taxonomy and ecology in the local
context. We compiled the list based on museum collections material (the Zoological Reference Collection), primary literature sources, and verified records from known overseas repositories. We documented
a total of 409 nominal species and subspecies, also a few notable morphospecies, from 10 subfamilies
and 100 genera. These include new records for 121 species and 10 genera. Another 96 species andsubspecies have types designated from Singapore; of these, 34 are currently considered as endemic. We
also raised nine subspecies to species and synonymized two species, providing reasons justifying eachstatus change: 1) Camponotus (Tanaemyrmex) carinifer stat. n., 2) Camponotus (Tanaemyrmex) tinctus
nom. rev., 3) Paraparatrechina malaccana stat. n., 4) Aphaenogaster simulans Forel, 1915 stat. n., 5)
Myrmicaria adpressipilosa stat. n., 6) Vollenhovia minuta stat. n., 7) Vollenhovia brevicornis (Emery,
1893) = V. fridae Forel, 1913 syn. n., 8) Hypoponera javana stat. n., 9) Hypoponera singaporensis
stat. n., 10) Mesoponera javana stat. n. Most species are considered native to Indomalaya, including 13
cosmopolitan tramps. Only 10 other species are presumed exotic to the region. At the time of writing,Singapore can be deemed the city with the highest recorded ant diversity in the world. Despite the sheer

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Wendy Y. Wang, Eunice J.Y. Soh, Gordon W.J. Yong, Mark K.L. Wong, Benoit Guénard, Evan P. Economo & Seiki Yamane

numbers, this list remains incomplete, with more species awaiting discovery or taxonomic resolution
in future. The immense diversity of ants in Singapore is mainly threatened by continued decimation ofremnant forest habitats and encroaching urban developments.
Keywords
Citation

Southeast Asia, biodiversity, natural history, museum collections
Wendy Y. Wang, Eunice J.Y. Soh, Gordon W.J. Yong, Mark K.L. Wong, Benoit Guénard, Evan
P. Economo & Seiki Yamane (2022). Remarkable diversity in a little red dot: a comprehensive
checklist of known ant species in Singapore (Hymenoptera: Formicidae) with notes on ecology
and taxonomy. Asian Myrmecology 15: e015006
Copyright
This article is distributed under a Creative Commons Attribution License CCBY4.0
Communicating Adam Cronin
EditorINTRODUCTION
The Republic of Singapore is an independent tropical island nation in Southeast Asia, the colloquial‘little red dot’ situated at the southernmost tip ofthe Malay Peninsula on the Equator. Singapore
has a total land area of approximately 720 square
kilometres (Singapore Land Authority, 2021),
including 64 smaller offshore islands. Despite
its small area and relative youth as a recognizedsovereignty (albeit initially governed as a British colony), substantial historical records since
its founding attest to the island nation’s diverse
flora and fauna, including the ants (see Brook et
al. 2003, Supplementary Information). Sir Thomas
Stamford Raffles founded Singapore as a colonial
British trading outpost in 1819, under the name of
the British East India Company.
A spill-over effect of increased commerceand human traffic that followed the colony’s establishment in 1819, was that faunal material fromSingapore was made available to foreign experts
residing elsewhere. The earliest formal records ofants identified from Singapore date back to theearly-mid 1800s to early 1900s. These comprised
mostly scattered, independent species descriptions
based on material from multiple collectors, andwere usually nested within publications of broader
geographic scope.
The first arguably comprehensive list ofants from Singapore, though not a ‘checklist’ bystrict definition, was created by Frederick Smith –then assistant curator in the zoological departmentof the British Museum – in 1857. He compileda larger catalogue of hymenopteran insects from
Borneo, Mount Ophir, Malacca, and Singapore col-lected by renowned explorer cum naturalist Alfred
Russel Wallace. Within this catalogue, Smith identified and roughly described 24 ant species fromSingapore. In 1858, Smith published another similar catalogue, this time documenting hymenopterans in the British Museum collections. He identified 29 ant species collected from Singapore in
the museum holdings. Over subsequent years, ant
species records from Singapore, including many
presumably new discoveries, were published discretely by various well-known entomologists suchas Carlo Menozzi (e.g., Menozzi 1926), Gustave
Mayr (e.g., Mayr 1897), Carlo Emery (e.g., Emery
1893b; 1896b; 1925), and Auguste-Henri Forel(e.g., Forel 1893; 1907b; 1911b; 1912b; 1913b).
In 1916 (some sources indicate 1915 to be
the correct publication date; we abide by 1916 inthis paper for consistency), the first species checklist dedicated to ants from Singapore was realized,with the publication of “Ameisen von Singapore”(Viehmeyer 1916), albeit including a few species
described from neighbouring countries. In thearticle, Dresden entomologist and schoolmaster
Hugo Viehmeyer identified 159 ant species fromSingapore, describing several in painstaking detail, based on material collected by Hans Friedrich
Overbeck, a German mercantile trader and Malayologist (Taylor 2014). Overbeck resided mainlyin Singapore before 1914 (Taylor 2014), during
which he amassed ant specimens mostly from his
own back garden (Overbeck 1924), and also selectlocalities including some in Malaysia or Indonesia.
In 1951, Chapman and Capco presented achecklist of ants of Asia, among which 143 species
were indicated to be found in Singapore, albeit
with misreported localities and nomenclatural er-

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

rors. This checklist, extensive as it seemed, paledin comparison to Viehmeyer’s (1916) seminal
work in terms of taxonomic rigour and descriptive detail. Following Chapman and Capco (1951),
no formal attempts have been made to properly
document Singapore’s whole ant fauna until now.
Given mounting evidence and recent
concerns over global insect declines (Saunders etal. 2020; Halsch et al. 2021), conservation bodies
worldwide have increasingly emphasized on theurgent need to document and understand insect
diversity at local scales. This applies even to urbanized cities, which have been shown to support
significant biodiversity despite lacking perceivably
good habitats (Brassard et al. 2021). Conservation
decisions are effective only if informed by accurate
data on species diversity and distributions. Thus,
species checklists are crucial resources for biological conservation, highlighting threatened taxa andtheir habitats that require prioritized protection.
We here present an updated and comprehensive checklist of ant species in Singapore, withdetails on an unprecedented scale. We document
all nominal species and subspecies identified fromavailable material and/or literature, from the 1800s
post-founding till present. We also include speci-

men collection data, notes on observed habitat andecology, and comments on taxonomy or other relevant information for every listed species.
MATERIAL AND METHODS
Material examined for the purposes of this checklist were from the Zoological Reference Collection (ZRC), housed in the Lee Kong Chian Natural
History Museum (previously known as the Raffles
Museum of Biodiversity Research), and the Seiki
Yamane Collection (SKYC). Invaluable contributions of material from the 1960s – early 1990s weremade by the late Professor Dennis H. Murphy, a fervent collector of hexapods and legendary entomologist from (what was then known as) the University of Malaya (Chan 1991). Murphy’s collections,distributed across the ZRC and other repositories
worldwide, have been indispensable for numerous
species discoveries in Singapore. Material available
for the compilation of this checklist were collectedin the 1960s until the late 2000s, mostly from nature
reserves and variable pockets of spontaneous orhuman-modified vegetation interspersing the larger
urban matrix on the main island, or scattered acrossthe surrounding offshore islands (Fig. 1).

Fig. 1. Map of Singapore with current day satellite imagery. The largest forest reserve, the Central Catchment Nature
Reserve, forms a major part of the green area at the centre of the map. Base map vector by Teo Siyang. Satellite
imagery based on Fair Use, from Google Maps – Imagery©2021 TerraMetrics, Map data©2021. Map projection:
SVY21. Inset: Map of Southeast Asia with red arrow indicating Singapore's geographic position.

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Spontaneous vegetation – defined asvegetation naturally developed without humanintervention – occupies less than 28% of total
land area in modern Singapore (Yee et al. 2016),
and is mostly found in protected reserves. Thesereserves encompass small tracts of inland primary
forest and substantial areas of abandoned agricultural lands (Corlett 1992; O’Dempsey 2014). The
largest of these reserves is the Central Catchment
Nature Reserve (CCNR) – comprising ca. 1622
ha. of gazetted catchment areas surrounding threemajor reservoirs – MacRitchie, Peirce and Seletar (Corlett 1992; O’Dempsey 2014). Besides the
CCNR, dispersed remnants of primary rainforest
are also found in the 71 ha. Bukit Timah Nature
Reserve (BTNR), and around ca. 15 ha. of freshwater swamp forest in an area known as Nee Soon
(Corlett 1992). Primary lowland dipterocarp forest
and freshwater swamp forest together occupy lessthan 1% of total land area (Yee et al. 2011, 2016).
Remnant spontaneous vegetation predominantly
consists of secondary vegetation or forests – floral
communities regenerated on previously exploitedland through natural succession (Yee et al. 2016).
The subsection ‘Material not physicallyexamined’ refers to accessioned specimens in otherformal repositories which we did not physicallyexamine for this study. These include not justmuseums, but also some private and well-curated
collections. All material listed in this checklist areof the worker caste, unless specified otherwise.
For the subsection ‘Literature’, only
primary literature sources explicitly stating thedirect examination and identification of physicalspecimens from Singapore were cited per relevant
species, for greater information reliability. Secondary literature sources that cite primary sources but
do not indicate involvement of actual specimens
were omitted for brevity.
Significant baseline data of existing species records were gleaned from the Global Ant
Biodiversity Database (GABI) (Guénard et al.
2017), albeit subject to extensive cross-checking
against other reputable online public databases
such as AntCat (Bolton 2021) and AntWeb (California Academy of Science, 2022), with error rectification where necessary.

Abbreviations of specimen repositories
ANIC

Australian National Insect Collection,
CSIRO, Australia
BMNH British Museum of Natural History,
London, United Kingdom
CASC
California Academy of Sciences, San
Francisco, USA
DEIC
Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany
FMNH The Field Museum of Natural History,
Chicago, USA
HKUC
Hong Kong University Collections,
Hong Kong, Republic of China
HNHM Hungarian Natural History Museum,
Budapest, Hungary
JKWC
James K. Wetterer Collection, Florida,USA
MCZ
Museum of Comparative Zoology,
Harvard University, Cambridge, Massachusetts, USA
MHNG Muséum d’Histoire Naturelle Geneva,
Geneva, Switzerland
MNHN Muséum National d’Histoire Naturelle,
Paris, France
MNHU Museum für Naturkunde der Humboldt-Universität, Berlin, Germany[may be synonym of ZMHB]
MSC
Mostafa Sharaf Collection, Riyadh,
Saudi Arabia
MSNG
Museo Civico di Storia Naturale di
Genova “Giacomo Doria”, Genoa,
Italy
NHMB Naturhistorisches Museum, Basel,
Switzerland
NHMW Naturhistorisches Museum Wien, Vienna, Austria
OUMNH Oxford University Museum of Natural
History, Oxford, United Kingdom
PSWC
Philip S. Ward Collection, Universityof California, Davis, C.A., USA
RBINS Royal Belgian Institute of Natural Sciences, Brussels, Belgium
SKYC
Seiki Yamane Collection, Kagoshima,
Japan
THNHM Thailand Natural History Museum, National Science Museum, Pathum Thani,
ThailandRemarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

UCDC
USNM
ZCW
ZMHB
ZRC

University of California, Davis, Collection, Davis, CA, USA
Smithsonian National Museum of
Natural History, United States National
Museum, Washington, D.C., USA
Zettel Collection Wien, Vienna, Austria
Zoologisches Museum der HumboldtUniversität zu Berlin, part of Museum
für Naturkunde Berlin, Germany
Zoological Reference Collection, Lee
Kong Chian Natural History Museum,
National University of Singapore, Singapore

RESULTS AND DISCUSSION
Ant diversity in Singapore
A total of 409 nominal ant species and subspecies
from 10 subfamilies were identified from ZRC
material, the available literature, or other known
curated records. Our list surpasses Viehmeyer’s(1916) list (i.e., 159 species) by ca. 150%. Of
these, 121 constitute new formal species records
from Singapore. Another 96 species (and subspecies) were first described and/or have designatedtypes from Singapore. Thirty-four of these are currently known only from Singapore and nowhere

else in the world. We also raised nine subspecies
to species, and synonymized two original species
based on our own assessment of available specimens and online type images, and original descriptions where possible.
We recorded 100 genera, including
10 new genus records: Amblyoponinae – Mystrium; Dolichoderinae – Ochetellus; Dorylinae– Parasyscia, Syscia; Ectatomminae – Holcoponera; Formicinae – Gesomyrmex; Myrmicinae Aphaenogaster, Gauromyrmex and Lordomyrma ;Ponerinae - Myopias. The top three most speciose
subfamilies were (species and subspecies counts inparentheses): Myrmicinae (158), Formicinae (117)and Ponerinae (61) (Fig.2).
The full checklist and corresponding
information mentioned are summarized in Table
1. Newly established species that were raised
from subspecies are indicated with suffixes ‘stat.
n.’ (novel status) or ‘nom. rev.’ (revived species
name) in the same table. Synonymized species
are also indicated with the suffix ‘syn. n.’(novel
synonymy). Detailed information on each species
is provided at the end of this section, under ‘Species Accounts’. Reasons justifying any taxonomicchanges can be found under ‘Remarks’ per relevant
species section.

Fig. 2. Number of nominal species and subspecies per subfamily in Singapore. Unnamed morphospecies are notincluded in species counts.

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Wendy Y. Wang, Eunice J.Y. Soh, Gordon W.J. Yong, Mark K.L. Wong, Benoit Guénard, Evan P. Economo & Seiki Yamane

Fig. 3. Accumulation graph of total known species (including subspecies) collected and/or recorded over timein years (1850 – present).

In addition to the full checklist, we havealso compiled a list of 91 species and subspecies
that were last known to be collected more than 100
years ago from the time of writing, i.e., 2022 (Table2). While it is possible that these species might havegone extinct locally, we cannot make a definite conclusion given the greater likelihood of incomplete
sampling and identification bottlenecks. Over thecourse of examining ZRC material, we noted thatmany specimens could not be confidently identified
to nominal species – some of these may correspondto species listed in Table 2.
Further, we flagged 14 species and subspecies currently known only from Singapore, butwhich have not been collected for more than acentury (Table 3). The current statuses of these
forms may require critical evaluation, and at leasta few might be synonyms of more widespread, recognized species. We listed possibly synonymous
species and their known biogeographic distributions where relevant.
The substantial volume of unidentifiablematerial also implies that the present checklist is
likely an under-representation of Singapore’s true
ant fauna. Moreover, the accumulation curve of
new species records over time appears to rise withno signs of reaching a plateau (Fig. 3), suggest-

ing that more species await discovery. We haveincluded details for a few outstanding unnamed
morphospecies that may warrant further taxonomic
attention in future, including the first local recordsof the doryline genus Syscia, and the myrmicine
genus Lordomyrma. These unnamed species are
excluded from the final checklist tally.
Exotics and natives
Singapore’s present topography is the culmination of a relentless trajectory of land use changes
against a historical backdrop of colonization andagriculture (O’Dempsey 2014), followed by rapid
urbanization and industrialization after the second
world war (Yee et al. 2011). Given this uniqueenvironmental context and Singapore’s status asa hub for international commerce, we expected
the local ant fauna to be dominated by exotics andcosmopolitan tramp species.
Contrary to expectations, we foundonly ten species considered exotic to Indomalaya– which Singapore is part of – including sevenrecognized cosmopolitan tramps defined by Wetterer (2015): Pheidole megacephala, Monomorium
monomorium, Solenopsis geminata, Strumigenys
eggersi, S. emmae, S. membranifera, S. rogeri,
Syllophopsis subcoeca, Technomyrmex difficilis
and Trichomyrmex destructor.

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Table 1. Summarized checklist of named ant species (and subspecies) found in Singapore (SG). For ‘Year First
Collected/Recorded’, the year of first collection is given precedence if both years are known. Species found onlyin Singapore (at the time of writing) are in bold.
Year First Type
New Lit.
Collected/ Locality
RecordRecordedSG

Subfamily

Scientific Name

Amblyoponinae

Myopopone castanea (Smith, 1860)1970

Amblyoponinae

Mystrium camillae Emery, 1889

2017

x

Amblyoponinae

Prionopelta kraepelini Forel, 1905

2012

x

Amblyoponinae

Stigmatomma besucheti (Baroni Urbani, 1978)

1964

Amblyoponinae

Stigmatomma reclinatum (Mayr, 1879)

1964

Dolichoderinae

Chronoxenus dalyi (Forel, 1895)

1916

Dolichoderinae

Chronoxenus wroughtonii (Forel, 1895)

1985

Dolichoderinae

Dolichoderus affinis Emery, 1889

1916

Dolichoderinae

Dolichoderus crawleyi Donisthorpe, 1917

1917

Dolichoderinae

Dolichoderus indrapurensis Forel, 1912

1968

Dolichoderinae

Dolichoderus sulcaticeps (Mayr, 1870)

1916

Dolichoderinae

Dolichoderus thoracicus Smith, 1860

1863

Dolichoderinae

Iridomyrmex anceps (Roger, 1863)

1916

Dolichoderinae

Ochetellus glaber (Mayr, 1862)

2007

Dolichoderinae

Philidris cordata (Smith, 1859)

1911

Dolichoderinae

Philidris laevigata (Emery, 1895)

1916

Dolichoderinae

Philidris myrmecodiae (Emery, 1887)

2014

Dolichoderinae

Tapinoma andamanense capsincola Forel, 1911

1916

Dolichoderinae

Tapinoma glaucum (Viehmeyer, 1916)

1916

Dolichoderinae

Tapinoma indicum Forel, 1895

1916

Dolichoderinae

Tapinoma melanocephalum (Fabricius, 1793)

1857

Dolichoderinae

Technomyrmex albipes (Smith, 1861)

1916

Dolichoderinae

Technomyrmex difficilis Forel, 1892

≤ 1948Dolichoderinae

Technomyrmex elatior Forel, 1902

1916

Dolichoderinae

Technomyrmex horni Forel, 1912

1989

Dolichoderinae

Technomyrmex kraepelini Forel, 1905

1967

Dolichoderinae

x
x
x
x

x

x
x

Technomyrmex pratensis (Smith, 1860)

1994

Dolichoderinae

Technomyrmex schimmeri Viehmeyer, 1916

1916

x

x

Dolichoderinae

Technomyrmex strenuus Mayr, 1872

1872

x [part]

Dolichoderinae

Technomyrmex textor Forel, 1909

2016

x

Dolichoderinae

Technomyrmex vitiensis Mann, 1921

1979

x

Dorylinae

Aenictus camposi Wheeler & Chapman, 1925

1967

x

Dorylinae

Aenictus cylindripetiolus Jaitrong & Yamane, 2013

2016

x

Dorylinae

Aenictus gracilis Emery, 1893

1967

Dorylinae

Aenictus seletarius Wong & Guénard, 2016

2015

Dorylinae

Aenictus shuckardi Forel, 1901

1916

Dorylinae

Aenictus yamanei Wiwatwitaya & Jaitrong, 2011

2017

x
x
x

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Subfamily
Dorylinae

Scientific Name

Year First Type
New Lit.
Collected/ Locality
RecordRecorded
SG

Dorylus laevigatus (Smith, 1857)

1916

Dorylinae

Lioponera hewitti (Wheeler, 1919)

2014

Dorylinae

Lioponera singaporensis (Viehmeyer, 1916)

1916

x

Dorylinae

Ooceraea biroi (Forel, 1907)

1907

x

Dorylinae

Parasyscia dohertyi (Emery, 1902)

1968

Dorylinae

Simopone bakeri Menozzi, 1926

1926

Ectatomminae

Holcoponera cribrata (Emery, 1900)

1988

Ectatomminae

Stictoponera bicolor (Emery, 1889)

2016

Ectatomminae

Stictoponera binghamii (Forel, 1900)

1916

Ectatomminae

Stictoponera coxalis (Roger, 1860)

1967

x

x
x
x
x

Ectatomminae

Stictoponera gabata (Lattke, 2004)

2016

Ectatomminae

Stictoponera hyalina (Lattke, 2004)

1967

x

Ectatomminae

Stictoponera ortostoma (Lattke, 2004)

1967

x

Ectatomminae

Stictoponera posteropsis Gregg, 1951

1981

Formicinae

Acropyga acutiventris Roger, 1862

1967

Formicinae

Acropyga inezae Forel, 19122019

Formicinae

Acropyga oceanica Emery, 1900

2004

x
x

Formicinae

Acropyga rubescens Forel, 1894

1916

Formicinae

Anoplolepis gracilipes (Smith, 1857)

1857

x

Formicinae

Camponotus (Karavaievia) overbecki Viehmeyer, 1916

1916

x

Formicinae

Camponotus (Myrmamblys) adustus Viehmeyer, 1916

1916

x

Formicinae

Camponotus (Myrmamblys) bedoti Emery, 1893

1907

Formicinae

Camponotus (Myrmamblys) bellus Forel, 1908

1916

Formicinae

Camponotus (Myrmamblys) ephippiatus Viehmeyer,
1916

1916

Formicinae

Camponotus (Myrmamblys) gretae Forel, 1902

1916

Formicinae

Camponotus (Myrmamblys) reticulatus Roger, 1863

1976

Formicinae

Camponotus (Myrmamblys) reticulatus sericellus
Viehmeyer, 1916

1916

x

Formicinae

Camponotus (Myrmosaulus) camelinus (Smith, 1857)

1857

x

Formicinae

Camponotus (Myrmosericus) parius Emery, 1889

1994

Formicinae

Camponotus (Myrmotarsus) nigricans enganensis Forel,
1916

1916

Formicinae

Camponotus (Myrmotarsus) quadrisectus hians Forel,
1907

1897

x

Formicinae

Camponotus (Myrmotarsus) quadrisectus margaritae
Forel, 1907

1897

x

Formicinae

Camponotus (Tanaemyrmex) albosparsus Bingham,
1903

1996

Formicinae

Camponotus (Tanaemyrmex) arrogans (Smith, 1858)

1858

x

x

x

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Subfamily

Scientific Name

Year First Type
New Lit.
Collected/ Locality
RecordRecorded
SG

Formicinae

Camponotus (Tanaemyrmex) carinifer Viehmeyer, 1916
stat.n.

1916

Formicinae

Camponotus (Tanaemyrmex) festinus eximius Emery,
1900

1916

Formicinae

Camponotus (Tanaemyrmex) festinus simaluranus Forel,
1915

1916

Formicinae

Camponotus (Tanaemyrmex) irritans (Smith, 1857)≤ 2000Formicinae

Camponotus (Tanaemyrmex) maculatus obfuscatus
Viehmeyer, 1916

1916

Formicinae

Camponotus (Tanaemyrmex) maculatus subnudus Emery, 1889

1916

Formicinae

Camponotus (Tanaemyrmex) maxwellensis Forel, 1913

1916

Formicinae

Camponotus (Tanaemyrmex) mitis (Smith, 1858)

1916

Formicinae

Camponotus (Tanaemyrmex) tinctus (Smith, 1858),
nom. rev.

1916

Formicinae

Camponotus (Tanaemyrmex) variegatus (Smith, 1858)

1858

Formicinae

Camponotus (Tanaemyrmex) variegatus fuscithorax
Dalla Torre, 1893

1916

Formicinae

Colobopsis badia (Smith, 1857)

1857

Formicinae

Colobopsis leonardi (Emery, 1889)

1916

Formicinae

Colobopsis leonardi gracilenta (Viehmeyer, 1916)

1916

x
x

Formicinae

Colobopsis moeschi lygaea (Viehmeyer, 1916)

1916

Formicinae

Colobopsis rothneyi (Forel, 1893)

1916

Formicinae

Colobopsis rothneyi krafti (Forel, 1901)

1901

Formicinae

Colobopsis vitrea (Smith, 1860)

1916

Formicinae

Colobopsis vitrea angustula (Emery, 1925)

1870

Formicinae

Colobopsis vitrea vittatula (Forel, 1913)

1916

Formicinae

Dinomyrmex gigas (Latreille, 1802)

1857

x

x

x [part]xx

Formicinae

Echinopla lineata Mayr, 1862

1975

Formicinae

Echinopla melanarctos Smith, 1857

1857

Formicinae

Echinopla rugosa André, 1892

1916

Formicinae

Euprenolepis procera (Emery, 1900)

2012

x

Formicinae

Gesomyrmex spatulatus Cole, 1949

1969

x

Formicinae

Lepisiota rothneyi splendida (Viehmeyer, 1916)

1916

Formicinae

Myrmoteras (Myagroteras) bakeri Wheeler, 1919

1965

Formicinae

Myrmoteras barbouri Creighton, 19301965

Formicinae

Nylanderia birmana (Forel, 1902)

2021

Formicinae

Nylanderia bourbonica (Forel, 1886)

2012

Formicinae

Nylanderia clandestina (Mayr, 1870)

1916

Formicinae

Nylanderia kraepelini (Forel, 1905)

2021

Formicinae

Nylanderia yerburyi (Forel, 1894)

1912

x

x
x
x

x

10 of 152

Wendy Y. Wang, Eunice J.Y. Soh, Gordon W.J. Yong, Mark K.L. Wong, Benoit Guénard, Evan P. Economo & Seiki Yamane

Subfamily

Scientific Name

Year First Type
New Lit.
Collected/ Locality
RecordRecorded
SG

Formicinae

Oecophylla smaragdina (Fabricius, 1775)

1916

Formicinae

Overbeckia subclavata Viehmeyer, 1916

1916

Formicinae

Paraparatrechina malaccana (Viehmeyer, 1916) stat.n.

1985

x
x

Formicinae

Paraparatrechina opaca (Emery, 1887)

2014

Formicinae

Paratrechina longicornis (Latreille, 1802)

1931

Formicinae

Plagiolepis bicolor Forel, 1901

1916

Formicinae

Plagiolepis exigua Forel, 1894

1916

Formicinae

Polyrhachis (Campomyrma) creusa Emery, 1897

1916

Formicinae

Polyrhachis (Chariomyrma) arcuata (Le Guillou, 1842)

1907

Formicinae

Polyrhachis (Chariomyrma) modesta Smith, 1857

1857

Formicinae

Polyrhachis (Cyrtomyrma) laevissima Smith, 1858

1916

Formicinae

Polyrhachis (Cyrtomyrma) lepida Kohout, 2006

2014

x

x
x

Formicinae

Polyrhachis (Myrma) beccarii Mayr, 18722015

Formicinae

Polyrhachis (Myrma) carbonaria Smith, 1857

1962

x

Formicinae

Polyrhachis (Myrma) hosei Donisthorpe, 1942

2014

x

Formicinae

Polyrhachis (Myrma) illaudata Walker, 18591973

Formicinae

Polyrhachis (Myrma) inermis Smith, 1858

2014

Formicinae

Polyrhachis (Myrma) nigropilosa Mayr, 1872

1926

Formicinae

Polyrhachis (Myrma) obesior Viehmeyer, 1916

1916

Formicinae

Polyrhachis (Myrma) orsylla Smith, 1861

1907

Formicinae

Polyrhachis (Myrma) orsylla ritsemai Mayr, 1883

1916

Formicinae

Polyrhachis (Myrma) proxima Roger, 1863

1893

Formicinae

Polyrhachis (Myrma) pruinosa Mayr, 1872

1913

Formicinae

Polyrhachis (Myrma) striata Mayr, 1862

1893

Formicinae

Polyrhachis (Myrma) striatorugosa Mayr, 1862

1989

Formicinae

Polyrhachis (Myrma) villipes Smith, 1857

1916

Formicinae

Polyrhachis (Myrma) vindex Smith, 18571962

Formicinae

Polyrhachis (Myrmatopa) charaxa Smith, 1860

1896

Formicinae

Polyrhachis (Myrmatopa) flavicornis Smith, 1857

1857

x

Formicinae

Polyrhachis (Myrmatopa) leviuscula Viehmeyer, 1916

1912

x

Formicinae

Polyrhachis (Myrmatopa) piliventris Smith, 1858

1858

x

Formicinae

Polyrhachis (Myrmatopa) varicolor Viehmeyer, 1916

1916

x

Formicinae

Polyrhachis (Myrmhopla) abdominalis Smith, 1858

1995

Formicinae

Polyrhachis (Myrmhopla) armata (Le Guillou, 1842)

1916

Formicinae

Polyrhachis (Myrmhopla) armata defensa Smith, 1857

1857

Formicinae

Polyrhachis (Myrmhopla) bicolor Smith, 1858

1916

Formicinae

Polyrhachis (Myrmhopla) bicolor aurinasis Forel, 1901

1907

Formicinae

Polyrhachis (Myrmhopla) chalybea Smith, 1857

1857

Formicinae

Polyrhachis (Myrmhopla) cryptoceroides Emery, 1887

2016

x
x

x
x

x

x [part]x

11 of 152

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Subfamily

Scientific Name

Year First Type
New Lit.
Collected/ Locality
RecordRecorded
SG

Formicinae

Polyrhachis (Myrmhopla) dives Smith, 1857

1857

x

Formicinae

Polyrhachis (Myrmhopla) hector Smith, 1857

1857

x

Formicinae

Polyrhachis (Myrmhopla) hippomanes ceylonensis Emery, 1893

1916

Formicinae

Polyrhachis (Myrmhopla) lucidula Emery, 1893

1916

Formicinae

Polyrhachis (Myrmhopla) muelleri Forel, 1893

1893

Formicinae

Polyrhachis (Myrmhopla) saevissima Smith, 1860

2012

x

Formicinae

Polyrhachis (Myrmhopla) thailandica Kohout, 2006

2016

x

x

Formicinae

Polyrhachis (Myrmhopla) tibialis parsis Emery, 1900

1916

Formicinae

Polyrhachis (Myrmothrinax) frauenfeldi Mayr, 1862

1916

Formicinae

Polyrhachis (Myrmothrinax) saigonensis Forel, 1886

1916

Formicinae

Polyrhachis (Myrmothrinax) textor Smith, 1857

1916

Formicinae

Polyrhachis (Myrmothrinax) thrinax inconstans Viehmeyer, 1916

1916

x

Formicinae

Polyrhachis (Myrmothrinax) thrinax overbecki Dorow,
1995

1916

x

Formicinae

Polyrhachis (Polyrhachis) bellicosa Smith, 1859

1916

Formicinae

Polyrhachis (Polyrhachis) olybria Forel, 1912

1967

Formicinae

Polyrhachis (Polyrhachis) ypsilon Emery, 1887

1875

Formicinae

Prenolepis jerdoni Emery, 1893

1916

x

Formicinae

Prenolepis subopaca Emery, 1900

1967

Formicinae

Pseudolasius badius Viehmeyer, 1916

1916

x

Formicinae

Pseudolasius circularis Viehmeyer, 1916

1916

x

Formicinae

Pseudolasius ludovici Forel, 1913

1916

Formicinae

Pseudolasius martini Forel, 1911

1916

Formicinae

Pseudolasius mayri duplicatus Viehmeyer, 1916

1916

Formicinae

Pseudolasius trimorphus Karavaiev, 1929

2018

Leptanillinae

Leptanilla havilandi Forel, 1901

1901

x

Leptanillinae

Leptanilla hypodracos Wong & Guénard, 2016

2015

x

Leptanillinae

Protanilla rafflesi Taylor, 1990

1964

x

Myrmicinae

Acanthomyrmex ferox Emery, 1893

1981

x

Myrmicinae

Aphaenogaster simulans Forel, 1915 stat. n.

2016

x

Myrmicinae

Calyptomyrmex beccarii Emery, 1887

1916

Myrmicinae

Calyptomyrmex loweryi Shattuck, 20112016

Myrmicinae

Cardiocondyla kagutsuchi Terayama, 1999

2003

Myrmicinae

Cardiocondyla obscurior Wheeler, 1929

2015

Myrmicinae

Cardiocondyla strigifrons Viehmeyer, 1922

2017

Myrmicinae

Cardiocondyla tjibodana Karavaiev, 1935

1995

Myrmicinae

Cardiocondyla wroughtonii (Forel, 1890)

1912

Myrmicinae

Carebara affinis (Jerdon, 1851)

1916

x
x

x
x
x

12 of 152

Wendy Y. Wang, Eunice J.Y. Soh, Gordon W.J. Yong, Mark K.L. Wong, Benoit Guénard, Evan P. Economo & Seiki Yamane

Subfamily

Scientific Name

Year First Type
New Lit.
Collected/ Locality
RecordRecorded
SG

Myrmicinae

Carebara castanea Smith, 1858

2015

x

Myrmicinae

Carebara diversa (Jerdon, 1851)

1916

Myrmicinae

Carebara overbecki (Viehmeyer, 1916)

1916

x

Myrmicinae

Carebara silenus (Smith, 1858)

1858

x

Myrmicinae

Cataulacus granulatus (Latreille, 1802)

1876

Myrmicinae

Cataulacus horridus Smith, 1857

1916

Myrmicinae

Cataulacus latissimus Emery, 1893

1974

Myrmicinae

Cataulacus praetextus Smith, 1867

1916

Myrmicinae

Crematogaster anthracina Smith, 1857

1857

Myrmicinae

Crematogaster aurita Karavaiev, 1935

2015

Myrmicinae

Crematogaster borneensis André, 1896

1916

Myrmicinae

Crematogaster brunnea ruginota Santschi, 1928

1916

Myrmicinae

Crematogaster coriaria Mayr, 1872

2017

Myrmicinae

Crematogaster decamera Forel, 1910

1916

Myrmicinae

Crematogaster difformis Smith, 1857

1857

x
x

x
x

Myrmicinae

Crematogaster dohrni artifex Mayr, 1878

1878

x

Myrmicinae

Crematogaster egidyi szaboi Forel, 1913

1898

x

Myrmicinae

Crematogaster ferrarii Emery, 1888

1916

Myrmicinae

Crematogaster inflata Smith, 1857

1857

x

Myrmicinae

Crematogaster linsenmairi Feldhaar, Maschwitz &Fiala, 2016

1916

x

Myrmicinae

Crematogaster modiglianii Emery, 1900

1985

Myrmicinae

Crematogaster overbecki Viehmeyer, 1916

1916

Myrmicinae

Crematogaster rogenhoferi Mayr, 1879

1916

Myrmicinae

Crematogaster sewardi Forel, 1901

1983

Myrmicinae

Crematogaster subnuda discinodis Emery, 1893

1893

Myrmicinae

Crematogaster treubi Emery, 1896

1916

Myrmicinae

Crematogaster yappi Forel, 1901

2012

x

Myrmicinae

Crematogaster (Orthocrema) bandarensis Forel, 1913

2016

x

Myrmicinae

Crematogaster (Orthocrema) longipilosa Forel, 1907

1912

Myrmicinae

Crematogaster (Orthocrema) myops Forel, 1911

1988

Myrmicinae

Crematogaster (Orthocrema) quadriruga Forel, 1911

1994

Myrmicinae

Dilobocondyla fulva Viehmeyer, 1916

1916

Myrmicinae

Erromyrma latinodis (Mayr, 1872)

1916

Myrmicinae

Eurhopalothrix heliscata Wilson & Brown, 1985

1983

x
x

x

x
x
x

Myrmicinae

Eurhopalothrix omnivaga Taylor, 1990

2015

x

Myrmicinae

Eurhopalothrix procera (Emery, 1897)

1991

x

Myrmicinae

Gauromyrmex acanthinus (Karavaiev, 1935)

2017

x

Myrmicinae

Liomyrmex gestroi (Emery, 1887)

1969

Myrmicinae

Lophomyrmex bedoti Emery, 18931916

13 of 152

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Subfamily

Scientific Name

Year First Type
New Lit.
Collected/ Locality
RecordRecorded
SG

Myrmicinae

Mayriella transfuga Baroni Urbani, 1977

1968

Myrmicinae

Meranoplus bicolor (Guérin-Méneville, 1844)

1916

Myrmicinae

Meranoplus malaysianus Schödl, 1998

2017

Myrmicinae

Meranoplus mucronatus Smith, 1857

1980

Myrmicinae

Metapone murphyi Wang, Yamada & Eguchi, 2019

1981

x
x
x

Myrmicinae

Monomorium atomum procax Forel, 1911

1911

x

Myrmicinae

Monomorium butteli demochrum Viehmeyer, 1916

1916

x

Myrmicinae

Monomorium floricola (Jerdon, 1851)

1916

Myrmicinae

Monomorium hospitum Viehmeyer, 1916

1916

Myrmicinae

Monomorium monomorium Bolton, 1987

2012

x

Myrmicinae

Monomorium pharaonis (Linnaeus, 1758)

1857

Myrmicinae

Myrmecina bandarensis Forel, 1913

2012

Myrmicinae

Myrmecina magnificens Wong & Guénard, 2016

2015

Myrmicinae

Myrmecina maryatiae Okido, Ogata & Hosoishi, 2020

2017

x

Myrmicinae

Myrmicaria adpressipilosa Santschi, 1928 stat. n.

1974

x

Myrmicinae

Myrmicaria arachnoides (Smith, 1857)

1916

Myrmicinae

Myrmicaria arachnoides lutea Emery, 1900

1922

Myrmicinae

Myrmicaria luteiventris Emery, 1900 [see Remarks in
text]

1922

Myrmicinae

Myrmicaria melanogaster Emery, 1900

1973

x

Myrmicinae

Paratopula demeta Bolton, 1988

2016

x

Myrmicinae

Paratopula oculata Smith, 1857

1988

Myrmicinae

Pheidole aglae Forel, 1913

1916

Myrmicinae

Pheidole aristotelis Forel, 1911

1968

Myrmicinae

Pheidole binghamii Forel, 1902

2016

Myrmicinae

Pheidole cariniceps Eguchi, 2001

2012

x
x

x

Myrmicinae

Pheidole clypeocornis Eguchi, 2001

2012

Myrmicinae

Pheidole elisae Emery, 1900

2016

Myrmicinae

Pheidole fervens Smith, 1858

1858

Myrmicinae

Pheidole hortensis Forel, 1913

2016

x

Myrmicinae

Pheidole magrettii Emery, 1887

2018

x

Myrmicinae

Pheidole megacephala (Fabricius, 1793)

1901

Myrmicinae

Pheidole nodgii Forel, 1905

1916

Myrmicinae

Pheidole nodgii verlatenensis Wheeler, 1937

2016

Myrmicinae

Pheidole parva Mayr, 1865

2012

Myrmicinae

Pheidole plagiaria Smith, 1860

1989

Myrmicinae

Pheidole plinii Forel, 1911

1911

Myrmicinae

Pheidole rinae Emery, 1900

1916

Myrmicinae

Pheidole rugifera Eguchi, 2001

2016

x

x

x
x

14 of 152

Wendy Y. Wang, Eunice J.Y. Soh, Gordon W.J. Yong, Mark K.L. Wong, Benoit Guénard, Evan P. Economo & Seiki Yamane

Subfamily

Scientific Name

Year First Type
New Lit.
Collected/ Locality
RecordRecorded
SG

Myrmicinae

Pheidole sexspinosa Mayr, 1870

1977

Myrmicinae

Pheidole singaporensis Özdikmen, 2010

1857

Myrmicinae

Pheidole singaporensis conicollis (Emery, 1900)

1916

x

Myrmicinae

Pheidole singaporensis continentis (Forel, 1911)

1916

Myrmicinae

Pheidole tandjongensis Forel, 1913

2016

x

Myrmicinae

Pheidole tjibodana Forel, 1905

2016

x

Myrmicinae

Pheidole umbonata Mayr, 1870

2012

x

Myrmicinae

Pristomyrmex bicolor Emery, 1900

1987

x

Myrmicinae

Pristomyrmex brevispinosus Emery, 1887

2016

x

Myrmicinae

Pristomyrmex costatus Wang, 2003

1994

Myrmicinae

Pristomyrmex picteti Emery, 1893

2003

Myrmicinae

Pristomyrmex punctatus (Smith, 1860)

1916

Myrmicinae

Proatta butteli Forel, 1912

1916

Myrmicinae

Rhopalomastix glabricephala Wang, Yong & Jaitrong,
2018

2016

x

Myrmicinae

Rhopalomastix javana Wheeler, 1929

2017

Myrmicinae

Rhopalomastix johorensis Wheeler, 1929

1916

Myrmicinae

Rhopalomastix murphyi Wang, Yong & Jaitrong, 2018

1981

x

Myrmicinae

Rhopalomastix striata Wang, Yong & Jaitrong, 2018

2016

x

Myrmicinae

Rhopalomastix tenebra Wang, Yong & Jaitrong, 2018

1983

x

Myrmicinae

Rostromyrmex pasohensis Rosciszewski, 1994

1988

Myrmicinae

Rotastruma recava Bolton, 1991

1973

Myrmicinae

Solenopsis geminata (Fabricius, 1804)

1897

Myrmicinae

Solenopsis overbecki Viehmeyer, 1916

1916

Myrmicinae

Strumigenys chapmani Brown, 1954

2016

Myrmicinae

Strumigenys eggersi Emery, 1890

2014

Myrmicinae

Strumigenys emmae (Emery, 1890)

1909

Myrmicinae

Strumigenys epyna Bolton 2000

2016

Myrmicinae

Strumigenys extemena (Taylor, 1968)

1965

Myrmicinae

Strumigenys godeffroyi Mayr, 1866

1916

Myrmicinae

Strumigenys juliae Forel, 1905

2000

Myrmicinae

Strumigenys koningsbergeri Forel, 1905

1990

Myrmicinae

Strumigenys kraepelini Forel, 1905

1968

Myrmicinae

Strumigenys membranifera Emery, 1869

2014

Myrmicinae

Strumigenys mitis (Brown, 2000)

2000

Myrmicinae

Strumigenys mutica Brown, 1949

2000

Myrmicinae

Strumigenys nanzanensis Lin & Wu, 1996

2016

Myrmicinae

Strumigenys nepalensis Baroni Urbani & De Andrade,
1994

2000

Myrmicinae

Strumigenys rofocala Bolton, 2000

2000

x

x
x [part]
x
x

x
x

x
x

x

15 of 152

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Subfamily

Scientific Name

Year First Type
New Lit.
Collected/ Locality
RecordRecorded
SG

Myrmicinae

Strumigenys roganas Bolton, 2000

2000

Myrmicinae

Strumigenys rogeri Emery, 1890

1993

Myrmicinae

Strumigenys signeae Forel, 1905

2000

Myrmicinae

Strumigenys sublaminata Brown, 1959

2017

x

Myrmicinae

Strumigenys sydorata Bolton, 2000

2017

x

Myrmicinae

Strumigenys szalayi Emery, 1897

1979

Myrmicinae

Syllophopsis australica (Forel, 1907)

2014

Myrmicinae

Syllophopsis sechellensis (Emery, 1894)

2014

x

x

Myrmicinae

Syllophopsis subcoeca (Emery, 1894)

2014

Myrmicinae

Tetramorium aptum Bolton, 1977

≤ 1990Myrmicinae

Tetramorium bicarinatum (Nylander, 1846)

1916

Myrmicinae

Tetramorium curtulum Emery, 1895

2014

x

Myrmicinae

Tetramorium eleates Forel, 1913

2017

x

Myrmicinae

Tetramorium insolens (Smith, 1861)

2015

x

Myrmicinae

Tetramorium kheperra (Bolton, 1976)

2014

x

Myrmicinae

Tetramorium lanuginosum Mayr, 1870

1916

Myrmicinae

Tetramorium meshena (Bolton, 1976)

1990

Myrmicinae

Tetramorium obtusidens Viehmeyer, 1916

1916

Myrmicinae

Tetramorium pacificum Mayr, 1870

1985

Myrmicinae

Tetramorium pulchellum Emery, 1897

1916

Myrmicinae

Tetramorium scabrum Mayr, 1879

1916

Myrmicinae

Tetramorium smithi Mayr, 1879

1916

Myrmicinae

Tetramorium tonganum Mayr, 1870

2014

Myrmicinae

Tetramorium walshi (Forel, 1890)

1912

Myrmicinae

Trichomyrmex destructor (Jerdon, 1851)

1899

Myrmicinae

Trichomyrmex mayri (Forel, 1902)

1911

Myrmicinae

Tyrannomyrmex rex Fernández, 2003

2015

Myrmicinae

Vollenhovia brevicornis (Emery, 1893) = V. fridae Forel,
1913, syn.nov.

1964

Myrmicinae

Vollenhovia minuta Viehmeyer, 1916 stat. n.

2015

Myrmicinae

Vollenhovia escherichi Forel, 1911

2016

Myrmicinae

Vollenhovia overbecki Viehmeyer, 1916

1916

Myrmicinae

Vollenhovia penetrans (Smith, 1857)

2016

Myrmicinae

Vollenhovia rufiventris Forel, 1901

1916

Ponerinae

Anochetus agilis Emery, 1901

1916

Ponerinae

Anochetus graeffei Mayr, 1870

1916

x

x
x

x

x
x
x

Ponerinae

Anochetus incultus Brown, 1978

1967

x

Ponerinae

Anochetus muzziolii Menozzi, 1932

2016

x

Ponerinae

Anochetus rugosus (Smith, 1857)

1857

Ponerinae

Anochetus tua Brown, 1978

1968

x
x

16 of 152

Wendy Y. Wang, Eunice J.Y. Soh, Gordon W.J. Yong, Mark K.L. Wong, Benoit Guénard, Evan P. Economo & Seiki Yamane

Subfamily

Scientific Name

Year First Type
New Lit.
Collected/ Locality
RecordRecordedSG

Ponerinae

Brachyponera luteipes (Mayr, 1862)

2014

x

Ponerinae

Brachyponera obscurans (Walker, 1859)

2015

Ponerinae

Buniapone amblyops (Emery, 1887)

1916

Ponerinae

Centromyrmex feae greeni Forel, 1901

1901

Ponerinae

Centromyrmex hamulatus (Karavaiev, 1925)

1973

x

Ponerinae

Cryptopone butteli Forel, 1913

1962

x

Ponerinae

Cryptopone testacea Emery, 1893

2019

Ponerinae

Diacamma geometricum (Smith, 1857)

1857

Ponerinae

Diacamma pallidum (Smith, 1858)

1989

Ponerinae

Diacamma rugosum Le Guillou, 1842

1907

x

x
x
x

Ponerinae

Diacamma vagans (Smith, 1860)

1893

Ponerinae

Ectomomyrmex leeuwenhoeki (Forel, 1896)

1916

Ponerinae

Ectomomyrmex overbecki (Viehmeyer, 1916)

1916

Ponerinae

Emeryopone buttelreepeni Forel, 1912

1968

Ponerinae

Euponera malayana (Wheeler, 1929)

1987

Ponerinae

Euponera sharpi Forel, 1901

1901

Ponerinae

Harpegnathos venator (Smith, 1858)

1974

Ponerinae

Hypoponera confinis (Roger, 1860)1912

Ponerinae

Hypoponera confinis epinotalis (Viehmeyer, 1916)

1916

Ponerinae

Hypoponera javana (Forel, 1905) stat. n.

1907

Ponerinae

Hypoponera malayana (Wheeler, 1929)

2019

Ponerinae

Hypoponera pygmaea (Forel, 1907)

2017

Ponerinae

Hypoponera singaporensis (Viehmeyer, 1916) stat. n.

1916

Ponerinae

Hypoponera truncata (Smith, 1860)

2021

x

Ponerinae

Hypoponera zwaluwenburgi (Wheeler, 1933)

1964

x

Ponerinae

Leptogenys borneensis Wheeler, 1919

1967

x

x

x
x
x
x

Ponerinae

Leptogenys chinensis (Mayr, 1840)

2016

x

Ponerinae

Leptogenys diminuta (Smith, 1857)

1964

x

Ponerinae

Leptogenys hemioptica Forel, 1901

1967

x

Ponerinae

Leptogenys iridescens (Smith, 1857)

1967

x

Ponerinae

Leptogenys kraepelini Forel, 1905

2014

Ponerinae

Leptogenys mutabilis (Smith, 1861)

1990

x

Ponerinae

Leptogenys myops (Emery, 1887)

1987

x

Ponerinae

Leptogenys pangui Xu, 2000

1975

x

Ponerinae

Leptogenys peuqueti (André, 1887)

1916

Ponerinae

Leptogenys pompiloides (Smith, 1857)

1857

Ponerinae

Leptogenys processionalis (Jerdon, 1851)

1962

Ponerinae

Mesoponera rubra (Smith, 1857)

1857

Ponerinae

Mesoponera javana (Forel, 1905) stat. n.

1969

x [part]xxx

17 of 152

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Subfamily

Scientific Name

Year First Type
New Lit.
Collected/ Locality
RecordRecordedSG

Ponerinae

Myopias mayri (Donisthorpe, 1932)

2018

x

Ponerinae

Odontomachus litoralis Wang, Yamada & Yamane, 2020

1977

Ponerinae

Odontomachus malignus Smith, 1859

2020

Ponerinae

Odontomachus rixosus Smith, 1857

1857

Ponerinae

Odontomachus simillimus Smith, 1858

1962

Ponerinae

Odontoponera denticulata (Smith, 1858)

1858

x

Ponerinae

Odontoponera transversa (Smith, 1857)

1857

x

Ponerinae

Parvaponera darwinii (Forel, 1893)

1916

x
x

Ponerinae

Platythyrea parallela (Smith, 1859)

1916

Ponerinae

Platythyrea sagei Forel, 1900

1916

Ponerinae

Ponera menglana Xu, 2001

1968

x

Ponerinae

Ponera swezeyi (Wheeler, 1933)

2005

x

Ponerinae

Pseudoneoponera havilandi (Forel, 1901)

1901

Ponerinae

Pseudoneoponera insularis (Emery, 1889)

1916

Ponerinae

Pseudoneoponera rufipes (Jerdon, 1851)

1967

x
x

Ponerinae

Pseudoneoponera tridentata (Smith, 1858)

1916

Proceratiinae

Discothyrea bryanti (Wheeler, 1917)

1967

x

Proceratiinae

Discothyrea sauteri Forel, 1912

1967

x

Proceratiinae

Probolomyrmex vieti Eguchi, Yoshimura & Yamane,
2006

2012

x
x

Proceratiinae

Probolomyrmex watanabei Tanaka, 1974

1968

Proceratiinae

Proceratium deelemani Perrault, 1981

1968

Proceratiinae

Proceratium malesianum De Andrade, 2003

2019

x

Proceratiinae

Proceratium papuanum Emery, 1897

2016

x

Pseudomyrmecinae Tetraponera allaborans (Walker, 1859)

1916

Pseudomyrmecinae Tetraponera attenuata Smith, 1877

1907

Pseudomyrmecinae Tetraponera crassiuscula (Emery, 1900)

1916

Pseudomyrmecinae Tetraponera difficilis (Emery, 1900)

1912

Pseudomyrmecinae Tetraponera extenuata Ward, 2001

1962

Pseudomyrmecinae Tetraponera inversinodis Ward, 2001

1993

Pseudomyrmecinae Tetraponera modesta (Smith, 1860)

1916

Pseudomyrmecinae Tetraponera nitida (Smith, 1860)

1916

Pseudomyrmecinae Tetraponera pilosa (Smith, 1858)

1916

Pseudomyrmecinae Tetraponera rufonigra (Jerdon, 1851)

1916

Pseudomyrmecinae Tetraponera vivax Ward, 2001

2001

Pseudomyrmecinae Tetraponera volucris Ward, 2001

2001

Total

x
x

x
x
96

121

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Table 2. List of species and subspecies last collected in Singapore > 100 years ago (from 2022).
Subfamily

Scientific Name

Latest Known
Collection Date

Dolichoderinae

Chronoxenus dalyi (Forel, 1895)≤ 1916

Dolichoderinae

Technomyrmex elatior Forel, 1902

≤ 1916

Dolichoderinae

Technomyrmex strenuus Mayr, 1872

≤ 1872

Dorylinae

Aenictus shuckardi Forel, 1901

≤ 1916

Dorylinae

Lioponera singaporensis (Viehmeyer, 1916)≤ 1916

Formicinae

Acropyga rubescens Forel, 1894

≤ 1916

Formicinae

Camponotus (Karavaievia) overbecki Viehmeyer, 1916

≤ 1916

Formicinae

Camponotus (Myrmamblys) adustus Viehmeyer, 1916

≤ 1916

Formicinae

Camponotus (Myrmamblys) bellus Forel, 1908

≤ 1916

Formicinae

Camponotus (Myrmamblys) ephippiatus Viehmeyer, 1916

≤ 1916

Formicinae

Camponotus (Myrmamblys) gretae Forel, 1902

≤ 1916

Formicinae

Camponotus (Myrmosaulus) camelinus (Smith, 1857)≤ 1857

Formicinae

Camponotus (Myrmotarsus) quadrisectus hians Forel, 1907

1897

Formicinae

Camponotus (Tanaemyrmex) festinus eximius Emery, 1900

≤ 1916

Formicinae

Camponotus (Tanaemyrmex) maculatus obfuscatus Viehmeyer, 1916

≤ 1916

Formicinae

Camponotus (Tanaemyrmex) maculatus subnudus Emery, 1889

≤ 1916

Formicinae

Camponotus (Tanaemyrmex) maxwellensis Forel, 1913

≤ 1916

Formicinae

Camponotus (Tanaemyrmex) mitis (Smith, 1858)≤ 1916

Formicinae

Camponotus (Tanaemyrmex) variegatus fuscithorax Dalla Torre, 1893

≤ 1916

Formicinae

Colobopsis badia (Smith, 1857)≤ 1916

Formicinae

Colobopsis leonardi gracilenta (Viehmeyer, 1916)≤ 1916

Formicinae

Colobopsis moeschi lygaea (Viehmeyer, 1916)≤ 1916

Formicinae

Colobopsis rothneyi (Forel, 1893)≤ 1916

Formicinae

Colobopsis rothneyi krafti (Forel, 1901)≤ 1916

Formicinae

Colobopsis vitrea angustula (Emery, 1925)≤ 1916

Formicinae

Colobopsis vitrea vittatula (Forel, 1913)≤ 1916

Formicinae

Echinopla melanarctos Smith, 1857

≤ 1857

Formicinae

Echinopla rugosa André, 1892

≤ 1916

Formicinae

Lepisiota rothneyi splendida (Viehmeyer, 1916)≤ 1916

Formicinae

Nylanderia clandestina (Mayr, 1870)≤ 1916

Formicinae

Nylanderia yerburyi (Forel, 1894)≤ 1912Formicinae

Overbeckia subclavata Viehmeyer, 1916

≤ 1916

Formicinae

Plagiolepis bicolor Forel, 1901

≤ 1916

Formicinae

Plagiolepis exigua Forel, 1894

≤ 1916

Formicinae

Polyrhachis (Campomyrma) creusa Emery, 1897

≤ 1916

Formicinae

Polyrhachis (Chariomyrma) modesta Smith, 1857

≤ 1857

Formicinae

Polyrhachis (Myrma) obesior Viehmeyer, 1916

≤ 1916

Formicinae

Polyrhachis (Myrma) orsylla Smith, 1861

≤ 1907

Formicinae

Polyrhachis (Myrma) orsylla ritsemai Mayr, 1883

≤ 1916

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Subfamily

Scientific Name

Latest Known
Collection Date

Formicinae

Polyrhachis (Myrma) striata Mayr, 1862

≤ 1893Formicinae

Polyrhachis (Myrmatopa) charaxa Smith, 1860

≤ 1912Formicinae

Polyrhachis (Myrmatopa) flavicornis Smith, 1857

≤ 1857

Formicinae

Polyrhachis (Myrmatopa) piliventris Smith, 1858

≤ 1858Formicinae

Polyrhachis (Myrmatopa) varicolor Viehmeyer, 1916

≤ 1916

Formicinae

Polyrhachis (Myrmhopla) armata defensa Smith, 1857

≤ 1857

Formicinae

Polyrhachis (Myrmhopla) bicolor aurinasis Forel, 1901

≤ 1907

Formicinae

Polyrhachis (Myrmhopla) hector Smith, 1857

≤ 1857

Formicinae

Polyrhachis (Myrmhopla) hippomanes ceylonensis Emery, 1893

≤ 1916

Formicinae

Polyrhachis (Myrmhopla) lucidula Emery, 1893

≤ 1916

Formicinae

Polyrhachis (Myrmhopla) muelleri Forel, 1893

≤ 1901

Formicinae

Polyrhachis (Myrmhopla) tibialis parsis Emery, 1900

≤ 1916

Formicinae

Polyrhachis (Myrmothrinax) saigonensis Forel, 1886

≤ 1916

Formicinae

Polyrhachis (Myrmothrinax) textor Smith, 1857

≤ 1916

Formicinae

Polyrhachis (Myrmothrinax) thrinax inconstans Viehmeyer, 1916

≤ 1916

Formicinae

Polyrhachis (Myrmothrinax) thrinax overbecki Dorow, 1995

≤ 1916

Formicinae

Polyrhachis (Polyrhachis) bellicosa Smith, 1859

≤ 1916

Formicinae

Polyrhachis (Polyrhachis) ypsilon Emery, 1887

1875

Formicinae

Pseudolasius badius Viehmeyer, 1916

≤ 1916

Formicinae

Pseudolasius ludovici Forel, 1913

≤ 1916

Formicinae

Pseudolasius martini Forel, 1911

≤ 1916

Formicinae

Pseudolasius mayri duplicatus Viehmeyer, 1916

≤ 1916

Leptanillinae

Leptanilla havilandi Forel, 1901

≤ 1901

Myrmicinae

Cardiocondyla strigifrons Viehmeyer, 1922

< 1920Myrmicinae

Crematogaster brunnea ruginota Santschi, 1928

≤ 1916

Myrmicinae

Crematogaster decamera Forel, 1910

≤ 1916

Myrmicinae

Crematogaster dohrni artifex Mayr, 1878

≤ 1916

Myrmicinae

Crematogaster egidyi szaboi Forel, 1913

1898

Myrmicinae

Crematogaster linsenmairi Feldhaar, Maschwitz & Fiala, 2016

≤ 1916

Myrmicinae

Crematogaster overbecki Viehmeyer, 1916

≤ 1916

Myrmicinae

Dilobocondyla fulva Viehmeyer, 1916

≤ 1916

Myrmicinae

Erromyrma latinodis (Mayr, 1872)≤ 1916

Myrmicinae

Monomorium atomum procax Forel, 1911

≤ 1916

Myrmicinae

Monomorium butteli demochrum Viehmeyer, 1916

≤ 1916

Myrmicinae

Monomorium hospitum Viehmeyer, 1916

≤ 1916

Myrmicinae

Myrmicaria arachnoides (Smith, 1857)≤ 1916

Myrmicinae

Myrmicaria arachnoides lutea Emery, 1900

≤ 1916

Myrmicinae

Pheidole nodgii Forel, 1905

≤ 1916

Myrmicinae

Pheidole rinae Emery, 1900

≤ 1916

Myrmicinae

Pheidole singaporensis conicollis (Emery, 1900)≤ 191619 of 152

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Subfamily

Latest Known
Collection Date

Scientific Name

Myrmicinae

Pheidole singaporensis continentis (Forel, 1911)≤ 1916

Myrmicinae

Tetramorium pulchellum Emery, 1897

≤ 1916

Myrmicinae

Tetramorium scabrum Mayr, 1879

≤ 1916

Myrmicinae

Trichomyrmex mayri (Forel, 1902)≤ 1911

Ponerinae

Anochetus rugosus (Smith, 1857)≤ 1857

Ponerinae

Centromyrmex feae greeni Forel, 1901

≤ 1901

Ponerinae

Hypoponera confinis (Roger, 1860)≤ 1912

Ponerinae

Hypoponera confinis epinotalis (Viehmeyer, 1916)≤ 1916

Ponerinae

Leptogenys pompiloides (Smith, 1857)≤ 1857

Ponerinae

Platythyrea sagei Forel, 1900

≤ 1916

Ponerinae

Pseudoneoponera havilandi (Forel, 1901)≤ 1901

Pseudomyrmecinae

Tetraponera crassiuscula (Emery, 1900)≤ 1916

Total

91

Table 3. List of species known only from Singapore (at time of writing) and not collected for > 100 years. Possiblesynonyms and their known regions of occurrence are also indicated were relevant. (*) - see comments on species
validity under ‘Remarks’ for each species in the main text.
Species/Subspecies

Possible Synonyms

Biogeographic Region/s

Camponotus maculatus obfuscatus

Camponotus maculatus, Camponotus
maculatus subnudus

Afrotropics, Indomalaya,
Malagasy, Nearctic, Neotropics, Oceania, Palearctic

Colobopsis rothneyi krafti

Colobopsis rothneyi

Indomalaya

Lepisiota rothneyi splendida*

Lepisiota rothneyi, Lepisiota rothneyi
sundaica

Indomalaya

Polyrhachis modesta

Unable to assess - only holotype queen is
known

NA

Pseudolasius badius

Unable to assess - only holotype queen is
known

NA

Leptanilla havilandi*

NA*

NA

Crematogaster egidyi szaboi*

Crematogaster egidyi, Crematogasteregidyi spinozai

Indomalaya, Palearctic

Monomorium atomum procax*

Monomorium atomum

Indomalaya, Palearctic

Monomorium butteli demochrum

Syllophopsis sechellensis; other Syllophopsis spp.

Afrotropics, Australasia,
Indomalaya, Malagasy,
Neotropics, Oceania

Monomorium hospitum*

Monomorium floricola

Afrotropics, Australasia,
Indomalaya, Malagasy, Nearctic, Neotropics, Oceania,
Palearctic

Centromyrmex feae greeni

Centromyrmex feae

Afrotropics, Indomalaya

Hypoponera confinis epinotalis

Hypoponera confinis

Australasia, Indomalaya,
Oceania

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Species/Subspecies

Possible Synonyms

Biogeographic Region/s

Pseudolasius mayri duplicatus

Unable to assess - only holotype queen
and type males are known

NA

Pseudoneoponera havilandi*

NA*

NA

Another 13 species are considered cosmopolitan tramps presumed native to Indomalaya(Wetterer 2015), although their exact native range
remains uncertain: Anoplolepis gracilipes, Cardiocondyla obscurior, Cardiocondyla wroughtonii,
Monomorium floricola, Monomorium pharaonis,
Nylanderia bourbonica, Ooceraea biroi, Paratrechina longicornis, Syllophopsis sechellensis,
Tapinoma melanocephalum, Technomyrmex vitiensis, Tetramorium bicarinatum, Tetramorium
lanuginosum. These species are common in urban
or semi-urban settings, sometimes in man-madeinfrastructure, where they often occur in abundance. Some species such as the yellow crazy
ant — Anoplolepis gracilipes — however, haveoccasionally been reported from primary or old
secondary forests (see Species Accounts).
Singapore’s complex land use history
probably helped shape a unique ant fauna comprising myriad species largely native to Indomalaya. These species tend to reside mainly in nature reserves or spontaneous vegetation fragments
scattered throughout the urban matrix or acrossoffshore islands. Aside from existing reserves, thevalue of forest remnants in supporting the native
ant fauna cannot be discounted. Secondary forests
or patches of spontaneous vegetation can be treasure troves of unexpected ant diversity. New species, such as Rhopalomastix spp. for example, were
only recently discovered in abandoned plantation
or waste woodland forests (see Species Accounts).
The nation’s first records of Lordomyrma were also
derived from abandoned wasteland forests about
to be cleared to make way for urban infrastructure.
Further, the rare arboreal genus Gesomyrmex, wasstumbled upon entirely by chance on the crown ofa large fallen tree in an urban residential district(see Species Accounts).
Future outlook
Ant diversity in Singapore is astounding – it exceeds all known records of ant diversity in other
cities worldwide at the time of writing (i.e., 6 – 213
species in 108 other cities; see Table 1 in Bras-

sard et al. 2021). Despite the impressive numbers,
the checklist remains, admittedly, incomplete. A
plethora of ant species that thrive in unexploredand more inaccessible domains such as the canopy,
deep soil layers, or restricted forests designatedas military training grounds, are possibly unaccounted for. Ants from cryptic strata usually require unique trapping techniques, and are rarely
encountered in regular ecological surveys or generalized sampling schemes. For example, elusive
hypogaeic species such as Aenictus seletarius
and Leptanilla hypdracos were discovered using
customized subterranean pitfall traps (Wong &Guénard, 2016a,c).
Substantial material that cannot be reliably identified to nominal species further exacerbate the situation. In particular, most available
specimens of hyperdiverse but taxonomically unresolved genera such as Camponotus, Nylanderia
and Tapinoma could only be differentiated to codedmorphospecies. In future when more taxonomic
insights are gained on these hyperdiverse groups
using advanced integrated approaches, the current
checklist should be revised.
The fact that even presumably degraded
forest remnants can support surprising arrays ofnative and/or new ant species, warrants an urgent
need to protect these fragments from impending
development and inevitable destruction. Many native species may likely be eliminated before they
can be discovered or described. Most species on
our list found in scattered forest fragments and notprotected reserves, also await a grim fate becauseof imminent urbanization. Precious diversity could
be forever lost with every patch of forest carelessly
cleared. Nevertheless, we hope that this compilation provides valuable insights on Singapore’s immense but potentially vulnerable insect fauna. Data
presented in the checklist can serve as a baseline
to support regional research, and also contributetowards informing future conservation decisions
in the country.

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SPECIES ACCOUNTSAMBLYOPONINAE (4 genera, 5 species)
Myopopone castanea (Smith, 1860)
Material examined. University Campus (Bukit
Timah), Oct 1970, D.H. Murphy leg., ZRC_
HYM_0000266; Mandai forest, 7 Jun 2018,
W. Wang leg., hand collection (night), ZRC_
ENT00007262.
Material not physically examined. ANIC32026171 - 026172, RWTacc.68.67 (ANIC).
Literature. Yong et al. (2017).
Localities: Mandai forest; Pulau Ubin; University
Campus (Bukit Timah).
Habitat/Ecology. This species was found in disturbed secondary forest or such forest fragments in semi-urban settings, sometimes insoil at the bases of large trees.
Mystrium camillae Emery, 1889
Material examined. Mandai Road, 8 Jun 2017,
G.W. Yong leg., ZRC_ENT00000924.
Material not physically examined.
CASENT0172325, ANTC6550 (ANIC).
Literature. None. New record.
Localities. Mandai Road – 12.5-mile Mandai
Road, near Bukit Mandai.
Habitat/Ecology. Found in secondary forest bush
litter.
Prionopelta kraepelini Forel, 1905
Material examined. Males, Prince George’s
Park Residences (NUS), 1.292389°N,
103.778694°E, 3-10 Jun 2015, M.S. Foo
and W. Wang leg., malaise trap, ZRC_
BDP0041602, 46318; Sungei Buloh Wetland
Reserve, 1°26’46.3”N 103°43’49.9”E, 7-14
Nov 2012, J. Puniamoorthy et al. leg., malaise
trap, ZRC_BDP0017406.
Material not physically examined. Unknown.
Literature. Wang et al. (2018a).
Localities. Prince George’s Park Residences (National University of Singapore, Kent Ridge
campus); Sungei Buloh Wetland Reserve.
Habitat/Ecology. This species was collected fromdisturbed secondary forest fragments in urban
or semi-urban settings, and also mangroves.

Remarks. This species was identified primarily
from males collected via malaise traps. Considering the hypogaeic nature of P. kraepelini,
it is unlikely that the species nests in actualmangroves – the tidal zone that undergoes
periodic inundation. Winged males collectedfrom mangrove malaise traps may have originated from drier grounds in adjacent backmangroves.
Stigmatomma besucheti (Baroni Urbani, 1978)
Material examined. None available.
Material not physically examined. ANIC32016286-16287(ANIC); CASENT0172194,
ANTC6446 (CASC); CASENT0195513,
ANIC32016287 (CASC – current repository).
Literature. Esteves & Fisher (2016).
Localities. Bukit Timah Nature Reserve (BTNR);
University Campus (Bukit Timah).
Habitat/Ecology. Found in Imperata grassland;also in degraded coastal hill forest on granite(i.e., BTNR).
Remarks. This species has not been recorded fromanywhere outside the Malagasy region except
for Singapore. The type series was collected in
Seychelles, specifically from La Digue island
in 1975, possibly from soil samples (Esteves& Fisher 2016). Morphological differences
have been noted between specimens from
Seychelles and Singapore, but because of inadequate material to ascertain character variation, these specimens were tentatively considered conspecific (Esteves & Fisher 2016). It isquite possible that S. besucheti may be morewidespread outside the Malagasy region, but
is rarely collected and/or identified becauseof its elusive hypogaeic nature.
Stigmatomma reclinatum (Mayr, 1879)
Material examined. Locality and collection
date unavailable, D.H. Murphy leg., ZRC_
HYM_0000265; Bukit Timah Nature Reserve,
BT06, 1.35284N, 103.77995E, 11 Jan 2017,
W. Wang leg., ZRC_HYM_0000571; Bukit
Timah Nature Reserve, 1 May 2015, W. Wangleg., ZRC_HYM_0000572; Central Catchment Nature Reserve, 10 Aug 2011, E.J.Y.
Soh leg., Gangsa 1, ZRC_HYM_0001751.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Material not physically examined. Male,CASENT0172223, ANTC6464 (ANIC).
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Central
Catchment Nature Reserve; University of Malaya (now known as the Bukit Timah campusof the National University of Singapore).
Habitat/Ecology. This species was found mainly
in primary or old/mature secondary forest,
usually in leaf litter or soil at bases of trees.
DOLICHODERINAE (7 genera, 26 species)
Chronoxenus dalyi (Forel, 1895)
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[both as ‘Bothriomyrmex wroughtoni subsp.
dalyi’].
Localities. Bukit Timah Road
Habitat/Ecology. This species was found in a garden, specifically at the foot of a hollow tree, in
passages like those made by termites.
Remarks. This species is only known from Singapore based on literature records at the timeof writing.
Chronoxenus wroughtonii (Forel, 1895)
Material examined. Alate queens, Pulau Semakau, 1°12’04.5”N 103°45’46.1”E, SMN2,
17-24 Oct 2013, J. Puniamoorthy et al. leg.,malaise trap, ZRC_BDP0015382; same locality as previous, 15-22 Nov 2012, J. Puniamoorthy et al. leg., malaise trap, ZRC_
BDP0017738; along Mandai Road, ex. Manis
javanica, 12 Jun 1985, collector unknown,
ZRC_ENT00007307 [Chronoxenus sp.1.of.
SKY]; locality unknown, 14 Apr 1993, Sk.
Yamane leg. (SKYC).
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Mandai Road; Pulau Semakau.
Habitat/Ecology. Individual workers were foundin Manis javanica (pangolin) stomach contents – the pangolin was roadkill found on
Mandai Road. Exact origins of these C.
wroughtonii specimens could not therefore

be verified, but might be postulated to be secondary forest distributed along Mandai Road.
Alate queens were also collected in replanted
mangrove forest on an offshore island landfill.
Remarks. Chronoxenus wroughtonii is currently
known from Central, South and East Asia, andparts of Southeast Asia including the Philippines, while its subspecies – C. wroughtonii
javanus (Forel, 1909), is known only fromIndonesia (Java) (antmaps.org - accessed Jan
2022; Janicki et al. 2016). We compared thespecimens with type images of both species
and subspecies on AntWeb, and found greater
similarities with C. wroughtonii. Hence, at thetime of writing we consider the two queens to
be C. wroughtonii, but this putative conclusion may be changed when more material ofboth species are made available for examination in future.
Dolichoderus affinis Emery, 1889
Material examined. Island Club, 30 Jun 1964,
D.H. Murphy leg., ZRC_HYM_0000799.
Material not physically examined. Queen types,
FOCOL2780-2781 (ZMHB). Non-types –Singapore Botanic Gardens, 5 Aug 2014, J.K.
Wetterer leg., vial #479 (JKWC).
Literature. Viehmeyer (1916), Overbeck (1924).
Localities. Queen type locality – Bukit Timah
Road. Island Club. Singapore Botanic Gardens.
Habitat/Ecology. This species was found in public
or private gardens, in one instance, under thebark of a branch of a dead tree.
Remarks. Type locality in Myanmar (Emery
1889), but queen type described from Singapore (Viehmeyer 1916).
Dolichoderus crawleyi Donisthorpe, 1917
Material examined. Namly Avenue, 28 Apr 1985,
D.H. Murphy leg., DHM-SG85-Doli2, ZRC_
ENT00027902.
Material not physically examined. Syntype;
CASENT0902970, ANTC22527 (BMNH).
Literature. Type – Donisthorpe (1917).
Localities. Specific type locality unknown. Namly
Avenue.

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Habitat/Ecology. The type series was found associated with Lecanium sp. (coccids) in hollow
stems of a Macaranga tree. Also found underbark of unknown trees.
Remarks. Type locality in Singapore.
Dolichoderus indrapurensis Forel, 1912
Material examined. None available.
Material not physically examined. ANIC32061444, RWTAcc.68.18 (ANIC).
Literature. None. New record.
Localities. Nee Soon
Habitat/Ecology. Specific information uncertain,
but possibly either swamp forest or old secondary forest inferring from locality.
Dolichoderus sulcaticeps (Mayr, 1870)
Material examined. End of Rifle Range Road,
stagnant pool, 28 Apr 1993, collector unknown, ZRC_HYM_0000246.
Material not physically examined. Male andqueen of Hypoclinea sulcaticeps, FOCOL2769-2770 (ZMHB).
Literature. Viehmeyer (1916), Overbeck (1924)[as Dolichoderus (Hypoclinea) sulcaticeps].
Localities. Bukit Timah Road; Rifle Range Road.
Habitat/Ecology. The ants were found in pasteboard nests underneath leaves; parts of nests
used to keep coccids.
Remarks. Type locality in Borneo (Mayr 1870),
but the male and queen were first describedfrom Singapore (Viehmeyer 1916).
Dolichoderus thoracicus Smith, 1860
Material examined. Rifle Range Road, Feb 2015,
R.Z.B. Quek leg., ZRC_ENT00007860; Pulau Ubin, Jalan Sam Heng, 1.412604°N,
103.99317°E, 4 Dec 2016, G.W. Yong and
M.K.L. Wong leg., ZRC_HYM_0001110; Pulau Ubin, Outward Bound School grounds,
1.416622°N, 103.99443°E, 4 Dec 2016,
G.W. Yong and M.K.L. Wong leg., ZRC_
HYM_0001111; Upper Thomson Nature Park,
9 Dec 2017, Sk. Yamane leg. (SKYC). Dealate
queens, Ulu Sembawang Park, 1.4177°N,

103.79063°E, 14 Jul 2017, W. Wang leg.,
ZRC_HYM_0001738.
Material not physically examined. Multiple
specimens in ZRC. Type of ‘Hypoclinea sellaris’, FOCOL2773 (ZMHB).
Literature. Roger (1863) [as type of ‘Hypoclinea
sellaris’]; Dill (2002).
Localities. Bukit Timah Nature Reserve; Lower
Peirce Reservoir; National University ofSingapore (Kent Ridge campus); Nee Soon;
Pulau Ubin; Rifle Range Road; Sime Road;
Ulu Sembawang Park; Upper Thomson Nature Park.
Habitat/Ecology. These ants were mostly foundin disturbed secondary and wasteland forests,
swamp forest, and also in an urban city park.
Remarks. A locally common and globally widespread tramp species, usually gregarious andnumerous where it occurs. Probably a species complex (Dill 2002), substantial morphological variation among different allopatricpopulations has been observed. At the time ofwriting, we treat these variants as conspecificuntil more compelling evidence for speciesdelineation is made available in future. If
more species are delimited from this complex,
the perceived trampy nature of D. thoracicus
may be disproved.
Iridomyrmex anceps (Roger, 1863)
Material examined. Pulau Ubin, 7 Jan 2014,
Sk. Yamane leg., ZRC_HYM_0000358– 359; Changi sand dunes, 21 Nov 1982,
D.H. Murphy leg., ZRC_HYM_0000631.
Alates, Pulau Semakau replanted mangrove,
1°12’04.5”N 103°45’46.1”E, SMN2, 6-13
Jun 2013, J. Puniamoorthy et al. leg., malaise trap, ZRC_BDP0015281; same locality
and collectors as previous, 17-24 Oct 2013,
ZRC_BDP0015381; same locality and collectors as previous, 21-28 Nov 2013, ZRC_
BDP0015369-15370; Pulau Semakau old mangrove fragment, 1°12’23.9”N 103°45’37.6”E,
SMO3, 30 Apr – 7 Mar 2013, J. Puniamoorthy
et al. leg., ZRC_BDP0016225; same locality
and collectors as previous, 22-29 Nov 2012,
ZRC_BDP0015244.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Material not physically examined. Non-types,
ANIC32-038822, 59573 (ANIC). Non-types,
J.K. Wetterer leg. (JKWC) - Kallang, Riverside Park, 29 Jul 2014, vial #372; Punggol, by MRT station, 30 Jul 2014, vial #390;
Pasir Panjang, by MRT station, 5 Aug 2014,
vial #482; Singapore Botanic Gardens, 5 Aug
2014, vial #479.
Literature. Viehmeyer (1916), Overbeck (1924).
Localities. Bukit Timah Road; Changi, Loyang
Industrial Estate; Pasir Panjang; Pasir Ris
Park; Pulau Semakau; Pulau Ubin; Punggol;
Riverside Park (Kallang); Singapore Botanic
Gardens.
Habitat/Ecology. Often found in gardens, urban parks, and disturbed secondary forests.
Winged alates collected from malaise traps
set up in both replanted and old-growth mangroves on an offshore island landfill.
Remarks. This species is afflicted with substantial taxonomic uncertainty, and different
geographic forms may actually comprise a
species complex. It is considered endemic
to the Indomalayan region where it was firstdescribed, and appears globally widespread.
It seems to be a rather resilient species, able
to adapt to and thrive in a broad range of different habitat types.
Ochetellus glaber (Mayr, 1862)
Material examined. Chestnut Avenue wasteland, 11 Jul 2009, H.K. Lua leg., LHK 505,
sweepnet, ZRC_HYM_0000182; Tuas West
Drive marshland, 31 Mar 2007, H.K. Lualeg., ZRC_HYM_0000183; Upper Thomson
Nature Park, 9 Dec 2017, Sk. Yamane leg.
(SKYC).
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Chestnut Avenue; Tuas; Upper Thomson Nature Park.
Habitat/Ecology. Associated mostly with young
secondary habitat in Singapore, including
wasteland forest fragments in semi-urban settings, and marshland. These ants were also
commonly observed at forest margins. Nests
were found in dead twigs, rotting wood orunder stones.

Remarks. This species may have been introduced
locally, though it can be considered native to
the Indo-Australian region that includes Singapore. This conjecture cannot be verified at
the time of writing because of deficient data.
Philidris cordata (Smith, 1859)
Material examined. Rifle Range Road, 6 Dec
2017, W. Wang leg., WW-SG17-Phil1,
ZRC_ENT00028313; Sungei Buloh Wetland
Reserve, 28 Mar 2018, W. Wang leg., WWSG18-Phil1, ZRC_ENT00000958; Bukit
Timah Nature Reserve, BT05, 1.35386°N,
103.77988°E, 28 Dec 2016, W. Wang leg.,
WW-SG16-010, ZRC_HYM_0000503;
Bukit Batok East Avenue 6, BB8, 1.34304°N,
103.76235°E, 28-30 Aug 2016, G.W. Yongleg., trunk pitfall trap, ZRC_HYM_0001544;
Upper Thomson Nature Park, UT5,
1.38311°N, 103.79839°E, 11-13 Sep 2016,
G.W. Yong leg., pitfall, ZRC_HYM_0001545;
Kent Ridge Park, 7 Dec 2017, W. Wang leg.,
WW-SG17-043, hand collection (day), ZRC_
ENT00014138.
Material not physically examined.
CASENT0909511, ANTC31950 (MHNG)[type of ‘Philidris cordata fuscus jactans’].
Literature. Forel (1911) [as type of ‘Philidris
cordata fuscus jactans’].
Localities. Bukit Batok East; Upper Thomson
Nature Park.
Habitat/Ecology. Found mostly in young secondary habitat in Singapore, including disturbed
scrub in parks and abandoned plantation forests, also sometimes in primary forest andmangroves. Nests have been found in fallen
twigs and behind bark of dead trees; semicarton nests were frequently found linked toextensive trunk trails made of similar substrate.
Remarks. Due to taxonomic ambiguities at thetime of writing, two subspecies — P. cordata
fusca and P. cordata protensa — have been included in this section. It is possible that thesesubspecies may be raised to species status in
future if more empirical evidence are madeavailable. Philidris cordata fusca appears
limited to primary and disturbed secondary
forests, while P. cordata protensa has only

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been found in mangroves. First local recordof this species is as type of ‘Philidris cordata
fuscus jactans (Forel, 1911)’ - now considered
an unavailable quadrinomial.
		
These ants are typically found in extreme
abundance, sometimes in semi-carton nest extensions or satellites on tree trunks or baseof trees, often linked by pervasive/extensive
trunk trails. Arguably, such trunk trails andcarton nest building behaviour may be only
demonstrated by one subspecies — P. cordata
fusca.
Philidris laevigata (Emery, 1895)
Material examined. Rifle Range Road, 6 Dec
2017, W. Wang leg., hand collection (day),
WW-SG17-045, ZRC_ENT00014140.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[as ‘Iridomyrmex levigatus’]; Baroni Urbani(1977) [as ‘Iridomyrmex laevigatus jactans’].
Localities. Ayer Terjun (archaic place name); Botanic Gardens; Bukit Timah Nature Reserve;
Chua Chu Kang; Pulau Bulang; Rifle Range
Road; Ulu Beri
Habitat/Ecology. Found mostly in primary or mature secondary habitat in Singapore, including
native-dominated secondary forest. Colonies
have been observed in (hollows of) bamboo
and hollow branches, in bark fissures or underloose bark of trees, covered up with earth. The
ants typically form carton trunk trails near andaround bases of trees.
Philidris myrmecodiae (Emery, 1887)
Material examined. MacRitchie Reservoir, 6
Jan 2014, Sk. Yamane leg., SG14-SKY-02,
ZRC_HYM_0000468.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. MacRitchie Reservoir
Habitat/Ecology. This species appears associatedwith primary and/or old/mature secondary
forests. Nests were found in dead twigs or
branches.
Remarks. This species was originally described
as a variant of Iridomyrmex cordatus (nowknown as Philidris cordata), and was ob-

served to consist of many ‘intermediary’forms with broad colour variation. We areunable to distinctly differentiate between Philidris cordata protensa and this species basedon original descriptions and available typeimages online, especially since both species
have been reported to have broad intra-species
variation including light- and dark-coloured
morphs. With more compelling evidence infuture, it is possible that the material treated as
P. myrmecodiae in this study may be P. cordata protensa or other species closely affiliatedwith P. cordata. Workers tend to be numerous
and aggressive where they occur; trunk trails
often extensive and constructed pervasively
around the central colony location.
Tapinoma andamanense capsincola Forel, 1911
Material examined. Males and workers, I-cubebuilding (NUS), 1.29347, 103.77633, MaySep 2015, M.S. Foo & W. Wang leg., malaise
trap, ZRC_BDP (multiple); Prince George’s
Park Residences (NUS), 1.29239, 103.77869,
1-8 Apr 2015, same collectors as previous,
ZRC_BDP0046251; University Hall (NUS),
1.29711, 103.77658, 1-16 Sep 2015, samecollectors as previous, ZRC_BDP0042894.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924).
Localities. Bukit Timah Road; National Universityof Singapore campus (multiple locations).
Habitat/Ecology. Colonies were found in a garden, specifically in a dry branch of a Jambu
tree, in withered bamboo, and in hollow twigs
of mangosteen trees. More recent specimens
were collected in malaise traps mostly set up
in disturbed secondary forest fragments, insemi-urban settings.
Remarks. At the time of writing, Tapinoma andamanense Forel, 1903 is only known by its
holotype queen collected from Andaman Island, India. Without queen specimens fromSingapore for comparison, we cannot justify
the raising of the subspecies capsincola tospecies. According to original descriptionsof T. andamanense capsincola, however, thesubspecies queen has a somewhat shorter headthat is broader posteriorly.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Tapinoma glaucum (Viehmeyer, 1916)
Material examined. Singapore, locality and collection date not specified, D.H. Murphy leg.,
ZRC_HYM_0000617.
Material not physically examined. Type, FOCOL2860 (ZMHB).
Literature. Type - Viehmeyer (1916) [as type of
Semonius glaucus]; Overbeck (1924) [as Semonius glaucus].
Localities. Bukit Timah Road [type]
Habitat/Ecology. The type series was collectedfrom a garden.
Remarks. Type locality in Singapore. While exact collection date is unknown, we estimate
the ZRC specimen to have been collected
between 1965 to 1990, when collector D.H.
Murphy was active. This minute species may
arguably be distinguished from other similarcongeners by the following combination:1)
its overall dark bluish-greenish shimmer, 2)very finely, densely reticulate head and bodysurfaces, 3) mesosoma slightly shorter thanhead, 4) pronotum in dorsal view nearly twice
as wide as long, 5) funicular segments 2-4(segment 1 being basal-most) distinctly wider
than long.
Tapinoma indicum Forel, 1895
Material examined. Alates, University Town
(NUS), 1.306222°N, 103.774583°E, 2027 May 2015, M.S. Foo and W. Wang leg.,malaise trap, ZRC_BDP0045129; same locality and collectors as previous, 1-8 Jul
2015, malaise trap, ZRC_BDP0045948;
Prince George’s Park Residences (NUS),
1.292389°N, 103.778694°E, 17-24 Jun 2015,
M.S. Foo and W. Wang leg., malaise trap,
ZRC_BDP0044384.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924).
Localities. Bukit Timah Road; Prince George’s
Park Residences, University Town (both partof the National University of Singapore campus).
Habitat/Ecology. This species was found in a garden - colony in decaying wood on the ground,in tree stumps slightly above ground surface,nesting between stem and loose bark, closed

up with earth. Alates were found in disturbed
secondary or wasteland forest patches, also
grassy patches adjoining urban human dwellings.
Remarks. Species similar to T. melanocephalum,
but may be differentiated partly by its generally lighter brown (head and mesosoma notblackish as in the latter) and relatively more
uniform body colour. The two species can also
be distinguished based on funicular segments– the third funicular segment (numbered fromthe basalmost segment) in T. indicum is wider
than long, whereas in T. melanocephalum thesame segment is distinctly longer than wide.
In addition, funicular segments 4-8 of T. indicum are each at least as wide as long, while in
T. melanochepalum these segments are clearly
longer than wide.
Tapinoma melanocephalum (Fabricius, 1793)
Material examined. Bukit Timah Nature Reserve, BT06, 1.35284°N, 103.77995°E, 11
Jan 2017, W. Wang leg., WW-SG17-005,
ZRC_HYM_0000573; queen, Lee Kong Chian Natural History Museum, level 1M lab, 14
Nov 2018, J. Tan leg., ZRC_ENT00007577;
males, Prince George’s Park Residences(NUS), 1.292389N, 103.778694E, 20-27 May
2015, M.S. Foo and W. Wang leg., malaise
trap, ZRC_BDP0046763; same locality andcollectors as previous, 3-10 Jun 2015, malaise trap, ZRC_BDP0041599; Sungei Buloh,
8 Jan 2014, Sk. Yamane leg., SG14-SKY-33(SKYC).
Material not physically examined. Syntype
of Myrmica pellucida, CASENT0903062,
ANTC22648 (BMNH). Non-types, J.K. Wetterer leg. (JKWC) – Bishan, by MRT station,
31 Jul 2014, vial #424; City Hall, by church,
29 Jul 2014, vial #374; Harbourfront, JulAug 2014, vials #385, 472; Hougang, 30 Jul
2014, vial #402; Kallang, Riverside Park, 29
Jul 2014, vial #372; Katong, 27-28 Jul 2014,
vials #328, 332, 345; Marine Parade, Jul-Aug
2014, vials #358, 368, 435; Mountbatten, by
MRT station, 4 Aug 2014, vial #465; Outram
Park, by MRT station, 29 Jul 2014, vial #380;
Pasir Panjang, by MRT station, 5 Aug 2014,
vial #483; Punggol, 30 Jul 2014, vials #390,

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392; Queenstown, 29 Jul 2014, vial #383; Serangoon, 30 Jul 2014, vial #389; Tai Seng, 31
Jul 2014, vial #422.
Literature. Viehmeyer (1916), Overbeck (1924);
Shattuck (1994); Tan & Corlett (2012); Wanget al.(2018a).
Localities. Bishan; Bukit Timah Nature Reserve;
Bukit Timah Road; City Hall; Harbourfront;
Hougang; Katong; Marine Parade; Mountbatten; National University of Singapore campus;
Outram Park; Pasir Panjang; Pulau Semakau;
Pulau Ubin; Punggol; Queenstown; Riverside
Park (Kallang); Serangoon; Sungei Buloh; Tai
Seng.
Habitat/Ecology. Found in a broad range of habitats. Garden – colonies found under loose bark,closed up with earth, and in branches lying
on the ground. Also collected from disturbed
secondary forests and mangrove backforests.
In mangroves, the ants were found nesting ina dead twig on a tree. Often observed and/orcollected in urban settings, such as parks andinside household compounds or residences.
Remarks. A globally widespread tropical tramp
species with unknown native range. Commonly known as the ‘ghost ant’, alluding to
its pale gaster and legs that contrast sharply
with blackish brown head and mesosoma.
Technomyrmex albipes (Smith, 1861)
Material examined. Pulau Ubin village, 7 Jan
2014, Sk. Yamane leg., SG14-SKY-17, ZRC_
HYM_0000437; Sungei Buloh, 8 Jan 2014,
Sk. Yamane leg., ZRC_HYM_0000438. Male,
Prince George’s Park Residences (NUS),
1.292389N, 103.778694E, 19-26 Aug 2015,
M.S. Foo and W. Wang leg., malaise trap,
ZRC_BDP0047424.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924);
Bolton (2007).
Localities. Bukit Timah Hill; Bukit Timah Road;
Nee Soon; Prince George’s Park Residences(NUS); Pulau Ubin; Sungei Buloh.
Habitat/Ecology. Colonies were found in a garden — in rotten wood, hollow branches andwithered bamboo. The ants were also foundin disturbed secondary, swamp forests, andmangroves, where a colony was found in adead twig on a tree.

Remarks. Known to be a highly successful tramp
species able to nest and forage in both groundlevel and arboreal environments (Bolton2007), also commonly found in parks andother urban settings elsewhere.
Technomyrmex difficilis Forel, 1892
Material examined. Alates and workers, Mandai
mangroves, 23 May 1979, D.H. Murphy leg.,
DHM-SG79-001, ZRC_ENT00048398.
Material not physically examined. Unknown.
Literature. Bolton (2007).
Localities. Botanic Gardens; Kent Ridge; Mandai
mangroves.
Habitat/Ecology. This species was mostly foundin urban parks and gardens. In mangroves, theants were found nesting in decayed branchesof Avicennia sp.
Remarks. Frequently confused with T. albipes, but
can be distinguished from the latter species
by the presence of setae on the head dorsum
behind the level of the posterior margin of theeye, also a longer and more slender promesonotum (Bolton 2007). Considered native
to the Malagasy region, the species is suggested to have started spreading across Southeast Asia and Oceania more than 60 years ago(Wetterer 2013). This species is estimatedto have been first collected from Singapore
around or earlier than 1948 (Wetterer 2013).
Technomyrmex elatior Forel, 1902
Material examined. None available.
Material not physically examined. Syntypes of‘Technomyrmex albipes var.cordiformis’, FOCOL2874 – 2875 (ZMHB).
Literature. Viehmeyer (1916), Overbeck (1924)[both as ‘Technomyrmex albipes var.cordiformis’]; Bolton (2007).
Localities. Ayer Terjun (archaic place name)
Habitat/Ecology. A colony of this species wasfound in a broken and hollowed piece of atree branch, forest type unknown.
Remarks. This species is only known from one
locality in Singapore — Ayer Terjun; area now
urbanized and developed.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Technomyrmex horni Forel 1912
Material examined. None available.
Material not physically examined.
CASENT0249786, PSW10256-3 (PSWC,
Davis).
Literature. Bolton (2007).
Localities. Bukit Timah
Habitat/Ecology. Specimen from Bukit Timah
indicated to be from rainforest - possibly the
specific location should be Bukit Timah Nature Reserve.
Remarks. This species and T. schimmeri are currently the only members of a complex withindistinct inter-species boundaries (Bolton2007). The two species are almost indistinguishable morphologically, taking into account the morphological variability among
different populations per species. Should morematerial and other forms of evidence, such
as DNA, be made available for analyses infuture, this treatment may be revised.
Technomyrmex kraepelini Forel, 1905
Material examined. Upper Peirce Reservoir,
10 Jan 2014, Sk. Yamane leg., SG14SKY-53, ZRC_HYM_0000464; same locality and collection data, SG14-SKY-51,
ZRC_HYM_0000465; Lower Peirce Reservoir, 28 Jul 1990, H.K. Lua leg., ZRC_
HYM_0000229. ZRC_BDP0016044;
University Town (NUS), 1.306222°N,
103.774583°E, 8-15 Jul 2015, M.S. Foo
and W. Wang leg., malaise trap, ZRC_
BDP0046474. Alate, Pulau Ubin, Sg. Assam,
1.42126°N, 103.94836°E, PU3, 27 Oct – 3
Nov 2012, J. Puniamoorthy et al. leg., malaise trap, ZRC_BDP0016044.
Material not physically examined. ANIC32044947, D3-9 (ANIC).
Literature. Bolton (2007); Tan & Corlett (2012);
Wang et al. (2018a).
Localities. Bukit Batok; Bukit Timah; Lower
Peirce Reservoir; Mandai Road; Seletar; Tuas
West Drive; University Town (NUS); Upper
Peirce Reservoir.
Habitat/Ecology. Typically found foraging at
ground level, often on/in leaf litter. Can befound in different types of secondary forest:

abandoned plantation, native-dominated andwaste woodland. Sometimes also found inmarshland.
Remarks. Winged alates of this species have been
identified from samples of malaise traps set up
in disturbed spontaneous vegetation patches
in urban or semi-urban settings, and in mangroves.
Technomyrmex pratensis (Smith, 1860)
Material examined. Locality and collector
data unavailable, NS138C, 1994, ZRC_
HYM_0000112; MacRitchie Reservoir, 6 Jan
2014, Sk. Yamane leg., SG14-SKY-01, ZRC_
HYM_00000467; Bukit Timah Nature Reserve, BT06, 1.35284°N, 103.77995°E, 11 Jan
2017, W. Wang leg., ZRC_HYM_0000580;
Upper Thomson Nature Park, 9 Dec 2017,ex.leaf litter, Sk. Yamane leg., SG-17-SKY-37,-40 (SKYC).
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; MacRitchie Reservoir; Upper Thomson Nature
Park.
Habitat/Ecology. In mature and/or abandoned
plantation secondary forests. Nests were
found in leaf litter, rotting wood or living trees
such as Macaranga sp.
Technomyrmex schimmeri Viehmeyer, 1916
Material examined. Upper Thomson Nature
Park, 9 Dec 2017, W. Wang leg., ZRC_
ENT00000776; Kent Ridge, KR02_05, 2017,
W.N. Lam leg., ZRC; Nee Soon, 17 Aug 2017,
W.N. Lam leg., ZRC; MacRitchie Reservoir,
9 Jan 2014, ex. leaf litter, Sk. Yamane leg.,SG14-SKY-36 (SKYC).
Material not physically examined. Types, including 1 queen, FOCOL2877 – 2879 (ZMHB).
Literature. Type – Viehmeyer (1916). Overbeck(1924); Bolton (2007).
Localities. Type locality – Bukit Timah Road.
Bukit Batok; Kent Ridge; near I-cube building; MacRitchie Reservoir; Nee Soon; Upper
Thomson Nature Park.

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Habitat/Ecology. The type series was found inwithered bamboo in a garden. This species
was also collected from scrubland, and abandoned plantation secondary forest, usually inleaf litter sometimes at bases of trees. Specimens have also been collected from liquid
contents of Nepenthes pitchers.
Remarks. Type locality in Singapore.
Technomyrmex strenuus Mayr, 1872
Material examined. None available.
Material not physically examined. Syntype CASENT0903048, ANTC22633 (BMNH).
Literature. Syntype – Mayr (1872). Bolton(2007).
Localities. Unavailable.
Habitat/Ecology. Unknown in the Singapore context.
Remarks. Original type locality is in Borneo(Mayr 1872), but specimen from Singapore
also designated as syntype.
Technomyrmex textor Forel, 1909
Material examined. Mandai Road, M1D,
1.41258°N, 103.79839°E, 8 Jun 2017, W.
Wang leg., ZRC_ENT00000951; Mandai
Road, 21 Nov 2016, G.W. Yong leg., ZRC_
HYM_0001638.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Mandai Road.
Habitat/Ecology. Native-dominated secondary
forest. Some foragers collected behind barkof Macaranga gigantea tree.
Remarks. This species is distinguishable by its relatively small size and pale light-brown body,characters atypical of most other congeners.
Technomyrmex vitiensis Mann, 1921
Material examined. Mandai Road, 1.41468,
103.79779, ZRC_ENT00048978; alates, Pulau Semakau New Fragment, 2012 – 2013, J.
Puniamoorthy et al. leg., malaise trap, ZRC_
BDP multiple; Pulau Semakau Old Fragment, SMO3, Nov 2012, same collectors asprevious, malaise trap, ZRC_BDP0016539;
queen, Sungei Buloh Wetland Reserve, SB2,

1°26’47.7”N, 103°43’49.9”E, 12-19 Dec
2012, same collectors as previous, malaise
trap, ZRC_BDP0016345.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Mandai Road; Pulau Semakau; Sungei
Buloh Wetland Reserve.
Habitat/Ecology. This species was collected fromboth old growth and newly replanted mangroves, and in native-dominated secondary
forest more inland.
Remarks. Winged alates collected using malaisetraps set up in mangrove forests on an offshoreisland landfill – Pulau Semakau, part of theMangrove Insect Project (MIP) 2012 – 2013.
DORYLINAE (7 genera, 12 species)
Aenictus camposi Wheeler & Chapman, 1985
Material examined. Nee Soon Swamp Forest, atbase of pipeline, 01°23.062’N, 103°48.684’E,
Feb 2015, W. Wang leg., Cryo-Acam_NS1NS2, ZRC; Bukit Timah Nature Reserve, 28Mar. (year unavailable), D.H. Murphy leg.,C2-7, ZRC_HYM_0000005-6; Bukit Timah
Nature Reserve, 24 Aug 1967, D.H. Murphyleg., Pitfall B8-31, ZRC_HYM_0000256.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Nee Soon
Forest.
Habitat/Ecology. Found in forest, sometimes inleaf litter or part of bivouac under soil. Also
collected from ground pitfall traps.
Aenictus cylindripetiolus Jaitrong & Yamane,
2013
Material examined. Central Catchment Nature
Reserve (fringe), 1.41138°N, 103.80653°E,
29 Nov 2016, G.W. Yong & Ben Ho leg.,
ZRC_HYM_0000293.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Central Catchment Nature Reserve
Habitat/Ecology. This species was discovered
in a native-dominated secondary forest fragment, forming a fast-moving ant trail underleaf litter.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Remarks. Active ant trail found in the evening.
Aenictus gracilis Emery, 1893
Material examined. None available.
Material not physically examined. ANIC32059592, Shattuck.1345388838 (ANIC).
Literature. None. New record.
Localities. Bukit Timah Nature Reserve
Habitat/Ecology. The original label indicates ‘degraded coastal hill forest on granite’, but thisis not an entirely accurate description of theactual habitat type in BTNR. The ants were
probably found in either primary or old/mature secondary forest.
Aenictus seletarius Wong & Guénard, 2016
Material examined. Holotype, Seletar Trail,ca. 40 m, 1.392467°N, 103.801308°E,
25.vii.2015, M.K.L. Wong leg., subterranean pitfall trap, ZRC_ENT00000663; Seletar Trail, 1.39625°N, 103.802994°E, 15 Jul
2015, M.K.L. Wong leg., SPT15, pitfall trap,
ZRC_ENT00000669.
Material not physically examined. Unknown.
Literature. Type – Wong & Guénard (2016).
Localities. Seletar Trail (near Central Catchment
Nature Reserve)
Habitat/Ecology. This small species was collectedfrom rainforest, deep in soil, using subterranean pitfall traps.
Remarks. Type locality in Singapore.
Aenictus shuckardi Forel, 1901
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924).
Localities. Bukit Timah Road
Habitat/Ecology. Garden.
Remarks. In literature, this species was identified
only from a single male collected at a lit-lamp.
No workers are known to have been collectedfrom the same locality till present day.

Aenictus yamanei Wiwatwitaya & Jaitrong,
2011
Material examined. Nee Soon forest, 1.382794°N,
103.806499°E, 23 Oct 2017, W.N. Lam leg.,NS09-GL, ZRC_ENT00000748.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Nee Soon forest
Habitat/Ecology. Swamp forest, found in contentsof Nepenthes gracilis pitcher.
Remarks. Species identified from only a single
worker in pitcher contents.
Dorylus laevigatus (Smith, 1857)
Material examined. Lower Peirce Reservoir, ForestA, 7 Aug 1990, Zoology Dept. 3rd year project,
ZRC_ENT00000191; Central Catchment Nature Reserve, Venus Loop, SPT15, 1.355491,
103.815777, 15 Jun 2015, M.K.L. Wong leg.,
subterranean pitfall trap, ZRC_ENT00000671;
Rifle Range Road, 6 Dec 2017, Sk. Yamaneleg., SG17-SKY-02, ZRC_ENT00000879.
Male alates; 3 Pandan Valley, 16 Sep 2016,
Y.C. Tay leg., ZRC_HYM_0001734; Chun Tin
Road, 24 Dec 2009, C.M. Yang leg., ZRC_
ENT00014107-14108; Ang Mo Kio Drive,
ITE Headquarters, 29 Sep 2020, H.F. Loo& S.K. Koh leg., on carpet in pantry, ZRC_
ENT00028565; Pasir Panjang Road, 20 Jan
2021, J. Ho leg., ZRC_ENT00047779.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924) [asDorylus (Dichthadia) levigatus var. breviceps].
Localities. (Only localities where workers were
found are given here, as male occurrences are
not accurate inferences of nest locations) Central Catchment Nature Reserve; Lower Peirce
Reservoir; Rifle Range Road.
Habitat/Ecology. Primary and/or mature secondary forest — colonies were found in soil.
Remarks. Considered a kind of Old World ‘army
ant’ — a general term for ants characterizedby cooperative group foraging/predatory tactics and nomadic behaviour. Solitary malealates have been reported from urban residential or commercial areas away from forested
areas — these are isolated incidences of males
dispersed far from their original nest sources.

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Lioponera hewitti (Wheeler, 1919)
Material examined. Nee Soon forest, 30 Dec
2014, W. Wang leg., ZRC_ENT00007302.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Nee Soon
Habitat/Ecology. This species was found in secondary swamp forest, where individuals werecollected from rotting wood.
Remarks. Slight morphological differences fromL. hewitti in its original description, but thismay be variation between allopatric populations, thus we treat the Singapore specimens
as conspecific to those from Borneo. This tentative species record may be changed should
new evidence be made available in future indicating heterospecificity.
Lioponera singaporensis (Viehmeyer, 1916)
Material examined. None available.
Material not physically examined. Syntype of
Cerapachy singaporensis, CASENT0911265,
ANTC33507 (NHMB). Types of Phyracaces
singaporensis, FOCOL0335-336 (DEIC); FoCol0807-0809 (ZMHB).
Literature. Type – Viehmeyer (1916); Overbeck(1924) [both as Phyracaces singaporensis].
Localities. Bukit Timah Road
Habitat/Ecology. A colony of this species wasfound in the hollow branch of a mangosteen
tree, in a garden.
Remarks. Type locality in Singapore.
Ooceraea biroi (Forel, 1907)
Material examined. Mandai, 1.40699, 103.77853,
19 Sep 2015, M.K.L. Wong leg., leaf litter,
Berlese extraction, ZRC_ENT00000670;
Pearl’s Hill City Park, 21 Aug 2017, J. Choileg., Jchoi-SG17-01, ZRC_HYM_0001748;
MacRitchie forest, 30 Aug 1982, D.H. Murphy leg., ZRC_ENT00027945.
Material not physically examined. Paralectotype of Cerapachys (Syscia) biroi,
CASENT0907059, ANTC27237 (MHNG).
Syntype of C. biroi, CASENT0916710,
ANTC44062 (HNHM Budapest).

Literature. Type – Forel (1907) [Cerapachys(Syscia) biroi].
Localities. MacRitchie; Mandai; Pearl’s Hill City
Park.
Habitat/Ecology. Both mature and young secondary forests, also cultivated land in urban orsemi-urban settings such as parks. Nests were
often found in soil under rocks, or in old rotten
logs typically filled with soil or abandoned termite nest residue. Individuals were sometimes
collected from leaf litter.
Remarks. Type locality in Singapore.
Parasyscia dohertyi (Emery, 1902)
Material examined. Bukit Timah Nature Reserve,
Feb-Mar 1968, D.H. Murphy leg., A122-8,
ZRC_HYM_0000724
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve
Habitat/Ecology. A single worker was found ineither primary or old/mature secondary forest.
Remarks. This is the first local record of Parasyscia. The specimen mostly confers with the
holotype of P. dohertyi, based on online typeimages and descriptions in Wilson (1959).
However, we observed a few differences between the Singapore specimen and the holotype that may warrant a reassessment ofspecies in future, should more local material
be available: 1) head slightly wider, CI 91
compared to CI 88 in the holotype; 2) posteroventral corner of head projected distinctly
as rounded lobe (projection more shallow inthe holotype); 3) generally smaller in size,
petiole and postpetiole widths 0.58 and 0.68
mm respectively (0.66 and 0.80 mm in theholotype).
Simopone bakeri Menozzi, 1926
Material examined. Nee Soon swamp, 13
Jun 1994, D.H. Murphy leg., ZRC_
HYM_0000831.
Material not physically examined. Holotype(IEUB). Dealate queen, CASENT0173045,
ANTC8121 (MCZ).
Literature. Type – Menozzi (1926). Taylor (1966);
Brown (1975); Bolton & Fisher (2012).

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Localities. Nee Soon Swamp Forest
Habitat/Ecology. This arboreal species is associated with primary and/or mature secondary
forests in Singapore, including swamp forest.
Remarks. Rarely observed or collected mainly
because of its apparent arboreal nature. Type
locality in Singapore.
Syscia cf.sp.3.of.SKY (*excluded from official
named species count)Material examined. Queen and worker, Mandai
Lake Road, 1°24’36.5”N, 103°47’04.0”, 53m,
18 July 2019, J.S. Tan & N.L. Chin leg., Winkler extraction, ZRC_ENT00028290.
Material not physically examined. Unknown.
Literature. None. New genus record.
Localities. Mandai Lake Road
Habitat/Ecology. The ants were found in disturbed mature secondary forest, in leaf litterand upper soil substrate.
Remarks. This is the first record of the genus
Syscia in Singapore, though we were unfortunately unable to identify it to named species. The species confers mostly with Syscia
sp.3.of.SKY. Of known species, it looks mostsimilar to Syscia chaladthanyakiji Jaitrong etal., 2020, but with nine antennal segments (instead of 11 in S. chaladthanyakiji), and headwith strong posteroventral lamellate flange.
ECTATOMMINAE (2 genera, 8 species)
Holcoponera cribrata (Emery, 1900)
Material examined. Central Catchment Nature
Reserve, 14 Sep 2011, E.J.Y. Soh leg., MainddiT.15, ZRC_ENT00028674; Bukit Timah,
1.35130, 103.77859, 31 Jul 2015, M.K.L.
Wong leg., pitfall trap, ZRC_ENT00000673.
Material not physically examined. Specimens
collected by D.H. Murphy and R.W. Taylor
from Peirce Reservoir in 1988, examined by
Lattke (2004) but depositories unknown.
Literature. Lattke (2004) [as Gnamptogenys
cribrata].
Localities. Bukit Timah, Central Catchment Nature Reserve; Peirce Reservoir.
Habitat/Ecology. This species was found in primary or mature secondary forests in Singapore.

Remarks. Genus initially determined as Gnamptogenys, but present species combination
generated in Camacho et al. (2022) based on
phylogenetic evidence.
Stictoponera bicolor (Emery, 1889)
Material examined. Khatib wasteland, 1.41122,
103.82862, 20 Apr 2016, M.K.L. Wong leg.,
ZRC_ENT00000672.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Khatib
Habitat/Ecology. This species was found in a
waste woodland forest.
Remarks. At the time of writing, this species isknown from only one locality in Singapore.
Genus initially determined as Gnamptogenys,
but present species combination revived in
Camacho et al. (2022) based on phylogenetic
evidence.
Stictoponera binghamii (Forel, 1900)
Material examined. Mandai, Bird Park Buffer,
17 Jul 2019, J. Tan & N. Chin leg., JTNCMIS-002, Mandai Insect Survey, ZRC_
ENT00013543; Bukit Timah forest, Apr 1968,
D.H. Murphy leg., ZRC_HYM_0000611-612;
same locality as previous, 17 Mar 1967, D.H.
Murphy leg., ZRC_HYM_0000615-616.
Material not physically examined. Type male –FOCOL0867 (ZMHB).
Literature. Viehmeyer (1916), Overbeck (1924)[both as Stictoponera binghami (sic)].
Localities. Bukit Timah; Mandai.
Habitat/Ecology. Found mostly in primary or
mature secondary forests, as well as young
secondary habitat in Singapore. Individuals
have been collected from forest litter and soil
samples; nests can be found in cavities offallen rotting/decaying wood or branches. In
one instance, a nest was found in a large fallen
rotting branch alongside a Leptogenys colony.
Remarks. The first local record of this species wasa male type specimen identified and describedby Viehmeyer (1916), based on material collected by Hans Overbeck. Genus initiallydetermined as Gnamptogenys, but present
species combination revived in Camacho etal. (2022) based on phylogenetic evidence.

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Stictoponera coxalis (Roger, 1860)
Material examined. Bukit Timah Nature Reserve, 25 May 1967, D.H. Murphy leg., ZRC_
HYM_0000250; Mandai Road, 5 Dec 2016,
G.W.J. Yong leg., ZRC_HYM_0001005;
Dairy Farm, 22 Jun 2012, Zestin Soh leg.,
ZRC_ENT00028676.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Dairy
Farm; Mandai Road.
Habitat/Ecology. Found in primary or mature secondary forests in Singapore, including nativedominated secondary forest. Nests were foundin fallen rotting wood and leaf litter.
Remarks. Some examined ZRC material wereoriginally identified as ‘Gnamptogenys costata (Emery, 1889)’, but that was considereda junior synonym of ‘G. coxalis’ (Lattke &Delsinne 2016). Genus initially determined as
Gnamptogenys, but present species combination revived in Camacho et al. (2022) basedon phylogenetic evidence.
Stictoponera gabata (Lattke, 2004)
Material examined. Bukit Batok East Avenue
6, 1.34218, 103.76316, 27-29 Sep 2016,
G.W.J. Yong et al. leg., BB2GPc422, ZRC_
ENT00028675.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Batok East
Habitat/Ecology. Abandoned plantation secondary forest.
Remarks.This species is currently only known
from a single specimen in Singapore. Genus
originally described as Gnamptogenys, but
present species combination generated in
Camacho et al. (2022) based on phylogenetic
evidence.
Stictoponera hyalina (Lattke, 2004)
Material examined. None available.
Material not physically examined. Holotype –ANIC32-017682, Ab2-10 (ANIC). Non-type– ANIC32-017682, Ab2-10 (ANIC).

Literature. Type - Lattke (2004) [as Gnamptogenys hyalina].
Localities. Bukit Timah Nature Reserve
Habitat/Ecology. Associated with primary and/or
mature secondary forest; small colony foundin dry bracket fungus on log.
Remarks. Type locality in Singapore. Originallabel describes habitat as ‘degraded coastal
hill forest on granite’, but this does not accurately describe the inland BTNR forest. Genus initially determined as Gnamptogenys,
but present species combination generated in
Camacho et al. (2022) based on phylogenetic
evidence.
Stictoponera ortostoma (Lattke, 2004)
Material examined. Bukit Timah Forest, 17 Mar
1967, D.H. Murphy leg., forest litter/soil, Berlese extraction, #N-13, ZRC_HYM_0000614– 616.
Material not physically examined. Holotype– ANIC32-017689, N-13Ab4-20 (ANIC).
Non-type – ANIC32-017690, N-13Ab4-20(ANIC).
Literature. Type - Lattke (2004) [as Gnamptogenys ortostoma].
Localities. Bukit Timah Nature Reserve
Habitat/Ecology. Associated with primary and/or
mature secondary forest; in leaf litter.
Remarks. Type locality in Singapore. Originallabel describes habitat as ‘degraded coastal
hill forest on granite’, but this does not accurately describe the inland BTNR forest. Genus initially determined as Gnamptogenys,
but present species combination generated in
Camacho et al. (2022) based on phylogenetic
evidence.
Stictoponera posteropsis Gregg, 1951
Material examined. Bukit Timah Nature Reserve,collection date unknown, D.H. Murphy leg.,
pitfall B7-2, ZRC_HYM_0000613.
Material not physically examined. FMNHINS0002821931, FMHD_81-282 (FMNH).
Literature. None. New record.
Localities. Bukit Timah Nature Reserve

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Habitat/Ecology. Associated with primary and/or
mature secondary forests. Individuals can befound on the forest floor, and in tree buttresslitter. Genus initially determined as Gnamptogenys, but present species combination
revived in Camacho et al. (2022) based on
phylogenetic evidence.
FORMICINAE (19 genera, 117 species)
Acropyga acutiventris Roger, 1862
Material examined. Bukit Timah Nature Reserve,near BT08, 31 May 2017, W. Wang leg.,
WW-SG17-025, ZRC_ENT00000971; Central Catchment Nature Reserve, 16 Feb 2016,
G.W.J. Yong leg., ZRC_HYM_0000996;
Prince George’s Park Residences (National University of Singapore), 1.29239,
103.77869, 24 Mar – 1 Apr 2015, M.S. Foo &W. Wang leg., malaise trap, NUS0001, ZRC_
BDP0044267; Mandai Lake Road, Northern
Node buffer area (Rainforest Park), MIS_
L02_LL, 1°24’36.5”N, 103°47’04.0”, 53 ma.s.l., 18 Jul 2019, J.S. Tan & N.L. Chin leg.,
Winkler extraction, ZRC_ENT00028297;
same locality as previous, 8 Sep 2020, N.
Chin, L. Tan & C. Goh leg., NCTG_MIS_022,
ZRC_ENT00047801.
Material not physically examined.
ANIC32-018494, Shattuck1345243946(ANIC); ANIC32-018649, Shattuck1345244635/1345244638/1345244641(ANIC).
Literature. LaPolla (2004).
Localities. Bukit Timah Nature Reserve; Central
Catchment Nature Reserve; Mandai Lake
Road; Prince George’s Park Residences(NUS); Singapore Botanic Gardens.
Habitat/Ecology. Usually associated with primary and/or secondary forests at varying
levels of disturbance in Singapore, including back mangrove forest, and disturbed but
well-developed secondary forest fragments inurban or semi-urban settings (e.g. parkland).
Nests were found in soil, soil mounds or rotten
trunks and logs; individuals were often foundat high densities from leaf litter and top soil,
also under wood .

Acropyga inezae Forel, 1912
Material examined. Nee Soon Swamp Forest,
plot Q6, 3 Sep 2019, W. Wang leg., WWSG19-001, ZRC_ENT00013859.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Nee Soon Swamp Forest
Habitat/Ecology. Freshwater swamp forest – nest
found in hummock, soil matrix around treeroots.
Acropyga oceanica Emery, 1900
Material examined. None available.
Material not physically examined. Workers fromSingapore as indicated in LaPolla (2004), exact depository not specified.
Literature. LaPolla (2004).
Localities. Bukit Timah Nature Reserve
Habitat/Ecology. Primary and/or mature secondary forest.
Remarks. Habitat described as ‘degraded coastal
hill forest on granite’ transcribed from originallabels, however this is an inaccurate description of forest types in the BTNR.
Acropyga rubescens Forel, 1894
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[both as Acropyga acutiventris rubescens].
Localities. Bukit Timah Road
Habitat/Ecology. Garden.
Remarks. At the time of writing, this species appears to be known in Singapore based only on
literature records.
Anoplolepis gracilipes (Smith, 1857)
Material examined. Bukit Timah Nature Reserve,
ZRC_ENT00000343; same locality as previous, ZRC_ENT00000361; Central Catchment Nature Reserve, Venus Loop, 1.35549,
103.81578, ZRC_ENT00000729; Seletar Trail,
1.39097, 103.79952, ZRC_ENT00000730;
Pulau Ubin, ZRC_ENT0007859; same locality as previous, ZRC_HYM_0000378;
Mandai, ZRC_ENT00013534; Mandai,

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Northern Node, 1.40842, 103.78319, 54 ma.s.l., ZRC_ENT00013870; Mandai, Bird
Park Buffer, near L10, 1.40536, 103.78228,
73 m a.s.l., ZRC_ENT00013882; Kent
Ridge Park, ZRC_HYM_0000376; MacRitchie Reservoir, ZRC_HYM_0000377;
Pulau Sentosa, ZRC_HYM_0000911; Pulau Ubin, Outward Bound School grounds,
1.41662, 103.99443, ZRC_HYM_0001160;
Kranji Road, 1.42638, 103.75413, ZRC_
HYM_0001179; same locality as previous,
7 Sep 2016, ZRC_HYM_0001192, 1195;
same locality as previous, 5-7 Sep 2016, pitfall trap, ZRC_HYM_0001193-1194, 1196;
Mandai Road 3, M3D, 1.41119, 103.80513,
ZRC_HYM_0001180; Sunset Way, 1.32609,
103.77187, 22-24 Aug 2016, G.W. Yong leg.,
pitfall trap, ZRC_HYM_0001187; same locality as previous, 21-23 Aug 2016, G.W.
Yong leg., pitfall trap, ZRC_HYM_00011971198, 1200; same locality and collection
data as previous, trunk pitfall trap, ZRC_
HYM_0001199; Bukit Batok East Avenue 6,
1.34304, 103.76235, 27-29 Sep 2016, G.W.
Yong leg., pitfall trap, ZRC_HYM_0001201,
1203-1206; same locality as previous, 28-30
Aug 2016, G.W. Yong leg., pitfall trap, ZRC_
HYM_0001189; same locality as previous, 30
Aug – 1 Sep 2016, G.W. Yong leg., trunk pitfall trap, ZRC_HYM_0001190-1191; Seletar
Link, 7 Dec 2015, M.K.L. Wong leg., Winkler
extraction, ZRC_ENT00000730; same locality as previous, 16-18 Oct 2016, G.W. Yongleg., trunk pitfall trap, ZRC_HYM_0001217,
1220-1221, 1224; same locality as previous,
23-25 Oct 2016, G.W. Yong leg., pitfall trap,
ZRC_HYM_0001210 – 1211; same locality
as previous, 16 Oct 2016, G.W. Yong leg.,
ZRC_HYM_0001218, 1222, 1226; same locality as previous, 18 Oct 2016, G.W. Yongleg., ZRC_HYM_0001219, 1223, 1225, 1227;
same locality as previous, 23 Oct 2016, G.W.
Yong leg, ZRC_HYM_0001208, 1212, 12141215; same locality as previous, 25 Oct 2016,
G.W. Yong leg., ZRC_HYM_0001209, 1213,
1216; Upper Thomson Nature Park, 1.38311,
103.79835, 9-11 Oct 2016, G.W. Yong leg.,
pitfall trap, ZRC_HYM_0001265-1266; same
locality as previous, 2-4 Oct 2016, G.W. Yongleg., pitfall trap, ZRC_HYM_0001267 –

1269; same locality as previous, 4 Oct 2016,
G.W. Yong leg., ZRC_HYM_0001270.
Material not physically examined. Syntypes –CASENT0102951, ANTC5313 (OUMNH);
CASENT0103001, ANTC5357 (OUMNH);
CASENT0903237, ANTC23099 (BMNH)
Non-types - numerous specimens frommultiple localities, both catalogued anduncatalogued (ZRC); ANIC32-012878,
RWTacc.68.25 (ANIC); ANIC32-012879,
RWTacc.68.8 (ANIC).
Literature. Type – Smith (1857) [as Formica
gracilipes]. Viehmeyer (1916), Overbeck(1924) [both as Plagiolepis longipes]; Wetterer (2005); Tan & Corlett (2012); Yong etal. (2017).
Localities. Bukit Batok East Avenue; Bukit Timah
Nature Reserve; Central Catchment Nature
Reserve; Kranji Road; MacRitchie Reservoir;
Mandai; National University of Singapore(Kent Ridge/Clementi campus); Nee Soon;
Pulau Ubin; Seletar Link; Seletar Trail; Sentosa; Singapore Botanic Gardens; Sunset Way;
Upper Thomson Nature Park.
Habitat/Ecology. Broad range of different habitatsof varying levels of disturbance in Singapore,including primary and mature secondary forests, degraded wasteland forests and spontaneous vegetation fragments in urban or semi-urban settings. These ants have very generalized
foraging and nesting habits, typically found athigh densities and extremely pervasive where
they occur. Nests can be found under stones orin fallen rotting wood and logs.
Remarks. Singapore is the type locality of thisinfamous global invasive species. Anoplolepis gracilipes (Smith, 1857) was originally ajunior synonym of Anoplolepis longipes (Jerdon, 1851) (synonymy by Emery 1887); hence
Anoplolepis gracilipes (Smith, 1857) became
the first available replacement name for Anoplolepis longipes (Jerdon, 1851) (Bolton,
1995: 67), since the latter was identified as a
junior primary homonym of Pheidole longipes(Latreille, 1802). Colonies can be polydomous
and/or polygynous (forming ‘supercolonies’),with overwhelmingly numerous queens andworkers. Colony dispersal by ‘budding’ hasbeen documented for this highly invasive
species.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Camponotus (Karavaievia) overbecki Viehmeyer,
1916
Material examined. None available.
Material not physically examined. Types –CASENT0910416, ANTC32748 (MHNG);
FOCOL2490-2493 [as types of Camponotus(Myrmosphincta) dolichoderoides subsp.
overbecki] (ZMHB).
Literature. Type – Viehmeyer (1916); Overbecki(1924) [both as Camponotus (Myrmosphincta) dolichoderoides subsp. overbecki].
Localities. Mandai Road.
Habitat/Ecology. These ants were originallyfound in pasteboard nests under the leavesof a tree.
Remarks. Type locality in Singapore.
Camponotus (Myrmamblys) adustus Viehmeyer,
1916
Material examined. None available.
Material not physically examined. Type – FOCOL2281 (ZMHB).
Literature. Type – Viehmeyer (1916); Overbeck(1924) [both as Camponotus (Myrmamblys)
bellus subsp. adustus].
Localities. Bukit Timah Road
Habitat/Ecology. Garden – type specimens werecaught with a sweepnet over foliage.
Remarks. Type locality in Singapore.
Camponotus (Myrmamblys) bedoti Emery, 1893
Material examined. Prince George’s Park Residences (NUS), 1.292389, 103.778694, MarSep 2015, M.S. Foo & W. Wang leg., malaise
trap, ZRC_BDP (multiple); Upper Peirce Reservoir, 10 Jan 2014, Sk. Yamane leg., SG14SKY-49, ZRC_HYM_0000445-446; Pulau
Semakau, 2012-2014, J. Puniamoorthy et
al. leg., malaise trap, mangroves, ZRC_BDP(multiple); Sungei Buloh Wetland Reserve,
SB1, 21-28 Nov 2012, J. Puniamoorthy et al.
leg, malaise trap, Reg. 29595, mangroves,
ZRC_BDP0017497; Pulau Ubin, 4 Dec 2016,
G.W. Yong leg., ZRC_ENT00027980.
Material not physically examined. Unknown.

Literature. Forel (1907) [as Camponotus reticulatus bedoti]; Viehmeyer (1916); Overbeck(1924); Wang et al. (2018a).
Localities. Bukit Timah Road; Prince George’s
Park Residences (NUS); Pulau Semakau;
Pulau Ubin; Sungei Buloh Wetland Reserve;
Upper Peirce Reservoir.
Habitat/Ecology. Associated with both matureand young secondary habitats in Singapore,including mangrove back forest and disturbed
secondary forest fragments in urban or semiurban settings. Colonies have been found indead twigs on trees.
Camponotus (Myrmamblys) bellus Forel, 1908
Material examined. None available.
Material not physically examined. Minor workers, queens and males, FOCOL2289-2296(ZMHB).
Literature. First description of queen – Forel(1912). First descriptions of minor worker andmale – Viehmeyer (1916); Overbeck 1924.
Localities. Bukit Timah Road
Habitat/Ecology. Garden – colonies in witheredbamboo, and in hollow branches of trees.
Remarks. This species was first described basedon a soldier collected from Indonesia. The
queen, minor worker and male for the species
were described later on based on specimens
collected from Singapore.
Camponotus (Myrmamblys) ephippiatus Viehmeyer, 1916
Material examined. None available.
Material not physically examined. Type – FOCOL2280, FOCOL2280 (ZMHB).
Literature. Type – Viehmeyer (1916); Overbeck(1924).
Localities. Bukit Timah Road
Habitat/Ecology. Garden – type specimens werecaught with a sweepnet over foliage.
Remarks. Type locality in Singapore.

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Camponotus (Myrmamblys) gretae Forel, 1902
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[as Camponotus (Colobopsis) gretae].
Localities. Bukit Timah Road
Habitat/Ecology. Garden.
Remarks. The only literature record of this species
in Singapore is that of a single queen collectedfrom a lamp in a garden (Viehmeyer 1916;
Overbeck 1924).
Camponotus (Myrmamblys) reticulatus Roger,
1863
Material examined. Mandai mangrove, 2
Nov 1976, D.H. Murphy leg., ZRC_
HYM_0000945; alate queen and worker, same
locality as previous, 2 Oct 1970, D.H. Murphyleg., ZRC_HYM_0000633; Namly Avenue,
25 Apr 1985, D.H. Murphy leg., forest, ZRC_
HYM_0000946; Kent Ridge, 24 Jun 1985,
D.H. Murphy leg., ZRC_HYM_0000947-949.
Material not physically examined. Unknown.
Literature. None; new record.
Localities. Kent Ridge; Mandai mangrove; Namly
Avenue.
Habitat/Ecology. This species is usually associated with both mature and young secondary
habitats in Singapore, mostly disturbed forest
fragments in urban or semi-urban parklands,
and mangrove forests. The ants can be foundin damp logs, sometimes on the ground under Eugenia plants, in close association withplanthoppers (Flatidae). In mangroves, a nest
was found in a twig of Sonneratia.
Remarks. This species is superficially similar to
C. bedoti, especially in the unique convex lateral profile of the propodeal dorsum. Minor
workers of C. reticulatus can be differentiated from C. bedoti based on the following (C.
bedoti traits in parentheses): 1) largely dark
reddish-brown in colour, with front edge ofhead including malar area brownish-yellow(largely black, including the malar area), 2)
entire body generally densely reticulate-punctate, appearing rather dull or weakly shining(body largely finely reticulate-punctate andshiny, gaster sparsely punctate and shining,sometimes superficially reticulate), 3) bodycovered with dense appressed hairs or pubescence (body with much sparser pubescence, the gaster especially with dispersed
punctures bearing standing hairs), 4) dorsumof propodeum more strongly depressed, i.e.,
propodeal ‘saddle’ more distinct (propodeal
dorsum weakly depressed, propodeal saddle
indistinct).
Camponotus (Myrmamblys) reticulatus sericellus
Viehmeyer, 1916
Material examined. Queen, Prince George’s Park
Residences (NUS), 1.292389, 103.778694,
1-8 Apr 2015, M.S. Foo & W. Wang leg.,malaise trap, NUS0005, ZRC_BDP0046220;
Sime forest, collection date and collector unknown, ZRC_ENT00048388.
Material not physically examined. Types –CASENT0910532, ANTC32839 (MHNG);
FOCOL2283 – 2288 (ZMHB).
Literature. Type – Viehmeyer (1916); Overbeck(1924) [both as Camponotus (Myrmamblys)
bedoti subsp. sericellus]. Wang et al. (2018a).
Localities. Type locality – Bukit Timah Road.
Prince George’s Park Residences (NUS);
Sime forest.
Habitat/Ecology. Type specimens of this subspecies were collected from a garden, “between
bark and stem” on a Chiku (Manilkara zapota)
tree. More recent specimens were sampledfrom disturbed secondary forest fragments insemi-urban settings.
Remarks. Type locality in Singapore. Minor workers of C. reticulatus sericellus resemble thoseof C. reticulatus, though the latter appear tohave somewhat shorter antennal scapes. At
the time of writing, we have insufficient viable material (i.e., aged queen specimen of C.
reticulatus in poor condition, no major workers available for comparison) to confidently
assess distinguishing features between the twoforms. Thus, we maintain status quo for thesubspecies sericellus.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Camponotus (Myrmosaulus) camelinus (Smith,
1857)
Material examined. None available.
Material not physically examined. Syntype –CASENT0901905, ANTC21350 (OUMNH).
Non-type – CASENT0906945, ANTC27560(BMNH).
Literature. Type – Smith (1857) [as Formica
camelina].
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore context.
Remarks. Type locality in Singapore.
Camponotus (Myrmosericus) parius Emery,
1889
Material examined. Lee Kong Chian Natural History Museum (NUS), 29 Mar 2018, W. Wang leg.,
WW-SG18-Campo1, ZRC_ENT00000919;
MacRitchie Catchment Reservoir, 20 Apr
1994, collector unknown, NS 123B, ZRC_
HYM_0000109; MacRitchie Reservoir, 6 Jan
2014, Sk. Yamane leg., ZRC_HYM_0000404;
Upper Peirce Reservoir, 10 Jan 2014, Sk. Yamane leg. (SKYC); Pulau Ubin, 7 Jan 2014,
Sk. Yamane leg., ZRC_HYM_0000405; Choa
Chu Kang Avenue 2, Block 273, 1 Mar 2017,
D.J. Court leg., ZRC_HYM_0001723; University Hall (NUS), 1.297111, 103.776583,
12-19 Aug 2015, M.S. Foo & W. Wang leg.,malaise trap, NUS0084, ZRC_BDP0047532;
Prince George’s Park Residences (NUS),
1.292389, 103.778694, 8-15 Jul 2015, M.S.
Foo & W. Wang leg., malaise trap, NUS0061,
ZRC_BDP0045928; same locality as previous, 1-8 Sep 2015, M.S. Foo & W. Wang leg.,malaise trap, NUS0093, ZRC_BDP0047461;
Pulau Tekukor, 9 Dec 2016, G.W. Yongleg., ZRC_ENT00027978; St. John’s Island,
1.220167, 103.847472, 11 May 2016, J.S.
Ascher et al. leg., ZRC_ENT00047855; City
Hall, 29 Jul 2014, J.K. Wetterer leg., ZRC_
ENT00054652; Outram Park, same collector
and date as previous, ZRC_ENT00054653;
Serangoon, 30 Jul 2014, same collector as previous, ZRC_ENT00054654; Marine Parade, 1
Aug 2014, same collector as previous, ZRC_
ENT00054655; Kranji, 2 Aug 2014, same col-lector as previous, ZRC_ENT00054656; Singapore Botanic Gardens, 5 Aug 2014, samecollector as previous, ZRC_ENT00054659;
Pasir Panjang, same collector and date as previous, ZRC_ENT00054660.
Material not physically examined. Unknown.
Literature. Wang et al. (2018a) [misidentified as
Camponotus rufoglaucus].
Localities. Choa Chu Kang Avenue 2; City Hall;
Kranji; National University of Singapore(Kent Ridge/Clementi campus); MacRitchie
Reservoir; Marine Parade; Outram Park; Pasir
Panjang; Pulau Tekukor; Pulau Ubin; Serangoon; Singapore Botanic Gardens; St. John’sIsland; Upper Peirce Reservoir.
Habitat/Ecology. This species is often associatedwith young secondary habitat in Singapore,also commonly observed close to human infrastructure in urban or semi-urban settings.
Nests were typically seen in soil in cultivatedgrassland or grass patches near roadside trees.
Colony presence is often conspicuous withnarrow nest entrance usually surrounded bya relatively (much) wider outer ring of sandy
soil.
Remarks. Often mistaken for the other morewell-known black Camponotus species — C.
auriventris, C. parius can be distinguishedfrom the latter mainly by the absence of adeep metanotal groove (in C. auriventris themetanotal groove is visible as a deep indentation between mesonotum and propodeum inlateral view), also a smoother sculpture withfine whitish pubescence.
		Camponotus parius also closely resembles C. rufoglaucus in general habitus. Workers of the two species can be differentiated bycolour – C. parius is almost entirely uniformly
black, while C. rufoglaucus is typically blackwith varying amounts of reddish-brown. Body
colour, however, might be unreliable becauseof the broad variation apparent in the latter
species. The species may also be distinguishedbased on condition of standing pilosity. In
C. parius workers, standing hairs on entire
dorsum are sparse, laterally-projecting hairs
almost absent along sides of head in full-faceview except in anterior portion closer to mandibular base. In contrast, C. rufoglaucus workers have denser and more numerous standing

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hairs on entire dorsum, especially the mesosoma; in full-face view, sides of head linedwith many laterally-projecting hairs.
Camponotus (Myrmotarsus) nigricans enganensis Forel, 1916
Material examined. Alate queen, Lee Kong Chian
Natural History Museum (NUS), on staircase
linking to NUS Museum, 21 Sep 2020, J.S.
Tan leg., ZRC_ENT00028320.
Material not physically examined. FOCOL2338[as type of Camponotus (Myrmotarsus) nigricans subsp. nitidiceps] (ZMHB).
Literature. Viehmeyer (1916), Overbeck (1924)[both as Camponotus (Myrmotarsus) nigricans subsp. nitidiceps].
Localities. Bukit Timah Road. Single alate queen
found at LKCNHM building compound may
have dispersed from nest in unknown distantlocation, exact locality of original nest unverifiable.
Habitat/Ecology. Queens have been found attracted to lamps in a garden.
Remarks. This species was originally describedbased on the queen. Appearance of workers
currently unknown. May be more common
than might be inferred from relatively rareoccurrences of alate queens.
Camponotus (Myrmotarsus) quadrisectus hians
Forel, 1907
Material examined. None available.
Material not physically examined. Types –CASENT0910508-910509, ANTC328203282 (MHNG); CASENT0922412,
ANTC43615 (HNHM, Budapest).
Literature. Type – Forel (1907).
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore context.
Remarks. Type locality in Singapore.

Camponotus (Myrmotarsus) quadrisectus margaritae Forel, 1907
Material examined. Island Club forest, 20
May 1985, D.H. Murphy leg., ZRC_
HYM_0000890-892.
Material not physically examined. Syntypes –CASENT0910510, ANTC32822 (MHNG);
CASENT0922409, ANTC43612 (HNHM,
Budapest).
Literature. Type – Forel (1907).
Localities. Island Club forest (Central Catchment
Nature Reserve).
Habitat/Ecology. Workers of this species werecollected from primary or old/mature secondary forest, where they were found in a carton nest at the foot of a tree. The ants werereportedly found together with an unnamed
Crematogaster species.
Remarks. Type locality in Singapore.
Camponotus (Tanaemyrmex) albosparsus Bingham, 1903
Material examined. Prince George’s Park Residences (NUS), 1.292389, 103.778694, 8-15
Apr 2015, M.S. Foo & W. Wang leg., malaise trap, NUS0009, ZRC_BDP0046879;
same locality and collectors as previous, 5-12
Aug 2015, malaise trap, NUS0077, ZRC_
BDP0047381; Pulau Semakau, 2012-2014, J.
Puniamoorthy et al. leg., mangroves, malaise
trap, ZRC_BDP (multiple); Nee Soon Swamp
Forest, NS2, 28 Mar-4 Apr 2012, J. Puniamoorthy et al. leg., malaise trap Reg. 29158,
ZRC_BDP0015966; locality unknown, 23 Jul
1996, Sk. Yamane leg., ZRC_HYM_0000430431; Kent Ridge Park, 11 Jan 2014, Sk. Yamane leg. (SKYC).
Material not physically examined. Non-types,
J.K. Wetterer leg. (JKWC) – Amber Road,
Katong, 28 Jul 2014, vial #339; Pasir Panjang,
by MRT station, 5 Aug 2014, vial #484; Punggol, by MRT station, 30 Jul 2014, vial #390;
Serangoon, 30 Jul 2014, vial #389; Singapore
Botanic Gardens, 5 Aug 2014, vial #480.
Literature. Li (2006) [as Camponotus bedoti albosparsus]; Wang et al. (2018a).

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Localities. Amber Road (Katong); Kent Ridge
Park; Nee Soon Swamp Forest; Pasir Panjang;Prince George’s Park Residences (NUS); Pulau Semakau; Punggol; Serangoon; Singapore
Botanic Gardens.
Habitat/Ecology. Found in both mature and young
secondary habitats, including disturbed secondary forest fragments in urban or semiurban settings, mangrove and swamp forests
in Singapore. Individuals were usually foundin open areas around or near forest fringes oredges, sometimes under stones, and at bases oftrees, close to man-made infrastructure.
Remarks. Workers of this species are typicallydistinguished from other congeners by a pairof large pale-whitish patches/dots, on eachof the first two gastral tergites respectively.
Camponotus (Tanaemyrmex) arrogans (Smith,1858)
Material examined. Bukit Timah Nature Reserve,
BT03, 1.34913, 103.77914, 17 May 2017,
W. Wang leg., old secondary forest, WWSG17-053, ZRC_ENT00027972.
Material not physically examined. Syntype –CASENT0903598, ANTC23456 (BMNH).
Literature. Type – Smith (1858) [as Formica arrogans]. Emery (1896) [as Camponotus maculatus arrogans]; Viehmeyer (1916); Overbeck(1924).
Localities. Bukit Timah Nature Reserve
Habitat/Ecology. This species is associated withprimary and old or mature secondary habitatin Singapore. In old secondary forest, nestmates have been collected from the surface
of a buttress root of a large tree, presumably
in the vicinity of the actual nest. It is unclear
exactly where actual nests may be located,i.e., in tree cavities or amidst buttress litter.
Remarks. Type locality in Singapore.
Camponotus (Tanaemyrmex) carinifer Viehmeyer, 1916 stat. n.
Camponotus (Myrmoturba) tinctus var. carinifer
Viehmeyer, 1916: 156 (s.q.)
Subspecies of Camponotus irritans: Bolton, 1995:
91.
Material examined. Prince George’s Park Residences (NUS), 1.292389, 103.778694, 15-22

Apr 2015, M.S. Foo & W. Wang leg., malaise trap, NUS0013, ZRC_BDP0045404;
same locality and collectors as previous, 29
Jul-5 Aug 2015, malaise trap, NUS0073,
ZRC_BDP0047331; same locality and collectors as previous, 23-30 Sep 2015, malaise trap, NUS0105, ZRC_BDP0047294;
Mandai mangroves, 17 Nov 1978, D.H.
Murphy leg., ZRC_HYM_0000881; same
locality and collector as previous, 1975,
ZRC_HYM_0000882; Sungei Buloh Wetland
Reserve, 14 Mar 2018, M.S. Foo leg., backmangrove, ZRC_ENT00000911.
Material not physically examined. Types –CASENT0910154 [cotype major of C. (Myrmoturba) tinctus var. carinifer], ANTC32547(MHNG); FOCOL2391-2396 [queens andworkers of C. tinctus var. carinifer] (ZMHB).
Literature. Types - Viehmeyer (1916); Overbeck(1924) [both as Camponotus (Myrmoturba)
tinctus var. carinifer]. Wang et al. (2018a)[identified as Camponotus irritans].
Localities. Jurong Road; Mandai mangroves;
Prince George’s Park Residences (NUS);
Sungei Buloh Wetland Reserve.
Habitat/Ecology. This species was found mainly
in disturbed young or mature secondary forest fragments in semi-urban settings, close to
man-made infrastructure. The type series wascollected from a nest in a rotting dead tree
trunk along a road. In mangroves, a colony
was found nesting in a rotting branch of themangrove plant Rhizophora.
Remarks. Prior to this study, C. carinifer wastreated as a subspecies of Camponotus irritans(Smith, 1857). However, upon comparisonsof type images and original descriptions, we
believe C. carinifer is sufficiently distinct tobe considered a separate species.
		The original description of C. irritans
was based on “normal” minor workers (notmajors/soldiers), specifically a large worker(12.7 mm) and a smaller one (6.4 mm). We
noted that the maximum size for non-major
workers in C. irritans appears larger than inlocal minors of C. carinifer – the latter neverexceeds 10 mm in total length. Further, evenmajors of C. carinifer are at most around 10- 11mm in length, at least for specimens fromSingapore.

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Wendy Y. Wang, Eunice J.Y. Soh, Gordon W.J. Yong, Mark K.L. Wong, Benoit Guénard, Evan P. Economo & Seiki Yamane		Based on our observations, large minor
workers of C. carinifer can be distinguishedfrom those of C. irritans by the following (C.
irritans traits in parentheses): 1) mesosoma
not elongate, in dorsal view pronotum as wide
as head, rounded and not much narrowed anteriorly (mesosoma elongate and slender, pronotum narrower than head, distinctly tapered andnarrowed anteriorly), 2) posterior margin ofhead very weakly and broadly convex in fullface view (posterior margin of head strongly
convex), 3) petiole in profile narrower andmore squamiform, weakly tapered towards
apex (petiole longer, strongly tapered towardsapex), 4) apex of scape and most of antenna
flagellum light yellowish-brown, reddishbrown at the tips (apex of scape and entire
flagellum ferruginous).
		Camponotus carinifer is very similar toCamponotus tinctus (Smith, 1858) (nom. rev.,
see relevant species section below). In fact,
the minor workers are almost indistinguishable from each other. The two species can still
be differentiated based on major workers andqueens. We discuss this in greater detail under
Remarks for C. tinctus.
Camponotus (Tanaemyrmex) festinus eximius
Emery, 1900
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah Road
Habitat/Ecology. Garden.
Remarks. At the time of writing, this species isonly known from Singapore based on a single
literature record of a queen caught at a litlamp in a garden.
Camponotus (Tanaemyrmex) festinus simaluranus Forel, 1915
Material examined. Major worker, Nee Soon,
14 Apr 1993, Sk. Yamane leg., swamp forest(SKYC).
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[both as Camponotus (Dinomyrmex) inezae
var. simalurana].

Localities. Bukit Timah Road; Nee Soon.
Habitat/Ecology. In the only literature record ofthis subspecies in Singapore, the ants werereportedly found in the trunk of a garden
tree with cavities created by termites. Morerecently, a major worker was collected fromswamp forest.
Camponotus (Tanaemyrmex) irritans (Smith,
1857)
Material examined. Lee Kong Chian Natural History Museum bus stop, 13 Jan 2017,
S.H. Tan leg., nest in junction box, WWSG17-007, ZRC_HYM_0000575; KranjiRoad, Sep-Nov 2016, G.W. Yong et al. leg.,
ZRC_ENT00057823-57827; Upper Thomson
Nature Park, 11-13 Sep 2016, G.W. Yong etal. leg., ZRC_ENT00057828; Seletar Link,
23-25 Oct 2016, same collectors as previous,
ZRC_ENT00057829; Mandai Road, 29 Nov– 1 Dec 2016, G.W. Yong et al. leg., ZRC_
ENT00057830; Sunset Way, 31 Aug 2016,
G.W. Yong et al. leg., ZRC_ENT00057831.
Material not physically examined. Unknown.
Literature. Wang et al. (2018a) - but see Remarks
for Camponotus carinifer.
Localities. Kranji Road; Lee Kong Chian Natural
History Museum (NUS); Mandai Road; Seletar Link; Sunset Way.
Habitat/Ecology. These ants appear to be quite
tolerant of degraded or disturbed environments, able to thrive in areas near man-madeinfrastructure as well as young or mature secondary forest fragments. A colony was evenfound nesting inside an outdoor electricaljunction box next to a building in urban settings. The ants are commonly observed (unverified sightings), sometimes photographed,but not collected. The species is probably
more widespread locally than is represented
in formal museum holdings, not limited to thetwo localities listed in this section. In recent
times, these ants have apparently also become
increasingly popular in the local pet trade.
Remarks. As mentioned before (see Remarks for
C. carinifer), C. irritans was first describedbased on two minor workers – a large one(>12 mm) and another roughly half its size
(which Smith defined as truly ‘minor’). The

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

major worker was never formally described.
This is problematic with regards to speciesdelimitation, especially for species of the subgenus Tanaemyrmex, as the worker caste canbe extremely polymorphic.
		Within each colony sample we examined,
we observed a broad spectrum of intermediate worker forms, varying mainly in head
shape, size, and body colours. Nevertheless,the elongate and slender habitus, characteristic of C. irritans as it was first described,
persists across all variants, including teneralindividuals. It is difficult to clearly define theboundaries separating ‘majors’ from ‘minors’ because of the apparent continuum offorms. As expected, Smith’s (1857) originaldescription and the online type image of C.
irritans matched only with a few individuals
per sample. Major workers, also variable inhead shape, resemble those of Camponotus
carin Emery, 1889 and other close relatives.
Minor workers of different species of Camponotus can be almost identical to eachother. Some minor workers of the C. irritanscolonies we examined, for example, closely
matched the original description and typeimage for Camponotus mitis (Smith, 1858),
except for petiole shape. Consequently, we
cannot overlook the possibility that specimens
we currently recognize as “C. irritans” may infact belong to another similar species of thesubgenus Tanaemyrmex.
Camponotus (Tanaemyrmex) maculatus obfuscatus Viehmeyer, 1916
Material examined. None available.
Material not physically examined. Types – FOCOL2400-2402 (ZMHB) [as types of Camponotus (Myrmoturba) subnuda var. obfuscata]
Literature. Type – Viehmeyer (1916); Overbeck(1924) [both as Camponotus (Myrmoturba)
subnudus var. obfuscata].
Localities. Bukit Timah Road
Habitat/Ecology. Garden – type colony foundin withered bamboo. A solitary type queen
was also found in a rotten branch lying on
the ground.
Remarks. Type locality in Singapore.

Camponotus (Tanaemyrmex) maculatus subnudus Emery, 1889
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[both as Camponotus (Myrmoturba) subnudus].
Localities. Bukit Timah Road
Habitat/Ecology. Garden – a single queen collected at a lit-lamp.
Remarks. At the time of writing, there are only literature records of this subspecies in Singapore.
Camponotus (Tanaemyrmex) maxwellensis
Forel, 1913
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[both as Camponotus (Myrmoturba) maculatus subsp. maxwellensis].
Localities. Bukit Timah Road
Habitat/Ecology. Garden – a single queen collected in a hollow branch of a tree.
Remarks. At the time of writing, there are only
literature records of this species in Singapore.
Camponotus (Tanaemyrmex) mitis (Smith, 1858)
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[both as Camponotus (Myrmoturba) maculatus subsp. mitis].
Localities. Bukit Timah Road
Habitat/Ecology. Garden.
Remarks. At the time of writing, there are only
literature records of this species in Singapore.
Camponotus (Tanaemyrmex) tinctus (Smith,1858), nom. rev.
Formica tincta Smith, 1858: 21 (q.)
Viehmeyer, 1916; 155 (s.w.); Karavaiev, 1929b:
237 (m.)Combination in Camponotus: Mayr, 1862: 676.
Combination in Camponotus (Myrmoturba): Forel,
1914: 267.
Combination in Camponotus (Tanaemyrmex): Emery, 1925: 94.

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Status as species: Mayr, 1862: 676; Mayr, 1863:
401; Roger, 1863b: 3; Mayr, 1865: 30; Mayr,
1867: 39 (redescription); Smith, 1871: 304;
Dalla Torre, 1893: 254; Emery, 1896b: 371;
Viehmeyer, 1916: 154; Wheeler, 1935: 40.
Junior synonym of Camponotus luteus: Bingham,
1903: 379.
Subspecies of Camponotus maculatus: Emery,
1900a: 704; Stitz, 1925: 124.
Subspecies of Camponotus irritans: Emery,
1920: 7, 17; Stitz, 1925: 124; Emery, 1925:
94; Karavaiev, 1929b: 237; Stitz, 1932: 736;
Chapman & Capco, 1951: 248; Baltazar,
1966: 274; Bolton, 1995: 127.
Material examined. None available.
Material not physically examined. FOCOL2386-2387 [used in first descriptions ofCamponotus tinctus workers] (ZMHB).
Literature. Types - Viehmeyer (1916); Overbeck(1924) [both as Camponotus (Myrmoturba)
tinctus var. carinifer, and C. (Myrmoturba)
tinctus].
Localities. Bukit Timah Road
Habitat/Ecology. A queen of this species wascaught at a lit lamp in a garden. The ants
were also found nesting in decaying or rotten
branches.
Remarks. While Camponotus tinctus was firstdescribed based on a queen specimen from
Myanmar, major and minor workers of thespecies were described later by Viehmeyer(1916) based on specimens from Singapore.
Minor workers of this species can be distinguished from C. irritans using the characters
detailed under Remarks for C. carinifer (seeabove).
		Camponotus tinctus closely resembles C.
carinifer, though major workers and queens
may be differentiated by the following characters (C. carinifer traits in parentheses): 1)
clypeus without or with very weak and indistinct median carina, (clypeus with distinct median carina), 2) posterior head margin almost
flat, weakly and broadly concave (posterior
head margin deeply concave medially), 3)
in anterodorsal/oblique view, rear corners ofhead not forming rounded lobes, only slightly
projected posteriorly (rear corners of head
forming rounded lobes, strongly projected
posteriorly).

		Given the absence of more compellingobservable differences, it remains possiblethat C. carinifer and C. tinctus are actually
ecomorphs of a morphologically variable
species. In this case, DNA evidence coupled
to more material coverage of the two species
from other parts of Indomalaya, may be key
to resolving this issue.
Camponotus (Tanaemyrmex) variegatus (Smith,1858)
Material examined. Linden Drive, 6 Jun
1985, D.H. Murphy leg., on mango, ZRC_
HYM_0000884.
Material not physically examined. Unknown.
Literature. Type — Smith (1858) [as Formica
variegata].
Localities. Linden Drive
Habitat/Ecology. Unknown in Singapore context– habitat information written on original labelof ZRC specimen illegible.
Remarks. Both Ceylon (now known as Sri Lanka)
and Singapore were named as type localities
in the original description of this species(Smith 1858), no lectotype was designated.
Camponotus (Tanaemyrmex) variegatus fuscithorax Dalla Torre, 1893
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[both as Camponotus (Myrmoturba) maculatus subsp. mitis var. fuscithorax].
Localities. Bukit Timah Road
Habitat/Ecology. These ants were found in gardens – colonies were found in the end of abranch of a dead tree, in bamboo, and under an
old packing-case on the ground. Queens havealso been collected from lit-lamps in gardens.
Remarks. The soldier, queen and male of thissubspecies were first described by Viehmeyer (1916) based on specimens collected in
Singapore.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Colobopsis badia (Smith, 1857)
Material examined. None available.
Material not physically examined. Syntype –CASENT0901897, ANTC21342 (OUMNH)
Literature. Type – Smith (1857) [as Formica
badia]. Viehmeyer (1916), Overbeck (1924)[both as Camponotus (Colobopsis) badius].
Localities. Ayer Terjun (archaic place name);
Bukit Timah Hill; Mandai Road.
Habitat/Ecology. Typically associated with primary and old secondary forest, these ants
have been found in Nepenthes pitcher cups;
colonies have been collected from inside thewood of living trees.
Remarks. Type locality in Singapore.
Colobopsis leonardi (Emery, 1889)
Material examined. Sungei Buloh, 8 Jan 2013,
Sk. Yamane leg., lower vegetation, ZRC_
HYM_0000441; MacRitchie Reservoir, 6 Jan
2014, Sk. Yamane leg., ZRC_HYM_0000442;
Kent Ridge, 13 May 1985, D.H. Murphy leg.,roadside tree trunk, ZRC_HYM_0000886888; same locality as previous, 28 May 1985,
D.H. Murphy leg., ZRC_HYM_0000889;
Bukit Timah Nature Reserve, near BT02,
1.35599, 103.77397, 25 Jan 2017, W. Wangleg., ZRC_ENT00014152; Kranji mangrove,
2 Jul 1985, D.H. Murphy leg., dead twig
and tree bole, DHM-SG85-Colo1, ZRC_
ENT00027954.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[as Camponotus (Colobopsis) pilosus].
Localities. Bukit Timah Nature Reserve (= Bukit
Timah Hill); Bukit Timah Road; Kent Ridge;
Kranji; MacRitchie Reservoir; Sungei Buloh.
Habitat/Ecology. This species appears associatedmostly with primary and old or mature secondary habitats, including mangrove forest.
The ants were often found on trees or lower
vegetation; nests have been collected fromdead twigs and tree boles.

Colobopsis leonardi gracilenta (Viehmeyer,
1916)
Material examined. None available.
Material not physically examined. Type queen– FOCOL2515 (ZMHB) [as Camponotus(Colobopsis) pilosus subsp. gracilentus].
Literature. Type – Viehmeyer (1916); Overbeck(1924) [both as Camponotus (Colobopsis)
pilosus subsp. gracilentus].
Localities. Bukit Timah Road
Habitat/Ecology. Garden.
Remarks. This subspecies was first described andis only known based on a single queen collected from a lit-lamp in a garden; no workers
have yet to be formally described. Type locality in Singapore.
Colobopsis moeschi lygaea (Viehmeyer, 1916)
Material examined. None available.
Material not physically examined. Types – FOCOL2270-2271 (ZMHB) [as Camponotus(Myrmamblys) moeschi var. lygaea].
Literature. Type – Viehmeyer (1916); Overbeck(1924) [both as Camponotus (Myrmamblys)
moeschi var. lygaea].
Localities. Ayer Terjun (archaic place name)
Habitat/Ecology. No information available on
macrohabitat; ants of the type series were
found in Nepenthes pitcher cups. Nepenthes
pitcher plants are usually found in old or mature secondary forest.
Remarks. Type locality in Singapore.
Colobopsis rothneyi (Forel, 1893)
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[as Camponotus (Colobopsis) rothneyi].
Localities. Bukit Timah Hill
Habitat/Ecology. Deducing from its reported locality in Singapore, this species is probably
associated mostly with primary and/or maturesecondary forest. Nests have been found inhollow branches.
Remarks. At the time of writing, there are only
literature records of this species in Singapore.

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Colobopsis rothneyi krafti (Forel, 1901)
Material examined. None available.
Material not physically examined. Unknown.
Literature. Type queen – Forel (1901) [as Camponotus (Colobopsis) rothneyi r. krafti].
Localities. Unknown.
Habitat/Ecology. None available.
Remarks. Type locality in Singapore. This subspecies was first described and is only known
based on a single queen collected from Singapore by Dr. H. Kraft in late May 1894 (Forel
1901). Whereabouts of the physical typespecimen are unknown at the time of writing.
Colobopsis vitrea (Smith, 1860)
Material examined. Admiralty, 4 Feb 2015,
W. Wang leg., abandoned park, ZRC_
ENT00000974; Bukit Timah forest, 3
Nov 1973, D.H. Murphy leg., ZRC_
HYM_00000968; Tyersall Istana, 9 Jun 1985,
D.H. Murphy leg., ZRC_HYM_0000969;
Catchment forest (= Central Catchment Nature Reserve), 15 Sep 1964, D.H. Murphy leg.,trunk, ZRC_HYM_0000970; Mandai forest,
15 Jan 2020, W. Wang leg., horizontal trunk oftree, ZRC_ENT00027940; Pulau Ubin, PU1,
1°25’11.64’’N, 103°56’6.252’’E, 25 Aug-1
Sep 2012, J. Puniamoorthy et al. leg., malaise
trap, Reg. 29436, ZRC_BDP0015995; Sungei
Buloh Wetland Reserve, SB1, 1°26’46.3’’N,
103°43’49.9’’E, 16 m, 2012-2014, J. Puniamoorthy et al. leg., malaise trap, ZRC_BDP(multiple).
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[both as Camponotus (Colobopsis) vitreus].
Localities. Admiralty; Bukit Timah forest; Bukit
Timah Road; Central Catchment Nature Reserve; Mandai forest; Pulau Ubin; Sungei Buloh Wetland Reserve; Tyersall Istana.
Habitat/Ecology. This species is associated withboth mature and young secondary habitat in
Singapore, often disturbed secondary forest
fragments in urban or semi-urban settings,
also abandoned park forests. Alates have beencollected using malaise traps from mangroveback forests, but actual nesting sites in mangroves have yet to be found. Colonies havebeen found in withered bamboo, decayedwood, and hollow twigs of mangosteen
trees. Individuals were also found foraging
on trunks of living trees.
Colobopsis vitrea angustula (Emery, 1925)
Material examined. None available.
Material not physically examined.
CASENT0915605, ANTC34823 (NHMW)
Literature. Viehmeyer (1916), Overbeck (1924)[as Camponotus (Colobopsis) vitreus var. angustata].
Localities. Ayer Terjun (archaic place name);
Bukit Timah Road; Changi.
Habitat/Ecology. Colonies have been found in rotten wood and bamboo in gardens; nests seemto occupy deserted termite tunnels or cavities.
One colony was reported to be nesting in thestem of a fern, hollowed out to around fivefeet above ground, with small round entrance
holes scattered some distance apart (from eachother).
Remarks. The very first recorded specimen of
this species was an alate queen collected by
Ranson in 1870, from Changi.
Colobopsis vitrea vittatula (Forel, 1913)
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah Road
Habitat/Ecology. Garden.
Remarks. This subspecies was first described andis only known based on a single queen from
Sumatra. Likewise, the only known record of
the subspecies in Singapore is that of a single
queen collected from a lit-lamp in a garden.
Dinomyrmex gigas (Latreille, 1802)
Material examined. Alate queen, Bukit Batok St. 21, 28 May 2018, Faith leg., ZRC_
ENT00000990; Bukit Timah Nature Reserve,
North View Path, 15 Aug 2009, T.M. Leong etal. leg., ZRC_6_21680; Bukit Timah forest, 16
Jul 1965, P.H.L. leg., ZRC_HYM_0000939;
dealate queen, Holland Close, 11 Dec 2019, K.
Lim leg., ZRC_ENT00014092; Kent Ridge, 20

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Dec 1989, K. Yong leg., ZRC_HYM_0000156;
alate queen, Lee Kong Chian Natural History
Museum (NUS), ground floor, 21 Jan 2019, S.H.
Tan leg., ZRC_ENT00047825; alate queen,same locality as previous, main lobby, 12 Mar
2018, T. Mokan leg., ZRC_ENT00000900;
Lower Peirce Reservoir Road, Aug 1990, collector unknown, ZRC_ENT00000218; Lower
Peirce Reservoir, Forest B, 7 Aug 1990, collector unknown, ZRC_ENT00000096; Lower
Peirce Reservoir, Forest A, 8 Aug 1990, collector unknown, ZRC_ENT00000118; same
locality as previous, 25 Aug 1990, collector
unknown, ZRC_ENT00000185; same locality
as previous, 18 Aug 1990, collector unknown,
ZRC_HM_0000146; MacRitchie Catchment
Reservoir (Reserve), 25 Apr 1994, J.C. leg.,
NS 125B, ZRC_HYM_0000070; same locality, 9 Jan 2014, Sk. Yamane leg. (SKYC);
Nee Soon Swamp Forest, 7 Jun 1990, H.K.
Lua leg., 106, ZRC_HYM_0000134; locality unknown, 25 Dec 1967, G.W. Classy leg.,
ZRC_HYM_0000940-941, 943-944; locality
unknown, 2 Jan 1968, same collector as previous, ZRC_HYM_0000942; workers and alates,
Nee Soon Swamp Forest, 2012-2014, J. Puniamoorthy et al. leg., malaise trap, ZRC_BDP(multiple).
Material not physically examined.
CASENT0280278, PSW09574 (PSWC);
FMNHINS0000045763-45764, FMNH.Emu.
CE.IRN_163906 (FMNH, Chicago, USA).
Literature. Smith (1857) [as Formica gigas];
Viehmeyer (1916), Overbeck (1924) [as Camponotus (Dinomyrmex) gigas]; Tan & Corlett(2012) [as Camponotus gigas].
Localities. Bukit Timah Nature Reserve (= Bukit
Timah Hill); Kent Ridge; Lower Peirce Reservoir; MacRitchie Catchment Reserve; NeeSoon Swamp Forest.
Habitat/Ecology. This species is associated mostly with primary and old or mature secondary
habitat in Singapore, including swamp forests.
These large ants have been observed nestingin huge fallen logs or rotting wood close tobases of mature trees. Solitary workers wereusually found foraging over wide areas of forest, sometimes even in grassland adjacent toforest edges.

Remarks. Alates, typically queens, have occasionally been found in urban dwellings orresidential areas somewhat far away fromtheir original nest sites (i.e., forested areas).
These locations cannot be strictly consideredas ‘localities’ for the species, since they do
not correspond to where actual nests occur.
Queens are known to fly over long distancesaway from the source nest during dispersal.
Echinopla lineata Mayr, 1862
Material examined. Kent Ridge, 24 Jun 1985,
D.H. Murphy leg., ZRC_HYM_0000249;
Kent Ridge Park, 11 Jan 2014, Sk. Yamaneleg., ZRC_HYM_0000403; Kranji mangrove,
2 Jul 1985, D.H. Murphy leg., DHM-SG85Poly1, ZRC_ENT00027955; Linden Drive
kampong, 27 Mar 1975, D.H. Murphy leg.,
ZRC_HYM_0000918-919; Mandai mangroves, 18 Aug 1977, D.H. Murphy leg.,
ZRC_HYM_0000917; Pulau Ubin, 1 Aug
2016, W. Wang leg., ZRC_ENT00014151;
Seletar Link, 1.40518, 103.88555, 23 Oct
2016, G.W. Yong leg., SL7GH1076, ZRC_
HYM_0001181; Sungei Buloh, 8 Jan 2013,
Sk. Yamane leg., ZRC_HYM_0000402.
Material not physically examined.
CASENT0280335, PSW10245-4 (PSWC).
In Zettel & Laciny (2015): Singapore Zoo, 2
Nov 2003, H. Zettel leg. (ZCW).
Literature. Zettel & Laciny (2015).
Localities. Kent Ridge; Kent Ridge Park; Kranjimangrove; Lim Chu Kang; Linden Drive
kampong; Mandai mangroves; Pulau Ubin;
Seletar Link; Singapore Zoo (in Mandai);Sungei Buloh.
Habitat/Ecology. This arboreal species was foundmainly in mangroves and young secondary
forests of varying quality and levels of disturbance, such as waste woodland secondary forest fragments in urban or semi-urban
settings. In mangroves, the ants have beenfound on lower vegetation, or nesting in felled
Lumnitzera or hollow twigs of Lumnitzera. In
other forests, individuals have been found inhollow branches suspended in bushes under
Eugenia, also nesting in other suspended hollow branches.

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Echinopla melanarctos Smith, 1857

Gesomyrmex spatulatus Cole, 1949

Material examined. None available.
Material not physically examined. Holotype –CASENT0901878, ANTC21323 (OUMNH).
Literature. Type – Smith (1857).
Localities. Unknown.
Habitat/Ecology. No information available in the
Singapore context.
Remarks. Type locality in Singapore. At the timeof writing, this species is only known by its
primary type collected from Singapore.

Material examined. Nassim Road, 25 Sep 1969,
D.H. Murphy leg., DHM-SG69-Geso1/2,
ZRC_HYM_0000001-3.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Nassim Road
Habitat/Ecology. This species was found nestingin old cerambycid mines or borings in live
terminal twigs of a parasitic fig (Ficus sp.) ona large fallen tree. Some individuals were also
observed to be swiftly running under sunlighton the same fallen tree.
Remarks. Specimens examined confer more with
G. spatulatus sensu lato in terms of petiolar and clypeal shapes, though the species
G. kalshoveni Wheeler, 1929 appears more
likely in terms of known geographic distribution (Indonesia (Java), as opposed to India(Assam)). The Singapore specimens may infact be an entirely new species with charactersintermediate between G. kalshoveni and G.
spatulatus. Given the lack of more compelling
empirical evidence at the time of writing, we
tentatively treat these specimens as conspecific to G. spatulatus.

Echinopla rugosa André, 1892
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah Road
Habitat/Ecology. A colony was found in witheredbamboo in a garden.
Remarks. At the time of writing, this species isonly known from Singapore based on literature records.
Euprenolepis procera (Emery, 1900)
Material examined. Nee Soon Swamp Forest, NS1, 16-23 May 2012, J. Puniamoorthy et al. leg., malaise trap, Reg. 29246,
ZRC_BDP0015065; Mandai Road, 5-7 Dec
2016, G.W. Yong leg., pitfall trap, ZRC_
HYM_0000513; Mandai Road 1, M1D,
5-7 Dec 2016, G.W. Yong leg., pitfall trap,
M1D19GPd2571, ZRC_HYM_0001182;
Upper Thomson Nature Park, UT1, 4 Oct
2016, G.W. Yong leg., UT11GH1898, ZRC_
HYM_0001183.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Nee Soon Swamp Forest; Mandai
Road; Upper Thomson Nature Park.
Habitat/Ecology. This species is associated mostly with primary or mature secondary habitat, including swamp and native-dominatedsecondary forest, as well as young secondary
habitat such as abandoned plantation forests
in Singapore.
Remarks. These ants are usually ground-foraging
and were mostly collected using pitfall traps.

Lepisiota rothneyi splendida (Viehmeyer, 1916)
Material examined. None available.
Material not physically examined. Types – FOCOL2227-2229 (ZMHB) [as Plagiolepis rothneyi subsp. splendida]
Literature. Type – Viehmeyer (1916) [as Acantholepis rothneyi subsp. splendida]; Overbeck(1924) [as Plagiolepis rothneyi splendida].
Localities. Ayer Terjun (archaic place name);
Bukit Timah Road.
Habitat/Ecology. Type specimens were collectedfrom Nepenthes pitcher cups in a garden.
Remarks. Type locality in Singapore.
Myrmoteras (Myagroteras) bakeri Wheeler, 1919
Material examined. Bukit Timah forest, 1965,
D.H. Murphy leg., ZRC_HYM_0000866.
Material not physically examined. ANIC32014953, Shattuck1345232391 (ANIC).
Literature. None. New record.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Localities. Bukit Timah Nature Reserve (= Bukit
Timah forest).
Habitat/Ecology. This species was found in leaf
litter in primary and/or old secondary forests
in Singapore.
Myrmoteras barbouri Creighton, 1930
Material examined. None available.
Material not physically examined. ANIC32014921, Shattuck1345232397 (ANIC).
Literature. Moffett (1985, 1986).
Localities. Bukit Timah Nature Reserve
Habitat/Ecology. This species is associated withprimary and old or mature secondary forests in
Singapore. Individuals can be found in leaf litter on forest floors. These ants have also beenfound in disturbed primary forest, at around
100 m elevation - a nest was found in a soil
crevice beneath leaf litter.
Remarks. Voucher specimens of this species areunavailable in ZRC, or have not been identified from unsorted material. Original labels ofmaterial in ANIC describe habitat as ‘degraded coastal hill forest on granite’, but this may
not be an accurate description of the forest in
Bukit Timah Nature Reserve where specimens
were collected.
Nylanderia birmana (Forel, 1902)
Material examined. Lower Peirce Reservoir
Park, 1.39095, 103.81041, 17 Dec 2021, W.N.
Lam leg., ZRC_ENT00057789-57790; Upper
Peirce Reservoir Park, 1.38726, 103.81030,
same collector and date as previous, ZRC_
ENT00057788; workers and alate queen,same locality and collector as previous, 27
Dec 2021, ZRC_ENT00057791; MacRitchie
Reservoir Park, 1.35229, 103.82450, 21 Oct
2021, same collector as previous, ZRC_
ENT00057792.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Lower Peirce Reservoir; MacRitchie
Reservoir; Upper Peirce Reservoir.
Habitat/Ecology. This species was collected fromleaf litter in freshwater swamp forest, also
native-dominated primary and/or secondary
dryland forests.

Remarks. Nylanderia birmana resembles Paratrechina longicornis in body colour and general habitus, but can be distinguished from thelatter by the following (P. longicornis traits inparentheses): 1) head and mesosoma finely
and irregularly reticulate, weakly shining or
almost matte (smooth and moderately shining,
weakly shagreened), 2) antennal scapes witherect macrosetae (scapes lacking erect macrosetae), 3) mesosoma and legs relatively morerobust and larger, dorsal outline of promesonotum more strongly convex in lateral view
(body more slender, dorsal outline of promesonotum weakly convex in lateral view).
Nylanderia bourbonica (Forel, 1886)
Material examined. Alate queens, Changi,
1.36206, 104.02525, 7 May 2020, mercury
vapour lamp, ZRC; alate queen, Pulau Semakau New Fragment, SMN2, 1°12’04.5’’N,
103°45’46.1’’E, 21-28 Nov 2013, J. Puniamoorthy et al. leg., replanted mangroves, malaise trap, Reg. 30285, ZRC_BDP0015368;
alate queen, same locality and collector as previous, 13-20 Aug 2012, replanted mangroves,malaise trap, Reg. 29431, ZRC_BDP0015362;
male, Pulau Ubin, PU2, 1°24’39.1”N,
103°59’23.2”E, 22-29 Sep 2012, J. Puniamoorthy et al. leg., mangroves, malaise trap,
Reg. 29489, ZRC_BDP00008654.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Changi; Pulau Semakau; Pulau Ubin.
Habitat/Ecology. A recognized tramp species
that has spread nearly worldwide to tropicalor subtropical areas mainly via human commerce (Wetterer 1998), N. bourbonica identified from Singapore appears to occur in a
fairly broad range of habitats at varied levelsof human disturbance. These habitats include
disturbed secondary forest fragments in semiurban settings, also replanted mangroves andold growth mangrove forests.
Remarks. At the time of writing, the examinedmaterial (comprising only winged alates)
could not be confidently established as N.
bourbonica with absolute certainty. The species appears to be morphologically variable
across populations, and therein lies the pos-

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sibility that some of these populations may actually be different, albeit closely-related, species. In view of this taxonomic conundrum,
though there are slight differences between
the morphology of the Singapore specimens
and that of the type images, we tentatively
consider the Singapore specimens to be conspecific. Species validity should be reassessedin future when more empirical evidence such
as DNA is made available for global populations of N. bourbonica.

Habitat/Ecology. This species was collected fromleaf litter in primary and/or old or mature secondary dryland forests, and swamp forest. Individuals were often numerous when found.
Remarks. Nylanderia kraepelini can be distinguished from other congeners by: 1) its relative small size (i.e., 1.3 – 1.4 mm), 2) microreticulate head and gaster - in contrast with asmooth and shining mesosoma, 3) relatively
small eye — roughly 6-8 ommatidia along its
maximum diameter in profile.

Nylanderia clandestina (Mayr, 1870)

Nylanderia yerburyi (Forel, 1894)

Material examined. None available.
Material not physically examined. Queen andmale types – FOCOL2715-2716 (ZMHB) [asPrenolepis clandestina].
Literature. Queen and male types – Viehmeyer(1916); Overbeck (1924) [as Prenolepis (Nylanderia) clandestina].
Localities. Bukit Timah Road
Habitat/Ecology. A colony was found under thebark of a fallen branch on the ground in agarden.
Remarks. While the species was first describedbased on a worker collected from Java (Mayr
1870), the queen and male were only describedlater on by Viehmeyer (1916) using specimens
collected by Overbeck in Singapore.

Material examined. None available.
Material not physically examined. Unknown.
Literature. Forel (1912) [as Prenolepis yerburyi].
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore context.
Remarks. At the time of writing, there is only asingle literature record of this species in Singapore (i.e., Forel 1912).

Nylanderia kraepelini (Forel, 1905)
Material examined. Dairy Farm Nature Park,
1.36385, 103.77731, 15 Oct 2021, W.N. Lamleg., ZRC_ENT00057794; MacRitchie Reservoir Park, 12-27 Oct 2021, same collector as
previous, ZRC_ENT00057796-57797, ZRC_
ENT00057800-57802; Pulau Ubin, 1.41984,
103.93272, 17 Nov 2021, same collector as
previous, ZRC_ENT00057795; workers andalate queen, Upper Peirce Reservoir Park,
11-27 Oct 2021, same collector as previous,
ZRC_ENT00057793, ZRC_ENT0005779857799, ZRC_ENT00057803-57804.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Dairy Farm; MacRitchie Reservoir;
Pulau Ubin; Upper Peirce Reservoir.

Oecophylla smaragdina (Fabricius, 1775)
Material examined. Bukit Batok East Avenue
6, 1.34304, 103.76235, 5 Oct 2016, G.W.
Yong leg., BB6, ZRC_HYM_0001477-1478;
same locality and collector as previous, 29
Sep 2016, ZRC_HYM_0001479-1481; Hougang Avenue 4, 1.34561, 103.88884, 14 Dec
2016, G.W. Yong leg., HA1GH914, ZRC_
HYM_0001488; same locality and collector as previous, 12 Dec 2016, HA3TH932,
ZRC_HYM_0001489; Icube, National University of Singapore, 1.293472, 103.77633,
May-Sep 2015, M.S. Foo & W. Wang leg.,malaise trap, ZRC_BDP (multiple); Prince
George’s Park Residences (NUS), 1.29239,
103.77869, Apr-Jul 2015, same collectors asprevious, malaise trap, ZRC_BDP (multiple);
University Town (NUS), 1.30622, 103.77458,
May-Sep 2015, same collectors as previous,
malaise trap, ZRC_BDP (multiple); KranjiRoad, 1.42638, 103.75413, 8-10 Nov 2016,
G.W. Yong leg., KR1, ZRC_HYM_0001461– 1466; same locality and collector as previous, 8 Nov 2016, KR4, ZRC_HYM_0001467;

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

same locality and collector as previous, 1
Nov 2016, KR8, ZRC_HYM_0001460;
Pulau Ubin, 7 Jan 2014, Sk. Yamane leg.,
ZRC_HYM_0000379; Seletar Link, 1.40518,
103.88555, 25 Oct 2016, G.W. Yong leg.,
SL4GH1019, ZRC_HYM_0001490; Sunset
Way, 1.32609, 103.77187, Aug-Sep 2016,
G.W. Yong leg., SW2, ZRC_HYM_00014691470; same locality and collector as previous, 23 Sep 2016, SW4GH309, ZRC_
HYM_0001471; same locality, collection
date and collector as previous, SW6GH356,
ZRC_HYM_0001472; same locality andcollector as previous, 21-23 Sep 2016, pitfall
trap, SW7, ZRC_HYM_0001172, 1473-1475;
waste woodland beside Temasek Polytechnic,
1.34239, 103.93598, 13-15 Dec 2016, G.W.
Yong leg., TT1, ZRC_HYM_0001482-1485;
same locality, collection date and collector as
previous, TT3, ZRC_HYM_0001486-1487.
Pulau Semakau New Fragment, 1°12›04.5››N,
103°45›46.1››E, 2012, J. Puniamoorthy et
al. leg., replanted mangroves, malaise trap,
SMN2, ZRC_BDP (multiple).
Material not physically examined. Non-types,
ANIC32-043666, 043928-043929, Tom Gush
Collection 3698/3714 (ANIC); ANIC32043930, Tom Gush Collection 3682 (ANIC).
Non-types, J.K. Wetterer leg. (USNM) – Katong, Amber Road, 28 Jul 2014, vial #340,
349; Marine Parade, 28 Jul 2014, vial #357;
City Hall, by church, 29 Jul 2014, vial #378;
Dhoby Ghaut, Istana Park, 30 Jul 2014, vials#406 – 418; Punggol Point, 30 Jul 2014, vial
#393; Katong, 1 Aug 2014, vials #443, 450;
Promenade, by MRT station, 4 Aug 2014, vial
#468; Pasir Panjang, by MRT station, 5 Aug
2014, vial #484.
Literature. Viehmeyer (1916), Overbeck (1924);
Tan & Corlett (2012); Yong et al. (2017);
Wang et al. (2018).
Localities. Amber Road (Katong); Bukit Batok
East; Bukit Timah Road; City Hall; Dhoby
Ghaut; Hougang; Katong; Kranji; Marine
Parade; National University of Singapore(NUS); Pasir Panjang; Promenade; Pulau Semakau New Fragment; Pulau Ubin; Punggol
Point; Seletar Link; Sunset Way, etc.

Habitat/Ecology. This species can be found in a
broad range of different habitats with varying
levels of disturbance in Singapore, including
cultivated vegetation in semi-urban settings
such as urban parks or overhead planters, also
in both mature and young secondary forestsand even mangroves. In parks, individuals
have been seen on stone surfaces and foliage, sometimes in or on leaf litter. The species
is known to create rather conspicuous nests
comprising a ball-like structure of interwoven
leaves and silken substrate, typically in bushes
or shrubs pavement-side. Occasionally, large
nests have been spotted in the crowns of large
dipterocarp trees, which are typically mistaken for hornet hives by observers from afar.
Remarks. A highly aggressive and behaviourally
dominant species that is apparently used forpest control in plantations such as oil palm.
Overbeckia subclavata Viehmeyer, 1916
Material examined. None available.
Material not physically examined. Types– FOCOL2564-2568 (ZMHB) [w.q.m.];
FOCOL0130 (DEIC); CASENT0905180(MSNG) [male]; CASENT0178502(MCZ). Non-types – CASENT0101191-92,
ANTC3162-63 (NHMB, Basel, Switzerland).
Literature. Type – Viehmeyer (1916); Overbeck(1924).
Localities. Bukit Timah Road
Habitat/Ecology. Colonies, including that of thetype series, were found in withered bamboo,
and in hollow branches of mangosteen trees
in a garden.
Remarks. Type locality in Singapore.
Paraparatrechina malaccana (Viehmeyer, 1916)
stat. n.
Prenolepis (Nylanderia) butteli subsp. malaccana
Viehmeyer, 1916: 147 (w.)
Combination in Paratrechina (Nylanderia): Emery
1925: 220.
Combination in Paraparatrechina: LaPolla et al.
2010: 128.

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Material examined. MacRitchie Reservoir, 9
Jan 2014, Sk. Yamane leg., SG14-SKY-39,
ZRC_HYM_0000439; Island Club forest (sic;in MacRitchie Reservoir), 20 May 1985, D.H.
Murphy leg., ZRC_HYM_0000774; Central
Catchment Nature Reserve, 10 Sep 2011,
E.J.Y. Soh leg., ZRC_ENT00048008; Upper
Thomson Nature Park, Sep-Oct 2016, G.W.
Yong et al. leg., ZRC_ENT00057813-57817;
Bukit Batok East Avenue 6, 27-29 Sep 2016,
G.W. Yong et al. leg., ZRC_ENT00049050,
57818; Mandai Road, Nov-Dec 2016, G.W.
Yong et al. leg., ZRC_ENT00057819-57821;
Sunset Way, 22 Aug 2016, G.W. Yong leg.,
ZRC_ENT00049056-49057.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Batok; Central Catchment Nature Reserve; MacRitchie Reservoir; Mandai
Road; Sunset Way; Upper Thomson Nature
Park.
Habitat/Ecology. This species was found inboth young and old/mature secondary forest, largely on shrubs or foliage. Individuals
were sometimes collected by manual beatingof vegetation, and in one instance, in a pitfall
trap.
Remarks. Paraparatrechina malaccana wasoriginally recognized as a subspecies of P.
butteli (Forel, 1913). Based on our assessmentof local specimens and comparisons with online type images and original descriptions,
we believe that the subspecies malaccana isdistinct from P. butteli and should be raised
to species.
		Workers of both species can be differentiated based on the following (P. butteli traits
in parentheses): 1) head rounded in full-faceview, sides convex, eyes very wide apart fromeach other, positioned above transverse midline of head (head more elongate-oval, longer than wide, eyes positioned at transverse
midline of head), 2) head with bluish-green
sheen, rest of body mostly brownish with afaintly-metallic purple or violet sheen (headand body largely uniformly blackish brown orbrown and shiny, but shimmer not coloured asin P. malaccana), 3) meso- and meta- pleura
mostly smooth and shining (same lateral faces
finely reticulate and weakly shining), 4) mid-and hind tibiae and tarsi pale-whitish, distal
ends of femora articulating with those tibiae
also distinctly pale-whitish, tibiae with fainttraces of brown – intensity of brown traces
varies among individuals (mid- and hindtibiae brown, distal ends of femora mostly
pale-brown and not strongly contrasting withrest of brown femur, tarsi pale-whitish).
		
We have also observed isometric size
polymorphism amongst individuals from thesame sample, with the smallest individualsof total length ~ 1mm, and larger individuals
about 1.5 – 2 mm.
Paraparatrechina opaca (Emery, 1887)
Material examined. Upper Peirce Reservoir, 10
Jan 2014, Sk. Yamane leg., SG14-SKY-52,
ZRC_HYM_0000466; Upper Thomson
Nature Park, 1.38311, 103.79839, 9-11
Oct 2016, G.W. Yong leg., UT6, ZRC_
HYM_0001686-1689; same locality, collection date/s and collector as previous, UT1,ZRC_HYM_0001690-1691.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Upper Peirce Reservoir; Upper Thomson Nature Park.
Habitat/Ecology. This species is associated mostly with primary or mature secondary habitatin Singapore, also sometimes in young secondary habitat such as abandoned plantation
secondary forests. These ants may be foundforaging on tree trunks or in low foliage.
Remarks. Based on original descriptions (seeEmery 1900) and type images on AntWeb,
the subspecies metallescens differs from P.
opaca in terms of having pale leg joints andbasal half of antennal scape; in the type of
P. opaca, leg joints are as brown as the restof the leg, basal half of antennal scape notso pale. We have, however, observed thatthe aforementioned characters can occur inintermediate states among nest mates of P.
opaca, thus these traits may not accurately
distinguish between species/subspecies.
		Moreover, according to Emery (1900),
the subspecies metallescens can also be distinguished from P. opaca by being smaller(i.e., TL 2.25 mm compared to 2.75 mm) ,Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

and mesosoma having a purplish or greenish
metallic sheen. The mesosoma of P. opaca
specimens we examined however, also had ablue-greenish metallic sheen, despite them nothaving other supposedly diagnostic charactersof subspecies metallescens mentioned in theprevious paragraph.
		Considering the above observations, andthe isometric size variation shown in otherspecies of this genus, it is possible that thesubspecies metallescens may be a synonymous variant of P. opaca. We lack adequate
material from a wider geographic range necessary to make definite taxonomic conclusions
concerning the two forms in this study.
Paratrechina longicornis (Latreille, 1802)
Material examined. Bukit Timah Nature Reserve,
8 Sep 2005, collector unknown, yellow pan
trap, ZRC_HYM_0000163; CommonwealthAvenue West, 21 Nov 2006, HK Lua leg., LHK464, ZRC_HYM_0000167; Commonwealth
Road, 1.31115, 103.78171, 22-24 Nov 2016,
G.W. Yong leg., CW3/4, ZRC_HYM_00015251527; Hougang Avenue 4, 1.34561, 103.88884,
12-14 Dec 2016, G.W. Yong leg., trunk pitfall trap, HA4TPd952, ZRC_HYM_0001528;
workers and males, National University of Singapore campus, Apr-Sep 2015, M.S. Foo & W.
Wang leg., malaise trap, ZRC_BDP (multiple);
Kent Ridge Park, 11 Jan 2014, Sk. Yamane leg.,
ZRC_HYM_0000400; Kent Ridge, 6 Jan 1983,
D.H. Murphy leg., ZRC_HYM_0000621; NeeSoon Swamp Forest, 5 Apr 2005, P. Grootaertleg., malaise trap, PGT 25046-25048, ZRC_
HYM_0000166; Pulau Ubin, 7 Jan 2014, Sk.
Yamane leg., ZRC_HYM_0000401; Sungei
Buloh, Feb 2015, R.Z.B. Quek leg., mangrove,
ZRC_ENT00007862; Sunset Way, 1.32609,
103.77187, Aug-Sep 2016, G.W. Yong leg.,
SW1/4/5, ZRC_HYM_0001491-1512.
Material not physically examined. Non-types,
ANIC32-010873 (in timber imported fromSingapore), ANIC32-053856 (ANIC). Nontypes, J.K. Wetterer leg. (USNM) – Bishan,
31 Jul 2014, vial #423; Buona Vista, by MRT
station, 5 Aug 2014, vial #485; City Hall, by
church, Jul-Aug 2014, vials #376, 489; Dhoby

Ghaut, Istana Park, 30 Jul 2014, vial #403;
Hougang, 30 Jul 2014, vial #402; Kallang,Riverside Park, 29 Jul 2014, vial #373; Katong, Jul-Aug 2014, vials #328, 334, 349, 444,
447; Kranji, by MRT station, 2 Aug 2014, vial
#452; Little India, 27 Jul 2014, vial #330; Marine Parade, Jul-Aug 2014, vials #353, 370,
436; Mountbatten, by MRT station, 4 Aug
2014, vial #464; Pasir Panjang, by MRT station, 5 Aug 2014, vial #481; Promenade, by
MRT station, 4 Aug 2014, vial #468; Punggol, by MRT station, 30 Jul 2014, vial #391;
Queenstown, 29 Jul 2014, vials #382, 384;
Serangoon, 30 Jul 2014, vial #389; Singapore Botanic Gardens, by MRT station, 5 Aug
2014, vial #476; Tai Seng, by MRT station, 31
Jul 2014, vial #422.
Literature. Wetterer (2008); Tan & Corlett (2012);
Yong et al. (2017); Wang et al. (2018a).
Localities. Bukit Timah Nature Reserve; Commonwealth Avenue West; Commonwealth
Road; Dhoby Ghaut; Fort Canning Park;
Hougang; Katong; Kent Ridge; Kranji; Little
India; Marine Parade; Mountbatten; National
University of Singapore campus; Nee Soon
Swamp Forest; Pasir Panjang; Promenade;
Pulau Ubin; Punggol; Queenstown; Serangoon; Singapore Botanic Gardens; Sungei
Buloh; Sunset Way; Tai Seng.
Habitat/Ecology. This infamous invasive pest is
mostly associated with areas of or close to human disturbance in Singapore, such as parks
in semi-urban settings and urban residential
areas. The ants are also common in young secondary forest fragments and at forest edges,
for example waste woodlands and abandoned
plantation secondary forests. The species appears tolerant of a broad range of different
environment stressors, and has been collectedfrom inland forest fragments, swamp forest,
even mangroves.
Remarks. A well-known widespread global tramp
species spread by human commerce and often
ubiquitous in heavily disturbed environments,
the earliest known record of P. longicornis in
Singapore is a specimen in the BMNH dated
back to 1931 (Wetterer 2008).

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Plagiolepis bicolor Forel, 1901
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah Road
Habitat/Ecology. Colonies of this species were
found under bark or in hollow twigs of mangosteen trees.
Remarks. At the time of writing, there are only
literature records of this species in Singapore.
Plagiolepis exigua Forel, 1894
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah Road
Habitat/Ecology. Garden – colonies were foundin withered shoots of bamboo near ground,between long fibres of withered bamboo, in
dry fallen branches, and in hollow twigs ofmangosteen trees.
Remarks. At the time of writing, there are only
literature records of this species in Singapore.
Polyrhachis (Campomyrma) creusa Emery, 1897
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[both as Polyrhachis (Myrmhopla) hecuba].
Localities. Bukit Timah Road
Habitat/Ecology. A colony was found in witheredbamboo in a garden.
Remarks. At the time of writing, there are only
literature records of this species in Singapore.
Polyrhachis (Chariomyrma) arcuata (Le Guillou, 1842)
Material examined. University Campus (Bukit
Timah), 9 Oct 1975, D.H. Murphy leg., ZRC_
HYM_0000895; same locality and collector as
previous, 7 Nov 1977, ZRC_HYM_0000979.
Material not physically examined. Unknown.
Literature. Forel (1907); Viehmeyer (1916);
Overbeck (1924).
Localities. Bukit Timah Road; University Campus(Bukit Timah).

Habitat/Ecology. This species was found in gardens or patches of spontaneous vegetation insemi-urban settings.
Remarks. Polyrhachis arcuata might be morecommon than is represented by samples
available for examination and in literature,especially in parks or other cultivated or spontaneous vegetation patches in the larger urban
matrix.
Polyrhachis (Chariomyrma) modesta Smith,
1857
Material examined. None available.
Material not physically examined. Holotype
queen – CASENT0901853, ANTC21297(OUMNH).
Literature. Type – Smith (1857). Emery (1896).
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore context.
Remarks. Type locality in Singapore. This species was first described based on the holotype alate queen collected from Singapore.
Neither worker nor male descriptions havebeen published.
Polyrhachis (Cyrtomyrma) laevissima Smith,1858
Material examined. Tyersall Istana, 9 Jun 1985,
D.H. Murphy leg., ZRC_HYM_0000839-841;
Fort Siloso (Sentosa), 16 Dec 2016, G.W.
Yong leg., ZRC_HYM_0001045; Bukit Timah, railway track, 12 Dec 2008, H.K. Lualeg., ZRC_ENT00048390.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[as Polyrhachis levissima].
Localities. Bukit Timah; Fort Siloso (Sentosa);
Tyersall Istana.
Habitat/Ecology. This species appears to be associated with cultivated or spontaneous secondary vegetation in semi-urban settings.
Colonies were found in withered bamboo ingardens, also in shrub foliage surrounding anabandoned building.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Polyrhachis (Cyrtomyrma) lepida Kohout, 2006
Material examined. Mandai, 1.40144,
103.77702, 7 Apr 2016, collector unknown,
ZRC_ENT00028312; Upper Peirce Reservoir, 10 Jan 2014, Sk. Yamane leg., ZRC_
HYM_0000397-398; MacRitchie Reservoir,
9 Jan 2014, Sk. Yamane leg. (SKYC).
Material not physically examined. FMNHINS0002842043, tc1357710853 (FMNH).
Literature. None. New record.
Localities. MacRitchie Reservoir; Mandai; Upper
Peirce Reservoir.
Habitat/Ecology. This species is associated withprimary or mature secondary habitat in Singapore, including mature/old secondary forest.
Polyrhachis (Myrma) beccarii Mayr, 1872
Material examined. None available.
Material not physically examined. Unknown.
Literature. Mezger & Moreau (2015).
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore
context.
Remarks. At the time of writing, there is onlya single literature record of this species in
Singapore (see Mezger & Moreau 2015) –no specific locality or collection data were
given in publication. Considering that we areunable to verify and validate the source andidentity of the specimen used in Mezger &Moreau’s (2015) phylogeny, we cannot ruleout the possibility that this may have been amisidentified specimen.
Polyrhachis (Myrma) carbonaria Smith, 1857
Material examined. None available.
Material not physically examined.
CASENT0919849, ANTC42479 (CASC).
Literature. None. New record.
Localities. Bukit Timah (alt. 100 ft.; possibly
Bukit Timah Hill)
Habitat/Ecology. Unknown in the Singapore
context – exact information unavailable fromoriginal label of unexamined material. However, it may be inferred that the ant was foundin either mature secondary or primary forest,since it was possibly collected from Bukit
Timah Hill, which is an established nature
reserve with both forest types.
Remarks. At the time of writing, the only recordof this species is that of a single worker from
Bukit Timah collected in 1962 by E.S. Ross
and D.Q. Cavagnaro, species identity determined by the late Rudy Kohout.
Polyrhachis (Myrma) hosei Donisthorpe, 1942
Material examined. Kent Ridge Park, 11 Jan 2014,
Sk. Yamane leg., ZRC_HYM_00000480.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Kent Ridge Park
Habitat/Ecology. The ants were found foraging
on spontaneous vegetation in a semi-urban
park setting.
Remarks. This species is currently known fromSingapore based on specimens from only one
locality, but may be more ubiquitous than implied by apparent sampling occurrence.
Polyrhachis (Myrma) illaudata Walker, 1859
Material examined. Bukit Timah, Hindhede Drive, 17 Apr 1988, H.K. Lua leg.,
ZRC_ENT00000331; Bukit Timah Nature Reserve, BT01, 1.35764, 103.77491, 1
Feb 2017, W. Wang leg., WW-SG17-039,
ZRC_ENT00007569; Bukit Timah forest, 3 Nov 1973, D.H. Murphy leg., ZRC_
HYM_0000854, 857-860; Bukit Timah Nature Reserve, BT05, 1.35386, 103.77988,
ZRC_HYM_0001753; Bukit Timah (University Campus), 7 Nov 1977, D.H. Murphy leg.,
ZRC_HYM_0000861; Commonwealth Road,
1.31115, 103.78171, 22 Nov 2016, G.W.
Yong coll., tree trunk, CW2TH668, ZRC_
HYM_0001589; same locality and collector
as previous, 24 Nov 2016, CW3TH708, ZRC_
HYM_00001592; Kranji Road, 1.42638,
103.75413, 10 Nov 2016, G.W. Yong leg.,
KR10GH1560, ZRC_HYM_0001597; MacRitchie Reservoir, 9 Jan 2014, Sk. Yamaneleg., ZRC_HYM_0000389; Mandai Road,
M3D, 1.41119, 103.80513, Nov-Dec 2016,
G.W. Yong leg., ZRC_HYM_0001593-1594;

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Mandai Road, M1D, 1.41258, 103.79839,
Nov-Dec 2016, same collector as previous,
ZRC_HYM_0001599-1600; same locality as previous, 4 Oct 2018, W. Wang leg.,
ZRC_ENT00047826; Prince George’s Park
Residences (NUS), 1.292389, 103.778694, 6
May 2015, M.S. Foo & W. Wang leg., malaise trap, NUS0021, ZRC_BDP0041454,
41412; Seletar Link, SL3, 1.40518,
103.88555, 18 Oct 2016, G.W. Yong leg.,
ZRC_HYM_0001595-1596; Sunset Way,1.32609, 103.77187, 29-31 Aug 2016, G.W.
Yong leg., ZRC_HYM_0001553; same locality and collector as previous, 21-23 Sep 2016,
ZRC_HYM_0001590-1591; Upper Peirce
Reservoir headquarters [sic], 10 Jan 2014, Sk.
Yamane leg., ZRC_HYM_00000390; Upper
Thomson Nature Park, 1.38311, 103.79839,
19 Oct 2016, G.W. Yong leg., UT5GH1640,
ZRC_HYM_0001598.
Material not physically examined. Non-types,
FMNHINS0002842133, tc1357712761;
FMNHINS0003259695, tc1357712757(FMNH); Harbourfront, Marang Trail, 4 Aug
2014, J.K. Wetterer leg., vial #473 (JKWC).
Literature. Tan & Corlett (2012).
Localities. Bukit Timah Nature Reserve; Bukit
Timah (University Campus); Commonwealth
Road; Kranji Road; Harbourfront; MacRitchie
Reservoir; Mandai Road; Prince George’s
Park Residences (NUS); Seletar Link; Sunset
Way; Upper Peirce Reservoir; Upper Thomson Nature Park.
Habitat/Ecology. This species is associated occasionally with primary forests, mostly withmature and/or old secondary habitats, andinfrequently with young secondary forest
fragments of varying levels of disturbance inSingapore. These include native-dominatedsecondary forest, abandoned plantation andwaste woodland secondary forests. Individuals can be found foraging on leaf litter, deadwood, fallen logs, tree trunks or low foliage.
Remarks. This species has a wide range of variation in the development and shape of petiole,spines, and also pilosity. A critical revision isneeded to clarify its status as a single species.

Polyrhachis (Myrma) inermis Smith, 1858
Material examined. MacRitchie Reservoir, 6 Jan
2014, Sk. Yamane leg., ZRC_HYM_0000481.
Material not physically examined. FMNHINS0002842039, tc1357710822 (FMNH).
Literature. None. New record.
Localities. Kent Ridge Park; MacRitchie Reservoir.
Habitat/Ecology. This species appears to be associated with either young or mature secondary
forests in Singapore.
Polyrhachis (Myrma) nigropilosa Mayr, 1872
Material examined. MacRitchie, NS 184, 9
Jun 1995, S.L. Goh & B.K.H. Tan leg.,
ZRC_ENT00000373; Mandai, 1.4083,
103.77803, 12 Nov 2015, M.K.L. Wong leg.,
ZRC_ENT00000733; Mandai Road, 1.41155,
103.79015, 23 Nov 2016, G.W. Yong leg.,
M2D2GH1961, ZRC_HYM_0001177; Upper Peirce Reservoir, 10 Jan 2014, Sk. Yamaneleg., SG14-SKY-47, ZRC_HYM_0000462;
alate queen, Nee Soon Swamp Forest, NS2,
1°23’04.2”N, 103°48’40.7”E, 28 Nov-5 Dec
2012, J. Puniamoorthy et al. leg., malaise trap,
Reg. 29611, ZRC_BDP0009132.
Material not physically examined. Unknown.
Literature. Menozzi (1926) [as Polyrhachis nigropilosa conophthalma]; Mezger & Moreau(2015).
Localities. MacRitchie; Mandai; Nee Soon Swamp
Forest; Upper Peirce Reservoir.
Habitat/Ecology. This arboreal species is associated mostly with primary or mature secondary
habitats, such as native-dominated secondaryand swamp forests, as well as young secondary forests in Singapore. Nests have beenfound in decayed branches.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Polyrhachis (Myrma) obesior Viehmeyer, 1916
Material examined. None available.
Material not physically examined. Unknown.
Literature. Type – Viehmeyer (1916); Overbeck(1924) [both as Polyrhachis mayri obesior].
Kohout (1998).
Localities. Bukit Timah Road
Habitat/Ecology. Specimens of the type series
were collected in a garden.
Remarks. Type localities in Singapore and Malaysia (i.e., Gunong Angsi) (Viehmeyer 1916).
At the time of writing, the only record of thisspecies in Singapore is that of the type series.
Polyrhachis (Myrma) orsylla Smith, 1861
Material examined. None available.
Material not physically examined. Unknown.
Literature. Forel (1907).
Localities. Unknown.
Habitat/Ecology. No information available in the
Singapore context.
Remarks. Specimens identified as ‘P. orsyllus’ inViehmeyer (1916) and Overbeck (1924) werecollected by Overbeck in Gunong Angsi – alocality in Malaysia and not Singapore.
Polyrhachis (Myrma) orsylla ritsemai Mayr, 1883
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[as Polyrhachis orsyllus ritesmai].
Localities. Bukit Timah Road
Habitat/Ecology. The ants were found nesting inwithered bamboo in a garden.
Remarks. At the time of writing, there are only
literature records of this species in Singapore.
Polyrhachis (Myrma) proxima Roger, 1863
Material examined. Bukit Batok East Avenue 6,
BB2, 1.34304, 103.76235, Sep 2016, G.W.
Yong leg., ZRC_HYM_0001557, 1567; Commonwealth Road, 1.31115, 103.78171, 24
Nov 2016, G.W. Yong leg., tree trunk, ZRC_
HYM_0001554, 1568; Hougang Avenue 4,
1.34561, 103.88884, 12 Dec 2016, G.W. Yongleg., HA5GH990, ZRC_HYM0001572; Kent

Ridge, scrub, 13 May 1985, D.H. Murphyleg., ZRC_HYM_0000927-929; Kranji Road,
1.42638, 103.75413, Nov 2016, G.W. Yongleg., tree trunk, ZRC_HYM_0001580-1582;
Lorong Banir, 29 Jun 1995, collector unknown,
NS 193, ZRC_ENT00000358; Mandai Road,
1.41119, 103.80513, Nov-Dec 2016, G.W.
Yong leg., M3D, ZRC_HYM_0001569-1571;
National University of Singapore campus
grounds, Apr-Sep 2015, M.S. Foo & W. Wangleg., malaise trap, ZRC_BDP (multiple); Seletar Link, 1.40518, 103.88555, Oct 2016, G.W.
Yong leg., ZRC_HYM_0001573-1579; Sunset Way, 1.32609, 103.77187, Aug-Sep 2016,
G.W. Yong leg., ZRC_HYM_0001555-1556,
1560-1566; Tyersall Istana, 9 Jun 1985, D.H.
Murphy leg., ZRC_HYM_0000863; Upper
Peirce Reservoir, 10 Jan 2014, Sk. Yamaneleg., ZRC_HYM_0000391; Upper Thomson
Nature Park, 1.38311, 103.79839, Sep-Oct
2016, G.W. Yong leg., ZRC_HYM_00015581559, 1583-1588.
Material not physically examined. FMNHINS0002842042, tc1357710848 (FMNH).
Literature. Forel (1893); Viehmeyer (1916);
Overbeck (1924); Wang et al. (2018a).
Localities. Bukit Batok East Avenue 6; Bukit Timah, Bukit Timah Nature Reserve; Commonwealth Road; Hougang Avenue 4; Kent Ridge;
Kranji Road; Lorong Banir; MacRitchie Reservoir; Mandai Road; National University ofSingapore campus grounds; Seletar Link;
Sunset Way; Tyersall Istana; Upper Peirce
Reservoir; Upper Thomson Nature Park.
Habitat/Ecology. This species is associatedmainly with both mature and young secondary habitats at varying levels of disturbancein Singapore, including native-dominated,
waste woodland and abandoned plantation
secondary forests, often fragments in a larger
semi-urban matrix. Individuals have also beenfound foraging in urban cultivated areas on
grassy patches, or semi-urban forest fringes
on foliage in scrubland.
Remarks. While species occurrences appear
widespread in Singapore, most forest fragments where P. proxima populations thrive
are highly likely under threat of future urban
development.

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Polyrhachis (Myrma) pruinosa Mayr, 1872

Polyrhachis (Myrma) villipes Smith, 1857

Material examined. Kent Ridge Park, 7 Dec
2017, Sk. Yamane leg., ZRC_ENT0000883.
Material not physically examined. Unknown.
Literature. Forel (1913); Viehmeyer (1916);
Overbeck (1924).
Localities. Ayer Terjun (archaic place name);
Bukit Timah; Kent Ridge Park; Jurong Road;
Mandai Road.
Habitat/Ecology. This species was found mostly
in disturbed secondary forests, frequently in
Nepenthes pitcher cups at Ayer Terjun.

Material examined. Bukit Timah forest,
3 Nov 1973, D.H. Murphy leg., ZRC_
HYM_0000966; Island Club forest, 20
May 1985, D.H. Murphy leg., ZRC_
HYM_0000967.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah forest; Bukit Timah Road;
Island Club forest.
Habitat/Ecology. This species appears associatedwith primary and/or mature secondary forest.
A single queen was also collected from a lit
lamp in a garden.

Polyrhachis (Myrma) striata Mayr, 1862
Material examined. None available.
Material not physically examined. Unknown.
Literature. Forel (1893).
Localities. Unknown.
Habitat/Ecology. Unknown in Singapore context.
Remarks. Specimens identified from Overbeck’s
collections as P. striata in Viehmeyer (1916)
were collected from Gunong Angsi in Malaysia, not Singapore. Thus, at the time of writing, there is only a single known literaturerecord of this species in Singapore, identified
from Arthur Müller’s collections (Forel 1893).
Polyrhachis (Myrma) striatorugosa Mayr, 1862
Material examined. ; Mandai Lake Road, 3 Nov
2016, G.W. Yong leg., ZRC_HYM_0001042;
Nee Soon Swamp Forest, 28 Dec 1989, Y.H.
Koo & K. Snyder leg., ZRC_ENT00047921.
Material not physically examined. FMNHINS0002842045, tc1357712546 (FMNH).
Literature. None. New record.
Localities. Mandai Lake Road; Nee Soon Swamp
Forest; Upper Peirce Reservoir.
Habitat/Ecology. This species was found in freshwater swamp forest, also primary or old secondary forest.
Remarks. A member of the P. relucens species
group that includes P. nigropilosa (Kohout2008), P. striatorugosa may be differentiatedfrom the latter by its finely striated gaster.
This species is also similar to P. sculpturata
Smith, 1860, but with a distinct (though small)intercalary tooth on dorsal margin of petiole.

Polyrhachis (Myrma) vindex Smith, 1857
Material examined. Bukit Timah Nature Reserve, 3 Nov 1973, D.H. Murphy leg., forest, ZRC_HYM_0000933; same locality
and collector as previous, area K5, 13 May
1967, ZRC_HYM_0000934; Sime Road
forest, 8 Aug 1978, D.H. Murphy leg.,
ZRC_HYM_0000932; Island Club forest,
20 May 1985, D.H. Murphy leg., ZRC_
HYM_0000936; Kent Ridge, 18 Mar 1985,
D.H. Murphy leg., ZRC_HYM_0000935;
same locality and collector as previous, 28
May 1985, ZRC_HYM_0000937.
Material not physically examined.
CASENT0217808, ANTC17997 (CASC).
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Island
Club forest; Kent Ridge; Seletar Reservoir;
Sime Road forest.
Habitat/Ecology. This species is associated mostly with primary or mature secondary forests
in Singapore, including native-dominated secondary forest. The ants have also been foundin disturbed young secondary forest fragments
in semi-urban settings. Individual workers are
sometimes collected from foliage or low vegetation via sweep-netting.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Polyrhachis (Myrmatopa) charaxa Smith, 1860
Material examined. None available.
Material not physically examined. Unknown.
Literature. Emery (1896); Viehmeyer (1912)[both as Polyrhachis charaxus].
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore context.
Remarks. At the time of writing, there are only
literature records of this species in Singapore.
Polyrhachis (Myrmatopa) flavicornis Smith,
1857
Material examined. None available.
Material not physically examined. Syntypes –CASENT0901857, ANTC21301 (OUMNH);
alate queen, CASENT0901877, ANTC21321(OUMNH).
Literature. Type – Smith (1857). Smith (1858);
Emery (1896).
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore
context.
Remarks. Type locality in Singapore, but specificlocality name and habitat unknown.
Polyrhachis (Myrmatopa) leviuscula Viehmeyer,
1916
Material examined. Sungei Buloh, 8 Jan 2014,
Sk. Yamane leg., SG14-SKY-27, ZRC_
HYM_0000461.
Material not physically examined. Types –CASENT0910788, ANTC33049 (MHNG);
FOCOL2554 (ZMHB) [as types of Polyrhachis (Myrmatopa) schang var. leviuscula].
Non-type – CASENT0906574, ANTC24598(BMNH).
Literature. Type – Viehmeyer (1916); Overbeck(1924) [both as P. schang var. leviuscula].
Localities. Buki Timah Road; Sungei Buloh.
Habitat/Ecology. Specimens of the type series
were collected with a sweep net in a garden,
thus we may infer that the ants forage on low
foliage. A colony ‘of a very similar form’ wasalso found on a palm leaf in the same garden(Overbeck 1924), but these have not beenverified to be conspecific to P. leviuscula.

		More recently (from the time of writing),
the ants were found nesting in a rolled leaf inback mangrove forest.
Remarks. Type locality in Singapore. Physically unexamined non-type material (i.e.,
CASENT0906574) was collected by Dr. G.E.
Brooke in 1912, preceding publication of theoriginal species description, and was initially
identified as P. schang var. laurae (now a junior synonym of P. leviuscula).
		The more recent specimens mostly confer with P. leviuscula in appearance, thoughwith slightly weaker dorsolateral mesonotal
projections/‘teeth’. The species resembles
Polyrhachis (Myrmatopa) solmsi Emery,
1887, but the latter has petiolar spines thatare almost supine and more strongly directed
posteriorly in profile view.
Polyrhachis (Myrmatopa) piliventris Smith, 1858
Material examined. None available.
Material not physically examined. Holotype
alate queen – CASENT0903307, ANTC23165(BMNH).
Literature. Type – Smith (1858). Emery (1896).
Localities. Unknown.
Habitat/Ecology. Unknown in Singapore context.
Remarks. Type locality in Singapore; this species
was first described based on the holotype alate
queen collected from Singapore. At the timeof writing, the holotype queen remains the
only known physical record of P. piliventris
in Singapore.
Polyrhachis (Myrmatopa) varicolor Viehmeyer,
1916
Material examined. None available.
Material not physically examined. Syntype series deposited at MNHU, Berlin.
Literature. Type – Viehmeyer (1916); Overbeck(1924) [both as Polyrhachis fruhstorferi
subsp. varicolor]. Kohout (2008) [raised to
species].
Localities. Bukit Timah Road
Habitat/Ecology. Specimens of the type series
were collected from a garden.

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Remarks. At the time of writing, the type series
is the only known record of this species in
Singapore. Type locality in Singapore.
Polyrhachis (Myrmhopla) abdominalis Smith,1858
Material examined. Bukit Timah, 1.347957,
103.779142, 29 Oct 2015, M.K.L. Wong leg.,
ZRC_ENT00000737; Bukit Timah, Hindhede
Drive, 30 May 1988, H.K. Lua leg., ZRC_
HYM_0000078; Hougang Avenue 4, HA5,
1.34561, 103.88884, 12-14 Dec 2016, G.W.
Yong leg., ZRC_HYM_0001601, 1603-1605;
Kent Ridge Park, 11 Jan 2014, Sk. Yamane,
ZRC_HYM_0000394; Kranji Road, 1.42638,
103.75413, 1-10 Nov 2016, G.W. Yong leg.,tree trunk, ZRC_HYM_0001609-1612;
Lorong Banir, 29 Jun 1995, collector unknown, NS 193, ZRC_ENT00000359; MacRitchie Reservoir, 6 Jan 2014, Sk. Yamaneleg., ZRC_HYM_0000395; Mandai Road,
1.41258, 103.79839, 5 Dec 2016, G.W. Yongleg., M1D19GH2572, ZRC_HYM_0001615;
Nee Soon Swamp Forest, plot Q9, 1.398925,
103.808608, 24 Sep 2019, W.N. Lam et al.
leg., nest in hollows of PVC pipes, WWSG19-007, ZRC_ENT00013881; Prince
George’s Park Residences (NUS), 1.292389,
103.778694, Apr-Sep 2015, M.S. Foo & W.
Wang leg., malaise trap, ZRC_BDP (multiple); Pulau Ubin, 14 Jun 2018, W. Wangleg., mangrove, nest in hollow of metal
piping of malaise trap, WW-SG18-Poly1,
ZRC_ENT00007299; Seletar Link, 1.40518,
103.88555, 16-15 Oct 2016, G.W. Yong leg.,
ZRC_HYM_0001606-1608; University Hall
(NUS), 1.297111, 103.776583, May-Aug
2015, M.S. Foo & W. Wang leg., malaise trap,
ZRC_BDP (multiple); Upper Thomson Nature Park, 1.38311, 103.79839, 9-11 Oct 2016,
G.W. Yong leg., ZRC_HYM_0001613-1614.
Material not physically examined. FMNHINS0002842037, tc1357710809; FMNHINS0002842049, tc1357710875 (FMNH).
Literature. Mezger & Moreau (2015); Wang etal. (2018a).

Localities. Bukit Timah Nature Reserve; Hougang
Avenue 4; Kent Ridge Park; Kranji Road; Lorong Banir; MacRitchie Reservoir; Mandai;
Nee Soon Swamp Forest; Prince George’s
Park Residences (NUS); Pulau Ubin; Seletar
Link; University Hall (NUS); Upper Peirce
Reservoir; Upper Thomson Nature Park.
Habitat/Ecology. This species is associatedmostly with both mature and young secondary habitats at varying levels of disturbance inSingapore. These include swamp forest, mangroves, native-dominated, abandoned plantation and waste woodland secondary forests.
Individual foragers are typically collectedfrom tree trunks, while both alates and workers have been collected using malaise traps
from disturbed secondary forest fragments insemi-urban settings. While considered arboreal, this species has been found nesting in unconventional places, e.g. in hollows of PVC or
metal pipes - parts of artifical infrastructures
such as malaise or leaf litter traps.
Polyrhachis (Myrmhopla) armata (Le Guillou,
1842)
Material examined. Bukit Timah forest, 3 Nov
1973, D.H. Murphy leg., ZRC_HYM_0000850,
852; Istana Club forest, 20 May 1985, D.H.
Murphy leg., ZRC_HYM_0000848-849; alate
queen, University Town (NUS), 26 Apr 2019,
N. Chin & J. Tan leg., ZRC_ENT00013540;
Prince George’s Park Residences, 1.292389,
103.778694, Aug 2015, M.S. Foo & W. Wangleg., malaise trap, ZRC_BDP0047321, 47427;
Singapore Botanical Gardens, Sep 1965, collector unknown, ZRC_HYM_0000853; Upper
Peirce Reservoir, 10 Jan 2014, Sk. Yamane leg.,
ZRC_HYM_0000392-393.
Material not physically examined.
CASENT0281398,
PSW10255-3(PSWC); FMNHINS0002842132, FMNHINS0003249694, ESJY-U-0024 (FMNH).
Literature. Viehmeyer (1916); Overbeck (1924);
Tan & Corlett (2012); Wang et al. (2018a).
Localities. Bukit Timah; Bukit Timah Hill; Istana
Club forest; Kent Ridge Park; Prince George’s
Park Residences (NUS); Singapore Botanical Gardens; University Town (NUS); Upper
Peirce Reservoir

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Habitat/Ecology. This arboreal species is associated mainly with mature and/or young secondary habitats, including disturbed secondary
forest fragments or even cultivated park areas
in urban or semi-urban settings. Individuals
including winged alates have sometimes beenfound in human infrastructure close to forest
fringes or fragments. In rainforest, the species
can be found foraging on lower vegetation.
Remarks. Polyrhachis armata appears similarin general appearance (and colouration) to P.
abdominalis, but can be distinguished fromthe latter by its coarser and more matte body
sculpture. Singapore is actually the type locality for a subspecies of P. armata - P. armata defensa [see next paragraph below]. It
is possible that subspecies is simply a morphological variant of the species, the moststriking variable feature being the conditionof the petiolar spines (length and thickness)– which can exhibit broad intranidal variation(W. Wang pers. obs.)
Polyrhachis (Myrmhopla) armata defensa Smith,
1857
Material examined. None available.
Material not physically examined. Syntype –CASENT0901845, ANTC21289 (OUMNH)[as Polyrhachis defensus].
Literature. Type – Smith (1857) [as P. defensus].
Emery (1925) [transferred to subspecies of
P. armata].
Localities. Unknown.
Habitat/Ecology. No information available in the
Singapore context.
Remarks. Type locality in Singapore. See Remarks in previous section (i.e., P. armata) on
distinction between P. armata and this subspecies.
Polyrhachis (Myrmhopla) bicolor Smith, 1858
Material examined. Kranji Road, 1.4238,
103.75413, 10 Nov 2016, G.W. Yong leg., tree
trunk, KR1TH1399, ZRC_HYM_0001177.2;
Sungei Buloh Wetland Reserve, 8 Jan 2014,
Sk. Yamane leg., nest between leaves,
SG14-SKY-28, ZRC_HYM_0000460;
Lim Chu Kang, 20 May 1976, C.S.J. leg.,

ZRC_HYM_0000906; Mandai mangrove,
2 Oct 1978, D.H. Murphy leg., ZRC_
HYM_0000976; same locality, collection dateand collector as previous, DHM-SG78-Poly1,
ZRC_ENT00014141; Prince George’s Park
Residences (NUS), 1.292389, 103.778694,
Apr 2015, M.S. Foo & W. Wang leg., malaise trap, ZRC_BDP0044269, 45587; Western Catchment, 1.408657, 103.701849,
20 Nov 2015, M.K.L. Wong leg., ZRC_
ENT00000731.
Material not physically examined.
CASENT0217750, ANTC1952 (CASC).
Literature. Viehmeyer (1916); Overbeck (1924);
Mezger & Moreau (2015).
Localities. Bukit Timah; Bukit Timah Road; Kranji Road; Lim Chu Kang; Mandai mangrove;
Sungei Buloh Wetland Reserve; Western
Catchment.
Habitat/Ecology. This species was mainly foundin mangroves, or young secondary habitats
including waste woodlands and disturbed
secondary forest fragments within urban orsemi-urban settings, in Singapore. In mangroves, nests were found between leaves, or in‘silk-webbed’ leaves of Sonneratia; individuals were sometimes collected from tree trunks
or foliage above ground.
Remarks. There is considerable morphologicalvariation between populations of this species. For the purposes of this checklist, we
tentatively treat all morphological variants asconspecific, pending more concrete empiricalevidence for species differentiation in future.
Polyrhachis (Myrmhopla) bicolor aurinasis
Forel, 1901
Material examined. None available.
Material not physically examined. Unknown.
Literature. Forel (1907).
Localities. Unknown.
Habitat/Ecology. No information available in the
Singapore context.
Remarks. The only literature record of this subspecies in Singapore is that of a single queen
collected by Biró (Forel 1907).

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Polyrhachis (Myrmhopla) chalybea Smith, 1857
Material examined. Jalan Kayu wasteland,
1 Feb 1975, D.H. Murphy leg., ZRC_
HYM_0000838; Lower Peirce Reservoir, Forest B, 10 Aug 1990, collector
unknown, ZRC_ENT00000203; Mandai
mangrove, 17 Oct 1978, D.H. Murphy leg.,
ZRC_HYM_0000836-837; Sime Road forest, 27 Jun 1982, D.H. Murphy leg., ZRC_
HYM_0000834; same locality as previous,
18 Jan 1976, D.H. Murphy leg., ZRC_
HYM_0000908; University Campus (Bukit
Timah), 17 Mar 1974, D.H. Murphy leg.,nutmeg tree, ZRC_HYM_0000835.
Material not physically examined. Unknown.
Literature. Type – Smith (1857) [as Polyrhachis
chalybeus].
Localities. Jalan Kayu; Lower Peirce Reservoir,
Forest B; Mandai mangrove; Sime Road; University campus (Bukit Timah).
Habitat/Ecology. This species is associated mainly with young secondary habitat of varying
levels of disturbance, including waste woodland secondary forest fragments in urban orsemi-urban settings, in Singapore. It is sometimes also found in mangrove back forest. Individuals were usually collected on trees ortree trunks.
Remarks. The holotype specimen depicted on AntWeb was inferred to be from Bacan island,
Indonesia (based on specimen label ‘Bac’),however, in the original description of ‘P. chalybeus’, only two other type localities were
given: Singapore and Malacca. It may be notable that an additional label on the holotype
specimen reads ‘Mal-ca’, which might refer to
Malacca, one of two type localities indicatedin Smith’s (1857) original account.
Polyrhachis (Myrmhopla) cryptoceroides Emery,
1887
Material examined. Rifle Range Road, 27 Jul
2016, G.W. Yong leg., ZRC_HYM_0001044.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Rifle Range Road
Habitat/Ecology. This species was found in old
secondary forest.

Remarks. Polyrhachis cryptoceroides can be
distinguished from other members of thecryptoceroides group by its medially deeply
emarginate anterior clypeal margin.
Polyrhachis (Myrmhopla) dives Smith, 1857
Material examined. Kent Ridge, 1981, D.H.
Murphy leg., ZRC_HYM_0000957; KranjiRoad, 10 Nov 2016, G.W. Yong leg., ZRC_
HYM_0001041.
Material not physically examined. Holotype –CASENT0903388, ANTC23251 (BMNH).
Literature. Type – Smith (1857). Kohout (2010).
Localities. Kent Ridge; Kranji Road.
Habitat/Ecology. This species is associated mostly with young secondary habitat in Singapore,including wasteland forest and disturbed
secondary forest fragments in urban or semiurban settings. Individuals have been foundforaging in shrubs and on grass. Known to bea polygynous, wide-ranging species that build
a distinctive carton nests on lower branches ofgrasses, trees and shrubs - foliage and twigs
are combined with larval silk to create thenest structure.
Remarks. Type locality in Singapore. The original description by Smith (1857) was based on
specimens collected by A.R. Wallace. Southeast Asian specimens of P. dives have been
observed to be slightly different morphologically from their Australian and New Guinean
counterparts - the former have a more deeply
emarginate anterior clypeal margin, and eyes
almost never protruding from the lateral cephalic outline.
Polyrhachis (Myrmhopla) hector Smith, 1857
Material examined. None available.
Material not physically examined. Holotype –CASENT0901811, ANTC21233 (OUMNH).
Literature. Type – Smith (1857). Emery (1896);
Viehmeyer (1912); Bolton (1974).
Localities. Unknown.
Habitat/Ecology. Unknown in Singapore context.
Remarks. Type locality in Singapore, but specificlocality name and habitat unknown.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Polyrhachis (Myrmhopla) hippomanes ceylonensis Emery, 1893

Polyrhachis (Myrmhopla) saevissima Smith,
1860

Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah Road
Habitat/Ecology. Garden.
Remarks. The only literature record of this subspecies in Singapore is that of a single queen
collected from a lamp in a garden (Viehmeyer
1916; Overbeck 1924).

Material examined. Upper Peirce Reservoir, 10
Jan 2014, Sk. Yamane leg., SG14-SKY-46,
ZRC_HYM_0000423; Pulau Ubin, PU3,
15-22 Dec 2012, J. Puniamoorthy et al.
leg., mangroves, malaise trap, Reg. 29646,
ZRC_BDP0016507; alate queen, Sungei
Buloh Wetland Reserve, SB1, 1°26’46.3”N,
103°43’49.9”E, 12-19 Dec 2012, J. Puniamoorthy et al. leg., mangroves, malaise trap,
Reg. 29634, ZRC_BDP0017223.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Pulau Ubin; Sungei Buloh Wetland
Reserve; Upper Peirce Reservoir.
Habitat/Ecology. This species is found in primaryor mature secondary habitats in Singapore,including mangroves where most occurrences have been reported. In primary or maturesecondary forests, nests were found between
leaves.
Remarks. At the time of writing, taxonomic validity of P. saevissima is uncertain with broad
morphological variation observed between
populations. While for the purposes of thischecklist we tentatively treat these variants
as conspecific, it is possible that these might
be proven to be different species with more
empirical evidence made available in future.

Polyrhachis (Myrmhopla) lucidula Emery, 1893
Material examined. None available.
Material not physically examined. Type queen– FOCOL2944 (ZMHB).
Literature. Types (queen and male) – Viehmeyer(1916); Overbeck (1924) [as Polyrhachis hippomanes subsp. lucidula].
Localities. Bukit Timah Road
Habitat/Ecology. Colonies of the species were
found in withered bamboo in a garden.
Remarks. The queen and male of this species werefirst described by Viehmeyer (1916) based on
specimens collected by Overbeck (1924) in
Singapore. Viehmeyer (1916) commented thatshould the species turn out to be a new ‘form’,
he would name it ‘lucidissima’ – a new varietyof the (then) subspecies lucidula.
Polyrhachis (Myrmhopla) muelleri Forel, 1893
Material examined. None available.
Material not physically examined. Syntype –CASENT0910868, ANTC33129 (MHNG).
Literature. Type – Forel (1893) [as Polyrhachis
mülleri]. Forel (1901, 1907); Emery (1925).
Localities. Unknown.
Habitat/Ecology. Unknown in Singapore context.
Remarks. Type locality in Singapore. The species
was first described as ‘P. mülleri’ in the original publication (Forel 1893), but the speciesname was later (since Emery 1925) adjusted
to the Latin alphabet, which comprises only
26 letters not including ‘ü’.

Polyrhachis (Myrmhopla) thailandica Kohout,
2006
Material examined. Mandai Road, 13 Jan 2017,
G.W. Yong leg., ZRC_HYM_0001043.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Mandai Road
Habitat/Ecology. This species was found in disturbed mature secondary forest.
Remarks. Polyrhachis thailandica can be differentiated from the similar-looking P. jerdonii
Forel, 1892 by the presence of a strong mediancarina on the clypeus.

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Polyrhachis (Myrmhopla) tibialis parsis Emery,
1900
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Ayer Terjun (archaic place name);
Bukit Timah Road.
Habitat/Ecology. These ants were found in a hollow fallen branch on the ground in a garden.
The species was also collected from Nepenthes pitcher cups, possibly in secondary forest
or similar spontaneous vegetation.
Remarks. At the time of writing, there are only
literature records of this subspecies in Singapore.
Polyrhachis (Myrmothrinax) frauenfeldi Mayr,
1862
Material examined. Mandai mangroves,
26 Jun 1979, D.H. Murphy leg., ZRC_
HYM_0000958.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah Road; Mandai mangroves.
Habitat/Ecology. An alate queen of this species
was collected from a lit lamp in a garden. The
ants have also been found in mangroves.
Polyrhachis (Myrmothrinax) saigonensis Forel,
1886
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[both as Polyrhachis thrinax subsp. saigonensis].
Localities. Bukit Timah Road.
Habitat/Ecology. Garden.
Remarks. The only literature record of this species
in Singapore is that of a single queen collectedfrom a lamp in a garden (Viehmeyer 1916;
Overbeck 1924).

Polyrhachis (Myrmothrinax) textor Smith, 1857
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Ayer Terjun (archaic place name)
Habitat/Ecology. Specimens of this species were
found in a Nepenthes pitcher cup.
Remarks. At the time of writing, there are only
literature records of this species in Singapore.
Polyrhachis (Myrmothrinax) thrinax inconstans
Viehmeyer, 1916
Material examined. None available.
Material not physically examined. Types –CASENT0910798, ANTC33059 (MHNG);
FOCOL2530-2531 (ZMHB).
Literature. Type – Viehmeyer (1916); Overbeck(1924).
Localities. Bukit Timah Road
Habitat/Ecology. Ants of this subspecies were
found in a pasteboard nest in the fold of apalm leaf in a garden; individual foragers werealso collected from around other parts of thegarden.
Remarks. Type locality in Singapore.
Polyrhachis (Myrmothrinax) thrinax overbecki
Dorow, 1995
Material examined. None available.
Material not physically examined. Types –CASENT0910797, ANTC33058 (MHNG);
FOCOL2527, FOCOL2529 (ZMHB); type
queen, FOCOL2528 (ZMHB) [all as types of
Polyrhachis thrinax var. nigripes].
Literature. Type – Viehmeyer (1916); Overbeck(1924) [as P. thrinax var. nigripes]. Dorow(1995).
Localities. Bukit Timah Road
Habitat/Ecology. A pasteboard nest of this subspecies was found in the twisted leaf of a
shrub in a garden.
Remarks. The present species epithet is a replacement name for P. thrinax nigripes Viehmeyer,
1916 (see Dorow 1995). Type locality in Singapore.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Polyrhachis (Polyrhachis) bellicosa Smith, 1859

Prenolepis jerdoni Emery, 1893

Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah Road
Habitat/Ecology. Specimens of this species werecollected from a garden.
Remarks. At the time of writing, this species isknown from Singapore based only on literature records.

Material examined. Bukit Timah Nature Reserve,
BT07, 1.35499, 103.78167, 28 Sep 2016, W.
Wang leg., leaf litter, winkler extraction,
ZRC_HYM_0000569; same locality, collection date and collector as previous, leaf litter,
winkler extraction, ZRC_HYM_0000570;
male, I-Cube building (NUS), 1.29347,
103.77633, 24 Mar-1 Apr 2015, M.S. Foo& W. Wang leg., malaise trap, NUS0002,
ZRC_BDP0044670; University Hall (NUS),
1.29711, 103.77658, 13-20 May 2015, M.S.
Foo & W. Wang leg., malaise trap, NUS0032,
ZRC_BDP0044799; males, Nee Soon Swamp
Forest, NS2, 1°23’04.2”N, 103°48’40.7”E, 28
Mar-4 Apr 2012, J. Puniamoorthy et al. leg.,malaise trap, Reg. 29158, ZRC_BDP0015151,
15168, 15924; same locality as previous,
1.38245, 103.80206, 46 m, 26 Apr 2018,
W. Wang leg., winkler extraction, NS_W2,
ZRC_ENT00000956.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924);
Wang et al. (2018a).
Localities. Ayer Terjun (archaic place name);
Bukit Timah Nature Reserve; Bukit Timah
Road; National University of Singapore campus; Nee Soon Swamp Forest.
Habitat/Ecology. This species is associatedmostly with old or mature secondary forests,
as well as young secondary habitats such
as Adinandra belukar dominated forest in
Singapore. Both alates (males and queens)
and workers have been collected, the former
in great abundance, using malaise traps inswamp forest. The ants can also be found in
disturbed secondary forest fragments in urban
or semi-urban settings.

Polyrhachis (Polyrhachis) olybria Forel, 1912
Material examined. Race Course forest,
22 Jun 1977, D.H. Murphy leg., ZRC_
HYM_0000971.
Material not physically examined. Bukit Timah
Nature Reserve, 30 Apr 1967, D.H. Murphyleg. (in Kohout 2014).
Literature. Kohout (2014).
Localities. Bukit Timah Nature Reserve; Race
Course forest.
Habitat/Ecology. This species is associated withprimary and/or mature secondary forests in
Singapore. Individuals have been found along
forest trails, or among low vegetation.
Remarks. Despite being considered a relatively
common species in Southeast Asia, P. olybria
appears limited to areas of good quality forest
in Singapore.
Polyrhachis (Polyrhachis) ypsilon Emery, 1887
Material examined. None available.
Material not physically examined. Lectotype –CASENT0905548, ANTC26437 (MSNG).
Literature. Type – Emery (1887). Hung (1970)[designation of lectotypes, paralectotypes];
Kohout (1998); Kohout (2014).
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore context.
Remarks. Type locality in Singapore. At the timeof writing, the only record of this species in
Singapore is that of its type series, collectedin 1875 by D’Albertis (Emery 1887).

Prenolepis subopaca Emery, 1900
Material examined. Bukit Timah Nature Reserve, 28 Mar 1967, D.H. Murphy leg., ZRC_
HYM_0000760; Mandai Track 15, 4 Jan
2016, G.W. Yong leg., ZRC_HYM_00010291031; Mandai Track, 4 Jan 2017, G.W. Yongleg., ZRC_HYM_0001168; Mandai Lake
Road, 4 Jan 2016, G.W. Yong leg., ZRC_
HYM_0001032; same locality and collector

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as previous, 5 Nov 2016, G.W. Yong leg.,
ZRC_ENT00047967; Upper Peirce Reservoir, 10 Jan 2014, Sk. Yamane leg. (SKYC);
Upper Thomson Nature Park, 1.38241,
103.81944, 9-11 Oct 2016, G.W. Yong et al.
leg., UT4TPc1672, ZRC_ENT00047920.
Material not physically examined. Bukit Timah,
1°21’N, 103°47’E, 50 m, 7 Apr 1989, P.S.
Ward leg.; Sembawang, 18 Jun 1967, D.H.
Murphy leg. [in Williams & LaPolla 2016].
Literature. Williams & LaPolla (2016); Peeters& Yong (2017).
Localities. Bukit Timah Nature Reserve; Mandai
Lake Road; Mandai Track; Sembawang; Upper Peirce Reservoir; Upper Thomson Nature
Park.
Habitat/Ecology. This species is associatedmostly with mature or old secondary forestsof varying levels of disturbance, including
abandoned plantation forest. Nests can befound in leaf litter; individuals were usuallyfound foraging arboreally.
Remarks. Prenolepis subopaca is morphologically similar to its congener P. jerdoni. The
former can be differentiated from the latterby its finely reticulate and dull (almost matte)
cuticular sculpture, and thick pubescencepresent in patches on pronotal and propodeal
dorsums, and also the gaster.
Pseudolasius badius Viehmeyer, 1916
Material examined. None available.
Material not physically examined. Type queen– FOCOL2677 (ZMHB).
Literature. Type – Viehmeyer (1916); Overbeck(1924).
Localities. Bukit Timah Road
Habitat/Ecology. The type queen was collected
at a lit-lamp in a garden.
Remarks. Type locality in Singapore. This species
was first described based on a single queen
collected in Singapore. At the time of writing,the type queen remains the only known recordof the species in Singapore.

Pseudolasius circularis Viehmeyer, 1916
Material examined. Alate queen, males andworkers, Nee Soon Swamp Forest, 16 Mar
1981, D.H. Murphy leg., DHM-SG81-Psel2,
ZRC_ENT00027952.
Material not physically examined. Type queen– FOCOL2676 (ZMHB).
Literature. Type – Viehmeyer (1916); Overbeck(1924).
Localities. Bukit Timah Road; Nee Soon Swamp
Forest.
Habitat/Ecology. The type queen was collected ata lit-lamp in a garden. More recently, a colony
was found nesting under a log on a raised pathin swamp forest.
Remarks. Type locality in Singapore. This species
was first described based on a single queen
collected in Singapore. The queen can be distinguished from other congeners mainly by
its expansive rounded mesosoma (in dorsalview), mesoscutum much wider than long,propodeum very short but wide with no distinction between lateral and posterior margins,
both of which form a continuous broadly convex outline in dorsal view. Workers exhibit
strong allometric polymorphism, with large
majors having massive sub-cordate heads withdeeply concave posterior margins, disproportionately small eyes and short antennal scapes;
smaller minor workers tend to have smaller
and less expanded heads, with antennal scapes
exceeding the occipital margin.
Pseudolasius ludovici Forel, 1913
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah Road
Habitat/Ecology. A single queen was collected ata lit-lamp in a garden.
Remarks. At the time of writing, there are only
literature records of this species in Singapore.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Pseudolasius martini Forel, 1911
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah Road
Habitat/Ecology. Males were collected at a litlamp in a garden.
Remarks. At the time of writing, there are only
literature records of this species in Singapore.
Pseudolasius mayri duplicatus Viehmeyer, 1916
Material examined. None available
Material not physically examined. Types– males, FOCOL2688-2689; queen, FOCOL2690 (ZMHB).
Literature. Type – Viehmeyer (1916); Overbeck(1924) [both as Pseudolasius mayri var. duplicata].
Localities. Bukit Timah Road
Habitat/Ecology. Type alates were collected at a
lit-lamp in a garden.
Remarks. Type locality in Singapore. This species
was first described based on a queen collectedfrom Singapore.
Pseudolasius trimorphus Karavaiev, 1929
Material examined. Nee Soon forest, 26 Apr
2018, W. Wang leg., WW-SG18-Pseul1,
ZRC_ENT00000949.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Nee Soon forest
Habitat/Ecology. This species was found in Adinandra belukar-dominated secondary forest;a nest was found under the bark of an underground tree root, with soil substrate.
LEPTANILLINAE (2 genera, 3 species)
Leptanilla havilandi Forel, 1901
Material examined. None available.
Material not physically examined. Types —CASENT0101455, ANTC3654 (MNHN);
CASENT0901483, ATC21005 (OUMNH);
CASENT0902773, ANTC22248 (BMNH);
CASENT0907032, ANTC27210 (MHNG);
FOCOL1126 (ZMHB).

Literature. Type – Forel (1901). Emery (1910).
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore
context; probably a hypogaeic subterranean
species as with most other Leptanilla species.
Remarks. Type locality in Singapore. Despite
not having been recorded from Singapore formore than a century since its initial discovery,
Leptanilla havilandi has been firmly established as a valid species discernible from its
congeners within Indomalaya (see: Baroni
Urbani 1977c, Ito et al. 2001, Wong & Guénard 2016). The scarcity of specimens may bepartly attributed to the species’ subterranean
nature and minute size, which render it elusiveand difficult to find.
Leptanilla hypodracos Wong & Guénard, 2016
Material examined. Holotype – Central Catchment Nature Reserve, ca. 55 m, 1.355,
103.815, 15 Jun 2015, M.K.L. Wong leg., subterranean pitfall trap, ZRC_ENT00000664.
Material not physically examined. Paratypes –same collection data as holotype, MW1506151.2, -1.3 (HKUC).
Literature. Wong & Guénard (2016).
Localities. Central Catchment Nature Reserve
Habitat/Ecology. The type series was collectedfrom underground using a baited subterranean
pitfall trap installed in a well-shaded fragmentof old secondary forest.
Remarks. Type locality in Singapore. At the timeof writing, the only record of this species in
Singapore is that of its type series.
Protanilla rafflesi Taylor (1990)
Material examined. Queen and workers, Bukit Timah Nature Reserve, D10-4, 28 Jan 1969, D.H.
Murphy leg., ZRC_HYM_0000023; same locality and collector as previous, 7 Dec 1968,
ZRC_HYM_0000028; same locality as previous, 1.34796, 103.77914, 29 Oct 2015, M.K.L.
Wong leg., ZRC_ENT00000674; Bukit Timah
Hill, slope, D11-7, collection date unknown,
D.H. Murphy leg., ZRC_HYM_0000027;
Lower Peirce Reservoir, Aug 1990, collector unknown, ZRC_HYM_0000244; samecollection data as previous (SKYC); Nee

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Soon forest, 1.38279, 103.80650, 25 Sep
2017, W. Wang leg., NSW1, Winkler extraction, ZRC_ENT00000910; same locality andcollector as previous, 1.38245, 103.80206,
46m, 26 Apr 2018, Winkler extraction, ZRC_
ENT00000953.
Material not physically examined. Holotype – CASENT0102375, ANTC4604(BMNH). Paratype – ANIC32-017901, Shattuck1345248029 (ANIC).
Literature. Type – Taylor in Bolton (1990). Xu(2012).
Localities. Bukit Timah; Bukit Timah Nature Reserve (a.k.a. Bukit Timah Hill); Lower Peirce
Reservoir Road; MacRitchie; Nee Soon forest.
Habitat/Ecology. This species is associated mostly with primary or mature secondary habitatin Singapore, including swamp forest and oldsecondary forest. Specimens have also beencollected from young secondary Adinandra
belukar-dominated forest. Usually found inleaf litter and woody ground debris, often collected via Winkler extraction.
Remarks. Type localities (i.e., MacRitchie andBukit Timah Nature Reserve) in Singapore.
MYRMICINAE (33 genera, 158 species)
Acanthomyrmex ferox Emery, 1893
Material examined. Queens and workers, NeeSoon Swamp Forest, 1.38252, 103.80208,
25 Sep 2017, W. Wang leg., NS_W1, Winkler extraction, WW-SG17-041, ZRC_
ENT00014121; Seletar Trail, 1.39192,
103.80081, 18 Oct 2015, M.K.L. Wong leg.,
Berlese extraction, ZRC_ENT00000683.
Material not physically examined. FMNHINS0002821982-2821990, FMHD_81-282(FMNH).
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Nee Soon
Swamp Forest; Seletar Trail.
Habitat/Ecology. This species is associated withprimary and/or mature secondary forests in
Singapore, including swamp forest. Individuals have been found in leaf or (tree) buttresslitter, also on the forest floor.

Aphaenogaster simulans Forel, 1915 stat. n.
Aphaenogaster (Deromyrma) feae r. simulans
Forel, 1915: 31 (w.q.m.)
Subspecies of Aphaenogaster feae: Emery, 1921:
65; Chapman & Capco, 1951: 133; Bolton,
1995: 73.
Material examined. Bukit Timah Nature Reserve,near BT01, 1.35764, 103.77491, 1 Feb 2017,
W. Wang leg., ZRC_ENT00007570; Mandai
Road, plot 1, 1.38252, 103.80208, 16 Nov
2016, G.W. Yong et al. leg., M1D4GH2159,
ZRC_ENT00047838; Nee Soon Swamp Forest, 1.38252, 103.80208, 25 Sep 2017, W.
Wang leg., NS_W1, leaf litter, Winkler extraction, ZRC_ENT00047839; same locality
as previous, Woodcutter’s trail, 4 Sep 2017,
W. Wang leg., ZRC_ENT00000985.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Mandai;
Nee Soon Swamp Forest.
Habitat/Ecology. This species is associated mostly with primary or mature secondary forests
in Singapore, including swamp forest, where
workers are usually found foraging on leaf
litter or in Nepenthes pitcher contents. At thetime of writing, a single occurrence of the species had also been recorded at an abandonedpark - young secondary forest.
Remarks. Originally described as a subspecies of
Aphaenogaster feae Emery, 1889, A. simulans
can be distinguished from the former basedon the following (A. feae characters in parentheses): 1) propodeal spines/denticles very
short and blunt, barely protruding from propodeal junction (propodeal spines relatively
longer, acute, distinctly protruding from propodeal junction), 2) petiolar node in dorsal
view roughly as long as wide, broadly ovate(petiolar node in dorsal view distinctly longer
than wide, elongate-ovate), 3) head in full faceview, excluding the narrowed neck, ca. 1.5
times longer than wide, area posterior to eyes
somewhat conical or like an inverted-cone(head in full face view, excluding the neck,only slightly longer than wide (< 1.5 times),
area posterior to eyes semi-circular).
		We believe that these observed differences, some of which were also mentioned inoriginal descriptions for both species, justify
the raising of simulans to species.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Calyptomyrmex beccarii Emery, 1893
Material examined. Bukit Timah Nature Reserve,
Jungle Fall Valley, 10 Apr 1976, D.H. Murphy leg., ZRC_HYM_0000623-624; Kent
Ridge Park, YM site 2, 7 Dec 2017, W. Wangleg., ZRC_ENT00027983; Mandai, 1.40699,
103.77853, 19 Sep 2015, M.K.L. Wong leg.,
Berlese extraction, ZRC_ENT00000684;
Mandai Lake Road, 3 Nov 2016, G.W. Yongleg., ZRC_HYM_0000990; same locality,
collection date and collector as previous,
ZRC_HYM_0001737.
Material not physically examined. FOCOL2181-2182 (ZMHB) [types of Calyptomyrmex beccarii var. glabratus].
Literature. Viehmeyer (1916), Overbeck (1924)[both as C. beccarii var. glabratus]; Shattuck(2011).
Localities. Bukit Timah Nature Reserve; Bukit
Timah Road; Kent Ridge Park; Mandai; Mandai Lake Road.
Habitat/Ecology. This species can be found mostly in young or old/mature secondary habitats
in Singapore, including waste woodlands anddisturbed secondary forest fragments in urban or semi-urban settings. A few occurrenceswere also recorded from cultivated vegetationsuch as gardens. Individuals have been collected from leaf litter, sometimes soil and root
substrate matrices at bases of trees.
Calyptomyrmex loweryi Shattuck, 2011Material examined. Bukit Batok East, 29 Sep
2016, G.W. Yong leg., ZRC_HYM_0000991.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Batok East
Habitat/Ecology. A single specimen was foundin mature or young secondary forest in Singapore. This species has only been found in
one locality to date.
Remarks. Calyptomyrmex loweryi is very similar
to the more widespread C. beccarii, but canbe distinguished from the latter by its relatively larger size, wider head (HW generally> 1.13 mm, see Shattuck 2011; 1.15 mm in
the Singapore specimen) and broader posterior margin of head. However, we noticed

some larger specimens of C. beccarii withmaximum head width slightly more than 1.10mm, unlike what was diagnosed in Shattuck(2011). These larger workers were from thesame colony as their smaller nestmates, whose
measurements better fulfilled the diagnosis for
C. beccarii. Thus, size should be used withcaution when differentiating between C. loweryi and beccarii.
Cardiocondyla kagutsuchi Terayama, 1999
Material examined. None available.
Material not physically examined. Unknown.
Literature. Seifert (2003).
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore context.
Remarks. At the time of writing, there is only asingle literature record of this species in Singapore.
Cardiocondyla obscurior Wheeler, 1929
Material examined. Icube, National Universityof Singapore (NUS), 1.29347, 103.77633, 29
Jul-5 Aug 2015, M.S. Foo & W. Wang leg.,malaise trap, NUS00074, ZRC_BDP0042671;
Prince George’s Park Residences (NUS),
1.29239, 103.77869, 24 Mar-1 Apr 2015, M.S.
Foo & W. Wang leg., malaise trap, NUS0001,
ZRC_BDP0044284, ZRC_BDP0044334;
same locality and collectors as previous, 29
Apr-6 May 2015, malaise trap, NUS0021,
ZRC_BDP0044156; University Hall (NUS),
1.297111, 103.77658, 1-8 Apr 2015, M.S.
Foo & W. Wang leg., malaise trap, NUS0008,
ZRC_BDP0045349; University Town (NUS),
1.30622, 103.77458, 5-12 Aug 2015, M.S.
Foo & W. Wang leg., malaise trap, NUS0079,
ZRC_BDP0042640.
Material not physically examined. Unknown.
Literature. Wang et al. (2018a).
Localities. National University of Singapore campus (Clementi/Kent Ridge).
Habitat/Ecology. This is an accomplished tramp
species presumably native to Southeast Asia,
mainly associated with disturbed secondary
forest fragments in urban or semi-urban settings in Singapore; it can also be found in

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urban cultivated grass- or parklands. While
the species is known to nest in dead twigs or
branch cavities of bushes/trees/shrubs, formal
records of local occurrences comprise mostly
alates collected via malaise traps. Like its
close relative (similarly also a tramp species)
C. wroughtonii, C. obscurior nests in vegetation above the surface rather than in soil.
Polygynous colonies with multiple queens are
known to be common.
Remarks. Given the morphological similarities
with the congener C. wroughtonii, which has
been recorded from Singapore since 1892, itis possible that at least some historical recordsof C. wroughtonii were actually misidentified
C. obscurior.
Cardicondyla strigifrons Viehmeyer, 1922
Material examined. None available.
Material not physically examined. In Seifert
et al. (2017): Singapore, 1.31, 103.83, Nest
sample collected pre-1920, H. Overbeck leg.
Literature. Seifert et al. (2017).
Localities. Bukit Timah Road
Habitat/Ecology. Unknown in the Singapore context.
Remarks. The name of the locality where this species was collected was roughly inferred fromcoordinates provided in Seifert et al. (2017).
Cardiocondyla tjibodana Karavaiev, 1935
Material examined. Bukit Timah Nature Reserve,
13 Dec 1995, H.K. Lua et al. leg., NS 212C,
ZRC_ENT00000347; Upper Peirce Reservoir,
10 Jan 2014, Sk. Yamane leg. (SKYC).
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Upper
Peirce Reservoir.
Habitat/Ecology. This species was found in either primary or mature secondary forests in
Singapore.
Remarks. In other countries, this species is known
to nest in shallow soil in more open disturbed
habitat with weakly herbaceous or even baregrounds. It is therefore possible that there are
more undocumented occurrences of the species in more exposed and disturbed habitats
locally.

Cardiocondyla wroughtonii (Forel, 1890)
Material examined. Kranji Road, 1.42638,
103.75413, 8 Nov 2016, G.W. Yong leg.,
KR1TH1392, ZRC_HYM_0001773; same
locality and collector as previous,7 Sep 2016,
ZRC_ENT00048403; Mandai Road, 1.4119,
103.80513, 29 Nov 2016, G.W. Yong leg.,
M3D4GH850, ZRC_HYM_0001772; Upper
Peirce Reservoir, 10 Jan 2014, Sk. Yamaneleg. (SKYC).
Material not physically examined.
CASENT0908352, ANTC29539 (MHNG)[syntype of C.wroughtonii quadraticeps];
FOCOL1612 (ZMHB) [type queen of
C.wroughtonii quadraticeps].
Literature. Forel (1912), Viehmeyer (1916),
Overbeck (1924) [all as C. wroughtonii quadraticeps]. Seifert (2003).
Localities. Bukit Timah Road; Kranji Road; Mandai Road; Upper Peirce Reservoir.
Habitat/Ecology. This species was found in bothmature native-dominated, and young waste
woodland secondary forests in Singapore, also
in gardens. Some specimens were collected
on tree trunks.
Remarks. Cardiocondyla wroughtonii is a recognized global tramp species that can thrive in arange of habitats including degraded vegetation in urban or semi-urban settings. Thus, it ishighly possible that actual species distributionin Singapore might be broader, and underrepresented by material available for examination
at the time of writing.
Cardiocondyla sp.4.of.SKY (*excluded from official named species count)Material examined. Queens and workers,
Central Catchment Nature Reserve, 4
Sep 2017, W. Wang leg., WW-SG17-020,
ZRC_HYM_0001757; queens and workers, Mandai forest, 15 Jan 2020, W. Wangleg., WW-SG20-001, ZRC_ENT00014129;
Kranji Road, 7 Sep 2016, G.W. Yong leg.,
ZRC_ENT00048406.
Localities. Central Catchment Nature Reserve;
Kranji Road; Mandai.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Habitat/Ecology. This species appears to be associated mostly with mature and/or old secondary forests. Nests were found in cavities
in bark of large fallen trees, or in fallen woodon ground.
Remarks. Though currently undescribed, thisunique species is given special mention here
as it appears quite common not just in Singapore, but in Borneo and throughout South
Thailand (Sk. Yamane 2020, pers. comm.).
Body generally uniformly dark brown, almost
entirely coarsely rugose-reticulate except for
the gaster which is smooth and shining. This
species appears affiliated to Cardiocondyla
thoracica (Smith, 1859), or is at least in thesame species group, based on similar features
such as: 1) anterior margin of pronotal discslightly elevated and projecting above the otherwise (near) continuous convex outline of themesosomal dorsum in profile, 2) metanotal
groove indistinct or almost absent, and 3) very
long propodeal spines with apices directed
posteriorly.
Carebara affinis (Jerdon, 1851)
Material examined. Lower Peirce Reservoir,
Forest A, 7 Aug 1990, Zoology Dept 3rd Yearproject students, ZRC_ENT00000192; same
locality as previous, Forest B, 25 Aug 1990,
Zoology Dept 3rd Year project students, ZRC_
ENT00000113; Lorong Banir, NS 192, 16 Jun
1995, Yang & Lua leg., ZRC_ENT00000376;
Bukit Timah, Hindhede Drive, 25 Dec 1989,
H.K. Lua leg., ZRC_HYM_00000130; Pulau Ubin, 7 Jan 2014, Sk. Yamane leg.,
ZRC_HYM_00000434-435, 482-483; University Campus Botany Garden, 11 Feb 1977,
D.H. Murphy leg., ZRC_HYM_0000913;
Mandai Road, 1.41119, 103.80513, 29 Nov1 Dec 2016, G.W. Yong leg., pitfall trap,
M3D3GPc806, ZRC_HYM_0001770; Bukit
Batok East Avenue 6, 1.34304, 103.76235,
3-5 Oct 2016, G.W. Yong leg., pitfall trap,
BB1GPd382, ZRC_HYM_0001771; Mandai forest, 7 Jun 2018, W. Wang leg., ZRC_
ENT00047996; male, University Town
(NUS), 1.30622, 103.77458, 29 Apr-6 May
2015, M.S. Foo & W. Wang leg., NUS0023,
malaise trap, ZRC_BDP0044771.

Material not physically examined. Unknown.
Literature. Viehmeyer (1916). Overbeck (1924),
Yamane (2003) [all as Pheidologeton affinis].
Wang et al. (2018a).
Localities. Bukit Batok East; Bukit Timah Hindhede Drive; Bukit Timah Road; Lorong Banir;
Lower Peirce Reservoir; Mandai; University
Town (NUS).
Habitat/Ecology. This species was found in bothmature and young secondary forests at varying levels of disturbances, including nativedominated and abandoned plantation secondary forests. Individuals have been collectedoccasionally in disturbed forest fragments insemi-urban settings, or in cultivated gardens.
The species has sometimes been collectedfrom leaf litter or decayed wood.
Remarks. Considerable morphological variation
was observed among different colonies of C.
affinis, but we lack convincing empirical evidence supporting further species delimitation
for this possible species complex at the time ofwriting. Thus, in this checklist, we tentatively
treat all morphological variants as conspecific.
Carebara castanea Smith, 1858
Material examined. Alate queen, Pulau Ubin,
1.41149, 103.97475, 22 Oct 2015, S.X. Chuiet al. leg., ZRC_ENT00047836; alate queen,
Kent Ridge Road, 1.29518, 103.77615, 2 Nov
2015, Y.G. Tan leg., ZRC_ENT00047837.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Kent Ridge Road; Pulau Ubin.
Habitat/Ecology. Alate queens were found, sometimes en masse, in secondary forest patches orfragments in semi-urban settings.
Remarks. At the time of writing, the only records of this species in Singapore are ofalate queens. The alate queen of C. castanea
closely resembles that of Carebara lignata
Westwood, 1840, however, its post-petiole
appears broadest (in dorsal view) anteriorly,
unlike in C. lignata.

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Carebara diversa (Jerdon, 1851)
Material examined. Bukit Timah Hindhede
Drive, 19 Nov 1989, H.K. Lua leg., ZRC_
HYM_0000158; Bukit Timah Nature Reserve,
13 Dec 1995, H.K. Lua et al. leg., NS 212C,
ZRC_ENT00000348; same locality as previous, 12 Oct 2015, D.J. Court leg., pitfall trap,
ZRC_ENT00048442; Commonwealth Avenue West, 6 Dec 2005, H.K. Lua leg., LHK457, ZRC_HYM_0000039-40; alate queens,
Holland Close (in HDB flat), 14 Apr 1990,
K. Lim leg., ZRC_HYM_0000201; National University of Singapore (Clementi/Kent
Ridge campus), Apr-Sep 2015, M.S. Foo &W. Wang leg., malaise trap, ZRC_BDP (multiple); Labrador beach, 29 Apr 1985, D.H.
Murphy leg., ZRC_ENT00027996; Lower
Peirce Reservoir, Forest A, 25 Aug 1990, collector unknown, ZRC_ENT0000137; Mandai,
Northern Node, 4 Sep 2019, J. Tan & N. Chinleg., large fallen rotting branch, MIS_NN_AC,
ZRC_ENT00027942; Mandai Lake Road,
Rainforest Park North, Buffer area, 1.40842,
103.78319, 54 m, 4 Sep 2019, J. Tan & N.
Chin leg., leaf litter, ZRC_ENT00013862;
Nee Soon Swamp Forest, 16 Apr 1992,
D.H. Murphy et al. leg., Y790, ZRC_
HYM_00000037; same locality as previous,
5 Apr 2005, P. Grootaert leg., malaise trap,
PGT 25046-25048, ZRC_HYM_0000038;
Neo Tiew Lane, 16 Jul 2009, H.K. Lua leg.,
LHK 513, ZRC_HYM_0000035; Pulau Ubin
(Outward Bound School grounds), 1.41662,
103.99443, 4 Dec 2016, G.W. Yong & M.K.L.
Wong leg., ZRC_HYM_0001051-1054; Seletar Link,1.40518, 103.88555, 16-18 Oct 2016,
G.W. Yong leg., pitfall trap, SL3GPd1195,
ZRC_HYM_0001769; Seletar Reservoir
Park, 12 May 1994, collector unknown, NS
132C, ZRC_HYM_0000108.
Material not physically examined. Non-types,
CASENT0195766, PSW09562 (PSWC).
Non-types, J.K. Wetterer leg. (JKWC) - Amber Road (Katong), 28 Jul 2014, vial #348;
Dhoby Ghaut, Istana Park, 30 Jul 2014, vial
#403; Harbourfront, 29 Jul 2014, vial #385;
Katong Park, 1 Aug 2014, vial #448; Outram
Park, 29 Jul 2014, vial #381; Punggol Point,
30 Jul 2014, vial #395; Singapore Botanic
Gardens, 5 Aug 2014, vial #475.

Literature. Viehmeyer (1916), Overbeck (1924),
Moffett (1987, 1988), Yamane (2003), Tan &Corlett (2012) [all as Pheidologeton diversus].
Wang et al. (2018a).
Localities. Bukit Timah Nature Reserve; Commonwealth Avenue West; Dhoby Ghaut; Harbourfront; Holland Close; Katong Park; Labrador beach; Lower Peirce Reservoir; Mandai;
Mandai Lake Road; National University ofSingapore campus; Nee Soon Swamp Forest;
Neo Tiew Lane; Outram Park; Pulau Ubin;
Punggol Point; Seletar Link; Seletar Reservoir
Park; Singapore Botanic Gardens.
Habitat/Ecology. This species was mainly foundat edges of mature/old secondary forest, in
disturbed waste woodland secondary forest
fragments, or swathes of cultivated vegetation such as gardens, in urban or semi-urban
settings. Colonies often form rather pervasive trunk trails on the ground (including
across human-made pavements), tree trunks,
and over foliage. These trunk trails are partof the species’ unique group hunting/foraging or swarm raiding strategies. These ants
are known to be extreme trophic generalists,
consuming a wide range of material including
animal carcasses, fruits, nuts, bark etc. Alatesof C. diversa frequently occur in abundanceat urban residential areas following a periodof prolonged heavy rain and humid weather.
Carebara overbecki (Viehmeyer, 1916)
Material examined. Pulau Ubin, 22 Oct 2016,
W. Wang leg., leaf litter, Winkler extraction,
ZRC_ENT00000988.
Material not physically examined. Types –CASENT0908912, ANTC31343 (MHNG);
FOCOL0278-279 (DEIC, Muncheberg); 3
males, 2 workers, FOCOL1918-1922 (ZMHB).
Literature. Type – Viehmeyer (1916); Overbeck(1924) [as Oligomyrmex overbecki]. Moffett(1986) [as Oligomyrmex overbecki].
Localities. Bukit Timah Road; Pulau Ubin.
Habitat/Ecology. This minute species is associated mostly with mature secondary forests in
Singapore. The ants can be found in leaf litteror under bark. Nests have been found in large
aged fallen tree trunks, also in root cavities ina soil mound, just next to a termite nest.
Remarks. Type locality in Singapore.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Carebara silenus (Smith, 1858)
Material examined. Male, University Hall (NUS),
1.29711, 103.77658, 6-13 May 2015, M.S.
Foo & W. Wang leg., malaise trap, NUS0028,
ZRC_BDP0041385.
Material not physically examined. Syntypes– CASENT0900737-900738, ANTC19789(BMNH)[types of Pheidologeton silenus].
Literature. Type – Smith (1858) [as Pheidole silenus]. Moffett (1988a), Yamane (2003) [bothas Pheidologeton silenus].
Localities. University Hall (NUS)
Habitat/Ecology. Similar foraging and/or hunting
tactics as with Carebara diversa, but lacking
the stable trunk trails unique to the latter (Moffett 1988a). There is no available informationon the habitat in which the type series was
obtained. This species may likely be found inmature or old secondary forest fragments, inurban or semi-urban settings, where it sometimes appears to thrive in sympatry with C.
diversa.
Remarks. Type locality in Singapore.
Cataulacus granulatus (Latreille, 1802)
Material examined. Bukit Batok, 28 Jan 2015, W.
Wang leg., ZRC_ENT00027963; Bukit Batok East Avenue 6, 1.34304, 103.76235, BB6GH076, ZRC_HYM_0001202.2; same locality and collector as previous, 30 Aug 2016,
G.W. Yong leg., ZRC_HYM_0001208.21209.2; Bukit Timah forest, 14 Aug
1968, D.H. Murphy leg., A124-4, ZRC_
HYM_0000691; Kent Ridge, 14 Apr 1985,
D.H. Murphy leg., ZRC_HYM_0000689;
Kent Ridge scrub, 18 May 1985, D.H. Murphy leg., ZRC_HYM_0000692; Lower Peirce
Reservoir, 21 Jul 1990, H.K. Lua leg., ZRC_
HYM_0000232; Sungei Buloh, 8 Jan 2013,
Sk. Yamane leg., ZRC_HYM_0000413; Pulau Sentosa, 2 Dec 1989, collector unknown(SKYC).
Material not physically examined.
CASENT0901453, ANTC20975 (OUMNH)[holotype of Cataulacus hispidus Smith,
1876, junior synonym of C. granulatus];
CASENT0922493, PSW09575-5 (PSWC).

Literature. Smith (1876) [as type of C. hispidus].
Forel (1912) [as C. hispidus]. Forel (1911),
Viehmeyer (1916), Overbeck (1924) [all as
C. granulatus subsp. hispidus]. Bolton (1974).
Localities. Bukit Batok; Bukit Timah; Bukit Timah Road; Kent Ridge; Lower Peirce Reservoir; Pulau Sentosa (now simply known as‘Sentosa’); Sungei Buloh; Thompson Road.
Habitat/Ecology. This species can be found in a
broad range of habitat types, including urban
or semi-urban scrub, waste woodland forest(abandoned park), mangrove back forest, andabandoned plantation (young) secondary forest. These ants are considered arboreal; specimens are often collected from low vegetationor foliage, sometimes on tree trunks or low
branches.
Remarks. The first local record of this species wasas the holotype of C. hispidus.
Cataulacus horridus (Smith, 1857)
Material examined. Bukit Timah forest, collection date unknown, D.H. Murphy leg., ZRC_
HYM_0000688.
Material not physically examined.
CASENT0217825, ANTC18013 (CASC);CASENT0922498, PSW09575-4 (PSWC).
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah forest
Habitat/Ecology. This arboreal species can befound in primary or mature/old forests in
Singapore. Individuals were collected fromlow vegetation, or on bark of trees.
Remarks. Examined ZRC material was originallyidentified as Cataulacus insularis Smith, 1857
– now a junior synonym of C. horridus.
Cataulacus latissimus (Emery, 1893)
Material examined. Central Catchment Nature
Reserve, Bukit Kalang service road, 3 Dec
2004, T.M. Leong leg., ZRC_ENT00048384;
Seletar Trail, 1.39532, 103.80228, 5 Mar 2016,
M.K.L. Wong leg., ZRC_ENT00000685.
Material not physically examined. Unknown.
Literature. Bolton (1974).
Localities. Central Catchment Nature Reserve;
Seletar Trail.

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Habitat/Ecology. This species was collected fromthe edges of primary or secondary forest, in
foliage along trails or roads.
Cataulacus praetextus (Smith, 1867)
Material examined. Mandai Road, plot 1, 1.41312,
103.80185, 6-8 Dec 2016, G.W. Yong et al.
leg., M1D18GPb2421, ZRC_ENT00048642;
Nee Soon Swamp Forest, NS1, 1°23’00.3”N,
103°48’46.5”E, 28 Mar-4 Apr 2012, J. Puniamoorthy et al. leg., malaise trap, Reg. 29157,
ZRC_BDP0014626.
Material not physically examined. Alate queen,
FOCOL2122 (ZMHB) [type queen of Cataulacus praetextus var. sumatrensis – now juniorsynonym of C. praetextus].
Literature. Viehmeyer (1916), Overbeck (1924)[both as C. praetextus var. sumatrensis].
Localities. Jurong Road; Mandai Road; Nee Soon
Swamp Forest.
Habitat/Ecology. Workers of this species were
found together with Tetramorium obtusidens
Viehmeyer, 1916 and Technomyrmex albipes(Smith, 1861) in the same withered branchof a roadside tree; workers have also beencollected from Nepenthes pitcher cups. The
species appears mostly associated with old/
mature secondary forests in Singapore, including swamp forest.
Crematogaster anthracina Smith, 1857
Material examined. Mandai mangroves,
4 Oct 1979, D.H. Murphy leg., ZRC_
HYM_0000699.
Material not physically examined. Syntype –CASENT0901434, ANTC20957 (OUMNH).
Non-type – CASENT0914071, ANTC27752(BMNH).
Literature. Type – Smith (1857) [as Crematogaster anthracinus].
Localities. Mandai mangroves
Habitat/Ecology. No habitat information is available for the type series. Non-type material ofthis species was collected from mangroves,where the ants were found nesting in a twig
of Sonneratia sp.
Remarks. Type locality in Singapore.

Crematogaster aurita Karavaiev, 1935
Material examined. Jalan Ulu Sembawang,
1.417, 103.769, Jul 2015, J.K.N. Tan leg.,
C190, JKT-SG15-001, ZRC_ENT00013939.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Jalan Ulu Sembawang
Habitat/Ecology. Uncertain in the Singapore context, but this species was possibly found inyoung secondary forest, as inferred from thereported locality.
Crematogaster borneensis André, 1896
Material examined. None available.
Material not physically examined.
CASENT0193115, BBB111 (BBBC).
Literature. Viehmeyer (1916), Overbeck (1924)[both as Crematogaster (Decacrema)borneensis macarangae].
Localities. Bukit Timah Hill; Central Catchment,
Sime Road.
Habitat/Ecology. This species is obligately associated with Macaranga plants, specifically hostspecies belonging to the sections Pachystemonand Pruinosae, which are commonly found insecondary forests in general. The ant-plant associations are not strictly species-specific, andC. borneensis is known to inhabit/colonize twoto seven different Macaranga host species overits entire distributional range. In Singapore,
the ants were found in the hollow internodesof shafts of Macaranga hypoleuca (Rchb.f. &Zoll.) Müll.Arg., and also with scale insects in
Macaranga bancana (Miq.) Müll.Arg.
Crematogaster brunnea ruginota Santschi, 1928
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[both as Crematogaster subnuda politula var.
ruginota].
Localities. Bukit Timah Road
Habitat/Ecology. Individuals were found in thestump of a branch on a living tree, in a garden.
Remarks. At the time of writing, there are only
literature records of this subspecies in Singapore.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Crematogaster coriaria Mayr, 1872
Material examined. Bukit Timah Nature Reserve, near BT01, 7 Jun 2017, W. Wang leg.,
WW-SG17-029, ZRC_ENT00000983; same
locality as previous, near BT02, 1.35599,
103.77397, 12 Jul 2017, W. Wang leg., WWSG17-016, ZRC_HYM_0001736; Bukit
Peirce, 10 May 2005, Sk. Yamane leg., nest
in dead twig on ground (SKYC).
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Peirce; Bukit Timah Nature Reserve.
Habitat/Ecology. This species is associated withprimary and old secondary forests in Singapore, and appears arboreal. Nests have beenfound in trunks of living trees (e.g. Lithocarpus sp.), and also in cavities of fallen branchesor twigs in old secondary forest.
Crematogaster decamera Forel, 1910
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah Road
Habitat/Ecology. A single queen was collectedfrom a lit-lamp in a garden.
Remarks. At the time of writing, there are only
literature records of this species in Singapore.
Crematogaster difformis Smith, 1857
Material examined. Upper Peirce Reservoir, 10 Jan 2014, Sk. Yamane leg., ZRC_
HYM_0000365-366.
Material not physically examined. Lectotype andparalectotypes (OUMNH).
Literature. Type – Smith (1857). Viehmeyer(1916), Overbeck (1924) [both misspelt as‘C. deformis’]; Hosoishi & Ogata (2009).
Localities. Ayer Terjun (archaic place name);
Bukit Timah Hill; Choa Chu Kang Road; Upper Peirce Reservoir.
Habitat/Ecology. This species was found in primary or mature secondary forests in Singapore.

Remarks. Type locality in Singapore. Very similar
to the more common C. sewardi, distinguishedmainly by the configuration or direction of
appressed hairs/setae on the first gastral tergite - in C. sewardi, the gastral hairs are allposteriorly-directed, whereas in C. difformis,
hairs in the posterior section (of the tergite)
are directed medially inwards towards thecentre of the tergite.
Crematogaster dohrni artifex Mayr, 1878 [erroneously printed as 1879 in AntCat and
AntWeb]Material examined. None available.
Material not physically examined. Syntype –CASENT0919683, ANTC41462 (NHMW)[type of C. artifex].
Literature. Type – Mayr (1878) [as C. artifex].
Viehmeyer (1916), Overbeck (1924) [both as
C. artifex].
Localities. Changi (type locality); Jurong Road.
Habitat/Ecology. The ants were found in Nepenthes pitcher cups, presumably in mature or old
secondary forest. Colonies were also foundhoused in fairly large pasteboard nests aroundbranches of trees and shrubs, with smaller satellite nests of similar substrate on neighbouring leaves sheltering scale insects (Coccidae).
Remarks. Type locality in Singapore. Crematogaster dohrni artifex was first described as C.
artifex Mayr, 1878; status as subspecies of C.
dohrni was first suggested by Forel (1902), but
only formally established by Emery (1922).
Crematogaster egidyi szaboi Forel, 1913
Material examined. None available.
Material not physically examined. Syntype –CASENT0908577, ANTC30926 (MHNG)
Literature. Type – Forel (1913).
Localities. Unknown.
Habitat/Ecology. Unknown in Singapore context.
Remarks. Type locality in Singapore. At the timeof writing, this subspecies is known only fromSingapore. While we lack more recent physical material for objective comparisons, basedon images on AntWeb, we observed the following differences between the types of C.
egidyi szaboi and C. egidyi (in parentheses):

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1) head in full face view wider than long (headin full face view either as long as wide orslightly longer than wide), 2) antennal scapemostly with dense appressed or decumbent
short hairs (scape with numerous erect orsuberect short hairs). Considering that theseapparent differences may be symptomatic of
variable imaging perspectives or conditions,
we refrain from making any definite taxonomic conclusions in this paper based on our
limited observations, i.e., maintain the subspecies szaboi.
Crematogaster ferrarii Emery, 1888
Material examined. Temasek Polytechnic - adjacent waste woodland, 1.34239, 103.93598, 13
Dec 2016, G.W. Yong leg., TT2GH587, ZRC_
HYM_0001767; Pulau Semakau Old Fragment, SMO2, 1°12’20.2”N, 103°45’35.8”E,
4-11 Oct 2012, J. Puniamoorthy et al. leg.,mangroves, malaise trap, Reg. 29512, ZRC_
BDP0016526; same locality and collectors asprevious, 27 Sep-4 Oct 2012, mangroves, malaise trap, Reg. 29499, ZRC_BDP0016412,-16414.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Chua [sic] Chu Kang Road; Pulau
Semakau Old Fragment; (beside) Temasek
Polytechnic.
Habitat/Ecology. This species was found in waste
woodland secondary forest, and also in oldgrowth mangroves; in the latter, specimens
were collected in malaise traps. Colonies havebeen found in withered bamboo.
Crematogaster inflata Smith, 1857
Material examined. Bukit Timah Forest,
20 Dec 1969, D.H. Murphy leg., ZRC_
HYM_0000595.
Material not physically examined. Lectotype andparalectotype workers (OUMNH) – examinedby Hosoishi & Ogata (2009).
Literature. Type – Smith (1857) [as Crematogaster inflatus]. Viehmeyer (1916), Overbeck(1924) [both as C. inflatus]; Hosoishi & Ogata(2009).

Localities. Bukit Timah Forest; Chua [sic] Chu
Kang Road.
Habitat/Ecology. This species is associated withprimary and mature or old secondary forests in
Singapore. Nests were observed to be made of‘earth’ or ‘pasteboard’, typically on branches
high up on (fallen) trees or in the high canopy.
The ants were typically collected from fallentrees or large branches. Sometimes nests appeared to be associated with scale insects(Coccidae) in fissures of tree bark, and underneath loose bark.
Remarks. Type locality in Singapore.
Crematogaster linsenmairi Feldhaar, Maschwitz& Fiala, 2016
Material examined. None available.
Material not physically examined. Types – FOCOL1645-1647 (ZMHB) [original syntypesof Crematogaster borneensis hosei tubuli (seeViehmeyer 1916)]
Literature. Type – Feldhaar et al. (2016). Viehmeyer (1916), Overbeck (1924) [both as C.
borneensis hosei var. tubuli].
Localities. Mandai Road
Habitat/Ecology. A member of the Macarangaassociated Crematogaster captiosa-subgroup(which also includes C. borneensis), this species colonizes mainly Macaranga hosts of thesection Pruinosae, more rarely colonizing thesection Pachystemon when the preferred hostspecies are absent (Feldhaar et al. 2016). Nests
have been reported to be found in hollow
shafts at the end (tips) of branches of large
Macaranga shrubs in Singapore.
Remarks. Type locality in Singapore. At the timeof writing, this species is known to only occur
in a single locality in Singapore.
Crematogaster modiglianii Emery, 1900
Material examined. Island Club forest, 20
May 1985, D.H. Murphy leg., ZRC_
HYM_0000709; Central Catchment Reservoir(sic), 10 May 2006, Sk. Yamane leg. (SKYC).
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Central Catchment Nature Reserve(CCNR); Island Club forest [part of CCNR].

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Habitat/Ecology. A colony of this species wasfound in a carton nest with Polyrhachis sp.,in primary or mature secondary forest. The
ants were also found nesting in hanging deadtwigs in similar forest.
Remarks. Previously reported literature recordsof the species in Singapore (i.e., Viehmeyer
1916, Overbeck 1924) actually refer to specimens from ‘St Barbe Island’, part of the Riau
Island Archipelago in Indonesia, and not in
Singapore. Thus, at the time of writing, this is
the only known physical record of the species
in Singapore.
Crematogaster overbecki Viehmeyer, 1916
Material examined. None available.
Material not physically examined. Types –CASENT0912734, ANTC36291 (NHMB, Basel); queen, male and worker, FOCOL0272-274(DEIC, Muncheberg); queen, male and workers, FOCOL1634-1637 (ZMHB).
Literature. Type – Viehmeyer (1916); Overbeck(1924).
Localities. Bukit Timah Road
Habitat/Ecology. A colony of this species wasfound in a dead branch of a tree in a garden,“honeycombed” by termites and beetle larvae.
Remarks. Type locality in Singapore. The only
records of this species in Singapore are thatof the type series.
Crematogaster rogenhoferi Mayr, 1879
Material examined. Temasek Polytechnic, adjacent waste woodland, 1.34239, 103.93598,
13 Dec 2016, G.W. Yong leg., TT2GH586,
ZRC_HYM_0001722.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah Road; (beside) Temasek
Polytechnic.
Habitat/Ecology. This species was found in a
waste woodland secondary forest fragment
in urban settings, also in a garden, where individuals were collected either in Nepenthes
pitcher cups, or on trees where they wereobserved “visiting” scale insects (Coccidae).

Crematogaster sewardi Forel, 1901
Material examined. Admiralty, 4 Feb 2015, W.
Wang leg., ZRC_ENT00048400; Bukit Timah
Nature Reserve, NS 202, 4 Dec 1995, H.K.
Lua et al. leg., ZRC_ENT00000357; same
locality as previous, 5 Feb 1983, D.H. Murphyleg., ZRC_HYM_0000705-707; Commonwealth Road, 1.31115, 103.78171, 24 Nov
2016, G.W. Yong leg., CW1TH633, ZRC_
HYM_0001762; Kent Ridge, 28 May 1985,
D.H. Murphy leg., ZRC_HYM_0000701;
Lower Peirce Reservoir, 21 Jul 1990, H.K.
Lua leg., ZRC_HYM_0000139; Mandai
forest, 15 Jan 2020, W. Wang leg., foraging
trail on tree trunk, WW-SG20-007, ZRC_
ENT00047968; National University of Singapore campus (Clementi/Kent Ridge), AprSep 2015, M.S. Foo & W. Wang leg., malaise
trap, ZRC_BDP (multiple); Nee Soon forest,
16 Aug 2017, W. Wang leg., WW-SG17-065,
ZRC_ENT00047969; Pulau Ubin, 7 Jan 2014,
Sk. Yamane leg., ZRC_HYM_0000361-363;
same locality as previous, PU3, 1-8 Sep 2012,
J. Puniamoorthy et al. leg., mangroves, malaise trap, Reg. 29451, ZRC_BDP001644716448; Sungei Buloh, Feb 2015, R.Z.B. Quekleg., mangroves, ZRC_ENT00007861; Upper
Peirce Reservoir, 10 Jan 2014, Sk. Yamaneleg., ZRC_HYM_0000360.
Material not physically examined. Locality unknown, 4 Dec 1995, Sk. Yamane leg. (SKYC);
locality unknown, 23 Jul 1996, Sk. Yamaneleg. (SKYC) [examined in Hosoishi & Ogata
2008, 2009].
Literature. Hosoishi & Ogata (2008, 2009); Wanget al. (2018a).
Localities. Bukit Timah Nature Reserve; Commonwealth Road; Kent Ridge; Lower Peirce
Reservoir; Mandai; National University ofSingapore campus (Clementi/Kent Ridge);
Pulau Ubin; Sungei Buloh; Upper Peirce
Reservoir.
Habitat/Ecology. This species was often foundin disturbed secondary forest fragments including waste woodland and abandoned park
forests, and also cultivated park areas withinsemi-urban settings or near urban dwellings.
Sometimes these ants were found in mangroves, typically on tree branches or tree
trunks. Nests have been found in tree holes.

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Remarks. As C. sewardi is morphologically very
similar to C. difformis, it is possible thatthe latter species as identified in Viehmeyer(1916) and Overbeck (1924), may have been
misidentified C. sewardi.
Crematogaster subnuda discinodis Emery, 1893
Material examined. Kranji Road, 1.42638,
103.75413, 7 Sep 2016, G.W. Yong leg., ZRC_
HYM_0001766; Labrador Hill, 9 May 1985,
D.H. Murphy leg., ZRC_HYM_0000704; Pulau Semakau, Apr-Oct 2012, J. Puniamoorthy
et al. leg., mangroves, malaise trap, ZRC_BDP(multiple); Pulau Ubin, Apr-Oct 2012, J. Puniamoorthy et al. leg., mangroves, malaise trap,
ZRC_BDP (multiple); Sungei Buloh Wetland
Reserve, Apr-Oct 2012, J. Puniamoorthy et al.
leg., malaise trap, ZRC_BDP (multiple); University Hall (NUS), 1.29711, 103.77658, M.S.
Foo & W. Wang leg., malaise trap, ZRC_BDP(multiple).
Material not physically examined. Syntype –CASENT0904488, ANTC24399 (MSNG).
Literature. Type – Emery (1893) [as Crematogaster discinodis]. Wang et al. (2018a).
Localities. Kranji Road; Labrador Hill; Pulau Semakau; Pulau Ubin; Sungei Buloh; University
Hall (NUS).
Habitat/Ecology. This species was often found inboth mature and young secondary forests in
Singapore, including waste woodland secondary forest, frequently fragments in semi-urban
or urban settings; nests have been found in
fallen twigs. Individuals have also been collected in fairly large numbers from mangroves
using malaise traps.
Remarks. Type locality in Singapore. Crematogaster subnuda discinodis appears similar
to C. subnuda Mayr, 1879, but can be distinguished by its largely smooth and shiny
integument, and overall lighter body colours,
relative to the more rugulose and darker bodyof the C. subnuda type.
		However, Emery (1900) observed thatsamples of C. subnuda collected from partsof Indonesia could vary widely in terms ofhead and mesosomal sculpture, and also

body colour. It is not stated if this variation
occurred between or within different colony
samples. If extreme variation does occur between intranidal workers, it would support thesynonymy of subspecies discinodis under C.
subnuda. At the time of writing, we do nothave adequate material (i.e., colony series ofC. subnuda from a broader geographic range)necessary for evaluation. Thus, at this point
we refrain from making any unjustified taxonomic changes to the current subspecies.
Crematogaster treubi Emery, 1896
Material examined. Bukit Timah Nature Reserve, 1.35499, 103.78167, 19 Jul 2017,
W. Wang leg., ZRC_ENT00047931; I-cubebuilding (NUS), 1.293472, 103.77633, 1-8
Apr 2015, M.S. Foo & W. Wang leg., malaise trap, ZRC_BDP0045999; Lower Peirce
Reservoir, Aug 1990, collector unknown,
ZRC_ENT00000144; Pulau Semakau New
Fragment, Apr-Oct 2012, J. Puniamoorthy et
al. leg., mangroves, malaise trap, ZRC_BDP(multiple); Pulau Ubin, Apr-Oct 2012, J. Puniamoorthy et al. leg., mangroves, malaise
trap, ZRC_BDP (multiple); same locality
as previous, 7 Jan 2014, Sk. Yamane leg.,
ZRC_HYM_0000368-370; Sungei Buloh
Wetland Reserve, Apr-Oct 2012, J. Puniamoorthy et al. leg., malaise trap, ZRC_BDP(multiple); same locality as previous, 1.44632,
103.73057, 5 Dec 2018, W. Wang leg.,
ZRC_ENT00007638; Sunset Way, 1.32609,
103.77187, 23 Sep 2016, G.W. Yong leg.,
ZRC_HYM_0001765; Upper Peirce Reservoir, 10 Jan 2014, Sk. Yamane leg., ZRC_
HYM_0000371-374.
Material not physically examined. Queens, FOCOL1632-1633 (ZMHB) [types of Crematogaster millardi].
Literature. Viehmeyer (1916), Overbeck (1924)[both as C. millardi]. Wang et al. (2018a).
Localities. Bukit Timah Nature Reserve; Bukit
Timah Road; I-cube building (NUS); Lower
and Upper Peirce Reservoir; Pulau Semakau
New Fragment; Pulau Ubin; Sungei Buloh
Wetland Reserve; Sunset Way.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Habitat/Ecology. This species is associated mostly with young or mature secondary forests in
Singapore, including abandoned plantation
forest and other disturbed fragments in urban
or semi-urban settings. These ants have also
been found in mangroves, nesting in mangrove tree trunks or foraging on Thalassina(mud lobster) mounds.
Remarks. This species was first recorded fromSingapore as types of ‘C. millardi’ (Viehmeyer 1916); the latter species was synonymized
as a junior synonym of C. treubi by Hosoishi& Ogata (2012).
Crematogaster yappi Forel, 1901
Material examined. Pulau Semakau, 2012-2013,
J. Puniamoorthy et al. leg., mangroves, malaise trap, ZRC_BDP (multiple).
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Pulau Semakau (Old and New Fragments).
Habitat/Ecology. This species was collected frommangroves and/or mangrove backforests in
Singapore, specifically on an offshore island
landfill. A larger proportion of specimens werefrom newly replanted mangroves. Abundant
workers were collected using malaise traps,
implying that the species dwells or forages on
elevated vegetation above ground.
Remarks. At the time of writing, this species waspreviously only known from its type locality
in Kedah, Malaysia. The species’ restrictedoccurrence in mangroves here seems anomalous relative to the original habitat in its type
locality, i.e., Gunung Inas, Kedah, which isan inland area of relatively high elevation(>1000 m a.s.l.). It might be possible that local populations of C. yappi were introduced
through human commerce, or via imported
plants used for mangrove reforestation.

Crematogaster (Orthocrema) bandarensis Forel,
1913
Material examined. Bukit Batok East Avenue 6,
BB2, 1.34304, 103.76235, Sep 2016, G.W.
Yong leg., ZRC_HYM_0001716-1717; Kent
Ridge Park, 7 Dec 2017, W. Wang leg.,WWSG17-028, ZRC_ENT00000982; KranjiRoad, 1.42638, 103.75413, 8-10 Nov 2016,
ZRC_HYM_0001718-1720; Upper Seletar
Reservoir, 4 Oct 2020, G.W. Yong leg., GYSG20-003, ZRC_ENT00028568.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Batok East; Kent Ridge Park;
Kranji Road; Upper Seletar Reservoir.
Habitat/Ecology. This species was found mainly
in young and/or mature secondary forests in
Singapore, including abandoned plantation
and waste woodland forests. Specimens were
often collected in ground pitfall traps. In one
occurrence, the ants were found nesting at thebase of a tree, amidst leaf litter and tree roots.
Crematogaster (Orthocrema) longipilosa Forel,
1907
Material examined. Bukit Timah Nature Reserve,
4 Dec 1995, H.K. Lua et al. leg., NS 201B,
ZRC_ENT00000367; Lower Peirce Reservoir,
28 Jul 1990, H.K. Lua leg., LHK 108, ZRC_
HYM_0000142; Mandai Lake Road, 5 Nov
2016, G.W. Yong leg., ZRC_ENT00048422;
Mandai Road, 1.41155, 103.79015, 21-23
Nov 2016, G.W. Yong leg., M2D1GPc1931,
ZRC_HYM_0001764.
Material not physically examined. Unknown.
Literature. Forel (1912); Viehmeyer (1916);
Overbeck (1924).
Localities. Ayer Terjun (archaic place name);
Bukit Timah Nature Reserve; Bukit Timah
Road; Lower Peirce Reservoir; Mandai.
Habitat/Ecology. This species was mostly foundin primary and/or mature secondary forests in
Singapore, including native-dominated secondary forest. The ants have sometimes beenfound in Nepenthes pitcher cups, and alsofrequently collected with ground pitfall traps.

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Crematogaster (Orthocrema) myops Forel, 1911
Material examined. Queens, males, and workers,
Bukit Timah Nature Reserve, 8 Aug 2017,
W. Wang leg., ZRC_HYM_0001745; queens
and workers, same locality as previous, 4 Jan
2017, W. Wang leg.,WW-SG17-003, ZRC_
HYM_0000566; queens and workers, same
locality as previous, 1.35386, 103.77988, 23
Aug 2017, W. Wang leg.,WW-SG17-063,
ZRC_ENT00047959; same locality as previous, 1.35764, 103.77491, 7 Jun 2017, W. Wangleg., WW-SG17-064, ZRC_ENT00047960.
Material not physically examined.
CASENT0914541, PSW09576-9 (PSWC).
Literature. Hosoishi et al. (2010).
Localities. Bukit Timah Nature Reserve
Habitat/Ecology. This small-eyed subterranean
species was found in primary and old/maturesecondary forests in Singapore. Nests were
often found in, or in soil under rotting wood,
also in soil substrate and debris at bases oftrees, usually large dipterocarps.
Remarks. At the time of writing, this species isonly known from one locality in Singapore.
Crematogaster (Orthocrema) quadriruga Forel,
1911
Material examined. National University ofSingapore campus (Clementi/Kent Ridge),
Apr-Sep 2015, M.S. Foo & W. Wang leg.,malaise trap, ZRC_BDP (multiple); Pulau
Semakau Old Fragment, SMO2, 1°12’20.2”N,
103°45’35.8”E, 5-12 Jul 2012, J. Puniamoorthy et al. leg., mangroves, malaise trap, Reg.
29343, ZRC_BDP0015230; Pulau Ubin, 7 Jan
2014, Sk. Yamane leg., ZRC_HYM_0000387388; Sungei Buloh Wetland Reserve, 14 Mar
2018, W. Wang leg., WW-SG18-Crem1,
ZRC_ENT00000905; Ulu Sembawang
Road, 17 May 1994, collector unknown, NS
135, ZRC_ENT00000269; Temasek Polytechnic, adjacent waste woodland, 1.34239,
103.93598, 15 Dec 2016, G.W. Yong leg.,
TT1TH548, ZRC_HYM_0001763.
Material not physically examined. Unknown.
Literature. Wang et al. (2018a).

Localities. National University of Singapore campus (Clementi/Kent Ridge); Pulau Semakau
Old Fragment; Pulau Ubin; Sungei Buloh
Wetland Reserve; Ulu Sembawang Road;(beside) Temasek Polytechnic.
Habitat/Ecology. This species was often found in
disturbed mature or young secondary forest
fragments in urban or semi-urban settings, including waste woodlands, where the ants were
found foraging on tree trunks. Crematogaster
quadriruga has also been found in mangroves,where individuals have been collected fromtrails on elevated branches of living trees.
Remarks. This small, yellowish species is similar in appearance to C. bandarensis, but canbe distinguished from the latter by its small
propodeal spiracle that is located relatively
apart from the metapleural gland bulla - in C.
bandarensis the propodeal spiracle is large
and touching the metapleural gland bulla.
Dilobocondyla fulva Viehmeyer, 1916
Material examined. None available.
Material not physically examined. Types –CASENT0908982, ANTC31413 (MHNG);
workers, queen and male, FOCOL1971-1974(ZMHB) [as types of D. cataulacoidea var.
fulva].
Literature. Type – Viehmeyer (1916), Overbeck(1924) [both as D. cataulacoidea var. fulva].
Wheeler (1924); Taylor (1991) [raised to species].
Localities. Bukit Timah Road
Habitat/Ecology. A presumably arboreal species, the type colony series was found in ahollowed-out pith channel of a twig of a mangosteen tree in a garden.
Remarks. Type locality in Singapore. This species
was originally described as a subspecies of
Dilobocondyla cataulacoidea (see Viehmeyer1916), before being raised to species status
by Taylor (1991). A few congeneric worker
specimens from the same locality (i.e., Bukit
Timah Nature Reserve, not equivalent to thetype locality) were determined to confer morewith D. sebesiana Wheeler, 1924 sensu lato
(uncertain since only the queen has been described for the latter species).

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Erromyrma latinodis (Mayr, 1872)
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[both as Monomorium latinoda (sic) – misspelling of M. latinode].
Localities. Bukit Timah Road
Habitat/Ecology. This species was reportedly
found in a garden.
Remarks. At the time of writing, this species isonly known from Singapore based on literature records.
Eurhopalothrix heliscata Wilson & Brown, 1985
Material examined. Alate queen, Bukit Timah
Nature Reserve, BT4, 1.35342, 103.77820, 1
Mar 2017, M.S. Foo leg., malaise trap, Reg.
B8274, ZRC_BDP0072108; Rifle Range
Road, 6 Dec 2017, F. Ito leg., WW-SG17-024,
ZRC_ENT00000775; males, National University of Singapore (NUS) campus, 6 Dec
2017, F. Ito leg., emerged in lab – May – Jun
2019, F117-143 (SKYC).
Material not physically examined. Paratype –CASENT0900938, ANTC20060 (BMNH).
Literature. Type – Wilson & Brown (1984).
Localities. Bukit Timah Nature Reserve; NUS campus (Kent Ridge/Clementi); Rifle Range Road.
Habitat/Ecology. This species is associated withprimary and old/mature secondary forests in
Singapore. Nests were typically found in soil
under rotting wood or fallen logs.
Remarks. Type locality in Singapore. The actual
publication year of the article in which thisspecies was first described is 1984, however
1985 appears to be the accepted publication
year of the species in literature.
Eurhopalothrix omnivaga Taylor, 1990
Material examined. Bukit Timah Nature Reserve, 25 Sep 2019, D.J. Court leg., ZRC_
ENT00013947; Nee Soon Swamp (Forest),
1.39932, 103.80954, 13 Sep 2015, M.K.L.
Wong leg., ZRC_ENT00000689.
Material not physically examined. Unknown.
Literature. None. New record.

Localities. Bukit Timah Nature Reserve; Nee Soon
Swamp Forest.
Habitat/Ecology. This species was found in primary and/or mature secondary forests in Singapore, including swamp forest. Specimens
were collected from soil and leaf litter.
Eurhopalothrix procera (Emery, 1897)
Material examined. Bukit Batok East, 29 Sep
2016, G.W. Yong leg., ZRC_ENT00014124;
alate queen, Bukit Timah Nature Reserve, 7 Sep 2016, M.S. foo leg., ZRC_
HYM_0001735; same locality as previous,
1.34796, 103.77914, 29 Oct 2015, M.K.L.
Wong leg., ZRC_ENT00000686; Loyang,
25 Dec 1991, D.H. Murphy leg., DHMSG91-Eur1, ZRC_ENT00027934; queens,male and workers, Mandai Track, 4 Jan
2017, G.W. Yong leg., GY-SG17-001, ZRC_
ENT00027987; Nee Soon, NS141A, 1994,
collector unknown, ZRC_ENT00027943;
Upper Thomson Nature Park, 1.38311,
103.79839, 19 Oct 2016, G.W. Yong leg.,
UT2GH1613, ZRC_HYM_0001641.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Batok East; Bukit Timah Nature
Reserve; Loyang; Mandai Track; Nee Soon;
Upper Thomson Nature Park.
Habitat/Ecology. This species is associated mostly with primary and secondary forests in Singapore, including young abandoned park andplantation secondary forests. Individuals were
often collected from rotting logs and leaf litter.
Nests were found under loose bark of fallentrees or logs.
Remarks. Often mistaken for and assumed to be
E. heliscata (especially since E. heliscata was
first described from Singapore), E. procera ismorphologically distinct, e.g. body sculpture
overall less coarse, body hairs sparse, generally finer and not explicitly clavate. Specimens in Singapore confer more with the formof E. procera malua (Mann, 1919) than the
primary types of E. procera, although the subspecies is now considered a junior synonym of
the latter (thus invalid). We observed distinct
differences in structure between original E.
procera and the currently invalid subspecies

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malua that may warrant future raising of thelatter to a separate species. For this checklist,
however, in the absence of more compelling
empirical evidence, we tentatively accept theconspecificity of E. procera and the subspecies malua.
Gauromyrmex acanthinus (Karavaiev, 1935)
Material examined. Bukit Timah Nature Reserve,near BT05, 1.35386, 103.77988, 23 Aug
2017, W. Wang leg., WW-SG17-018, ZRC_
HYM_0001747; same locality and collector
as previous, 1 Nov 2017, ZRC_EnT00047952.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve
Habitat/Ecology. This species is associated withprimary or old secondary forest in a single
locality. In one instance, the ants were foundnesting under the bark of a very large livingtree. They were also observed in large deadlogs or fallen rotting branches.
Liomyrmex gestroi (Emery, 1887)
Material examined. Bukit Timah Nature Reserve, ca. 50 m uphill of Main Path, 1.3489,
103.77736, 15 Mar 2017, W. Wang leg.,trunk cavity of very large fallen tree, ZRC_
HYM_0001169; Nassim Road, 25 Sep 1969,
D.H. Murphy leg., ZRC_HYM_0000018; Pulau Ubin, 1.41662, 103.99443, 4 Dec 2016,
G.W. Yong & M.K.L. Wong leg., ZRC_
HYM_0001072; Rifle Range Road, 27 Jul
2016, G.W. Yong leg., ZRC_HYM_00000994;
Racecourse forest, 11 Jan 1981, D.H. Murphyleg., DHM-SG81-Lio1, ZRC_ENT00027928.
Material not physically examined. Non-type
workers, specific locality unknown, Sk. Yamane leg. (SKYC) – in Rigato & Bolton(2011).
Literature. Rigato & Bolton (2011); Yong et al.
(2017).
Localities. Bukit Timah Nature Reserve; Nassim
Road; Pulau Ubin; Rifle Range Road; Racecourse Forest.

Habitat/Ecology. This species is associated withprimary or secondary forests in Singapore,including young secondary forests. The ants
have been found mostly in or under bark oflarge dead trees in old secondary forest. In one
instance, nestmates were found in the trunk
cavity of a very large, relatively fresh-fallentree. Though often reported elsewhere to be
associated with termites, no such associations
were observed or recorded for local occurrences of this species. The ants have also beenfound boring in rotten timber under bark ofstanding dead trees ca. 50 feet above ground,in deep bark scrapings of trees, and underfrais-filled bark.
Lophomyrmex bedoti Emery, 1893
Material examined. Locality and collection dateunknown (only ‘Singapore’ indicated on label), D.H. Murphy leg., ZRC_HYM_0000634;
locality unknown [but deduced as Nee Soonbased on ‘swamp forest’ on original label], 14
Apr 1993, Sk. Yamane leg. (SKYC).
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah Road; Nee Soon.
Habitat/Ecology. This species was found in therotten trunk of a fallen tree on the ground, ina garden. More recently these ants were alsocollected from swamp forest.
Lordomyrma cf.sp.7.of.SKY (*excluded fromofficial named species count)Material examined. Queens, males and workers,
Mandai, Northern Node, 1.40842, 103.78319,
54 m a.s.l., 25 Jul 2019, J. Tan & S. Tangleg., JTNC-MIC-005, ZRC_ENT00013559;
Mandai Bike Trail, 1.40469, 103.77653,
10 Mar 2016, M.K.L. Wong leg., ZRC_
ENT00000690; Mandai Track, 4 Jan 2017,
G.W. Yong leg., ZRC_ENT00027990; Pulau Ubin, 12 Jan 2015, W. Wang leg., ZRC_
ENT00000980; Upper Thomson Road, 11 Oct
2016, G.W. Yong leg., ZRC_HYM_0000997.
Material not physically examined. Unknown.
Literature. None. New genus record.
Localities. Mandai; Pulau Ubin; Upper Thomson
Road.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Habitat/Ecology. This species was collected fromold or mature secondary forests, including
abandoned plantation forest. The ants weresometimes found nesting in fallen rotting
wood, logs or tree branches.
Remarks. This is the first record of the genus
Lordomyrma in Singapore; unfortunately,
we were unable to identify local specimens
to known species at the time of writing. This
species should be closely related to L. reticulata or L. azumai based on general habitus
and strong rugulose-reticulate sculpture. It is
most similar morphologically to L. idianalethat was described from the Philippines, butwith small nodules on the pronotal humeri
amongst other more subtle differences.
Mayriella transfuga Baroni Urbani, 1977
Material examined. Bukit Timah Nature Reserve, BT07, 1.35499, 103.78167, 22 Dec
2016, W. Wang leg., soil, WW-SG16-005,
ZRC_HYM_0000353; same locality as
previous, 1.34796, 103.77914, 29 Oct
2015, M.K.L. Wong leg., Berlese extraction, ZRC_ENT00000691-692; Nee Soon,
1.38245, 103.80206, 46 m, 26 Apr 2018,
W. Wang leg., Winkler extraction, NS_W2,
ZRC_ENT00000954; Pulau Ubin, 22 Oct
2016, W. Wang leg., Winkler extraction,
ZRC_ENT00000987; same locality as previous, Jalan Sam Heng, 1.41260, 103.99317, 4
Dec 2016, G.W. Yong & M.K.L. Wong leg.,
Winkler extraction, ZRC_HYM_0001136.
Material not physically examined. ANIC32014977, Shattuck1345185593; ANIC32015009, Shattuck1345233190 (ANIC).
Literature. Taylor (1991); Shattuck & Barnett(2007).
Localities. Bukit Timah Nature Reserve; Nee
Soon; Pulau Ubin.
Habitat/Ecology. This species was found mostly
in primary and/or mature secondary forests, aswell as young secondary forests in Singapore,including Adinandra belukar and swamp forest. These minute ants were usually collectedfrom leaf litter or leaf mound berlesate.

Meranoplus bicolor (Guérin-Méneville, 1844)
Material examined. Botany Dept. Garden (NUS), 1 Nov 1990, H.K. Lua leg.,
ZRC_ENT00000260; Changi sand dunes,
21 Nov 1982, D.H. Murphy leg., ZRC_
HYM_0000696; Commonwealth Avenue
West, 6 Apr 2006, H.K. Lua leg., LHK
461, ZRC_HYM_0000187; I-cube building(NUS), 1.29347, 103.77633, 24 Jun-1 Jul
2015, M.S. Foo & W. Wang leg., malaise trap,
NUS0054, ZRC_BDP0044222; University
Town (NUS), 1.30622, 103.77458, Apr-May
2015, M.S. Foo & W. Wang leg., malaise trap,
ZRC_BDP multiple; Kranji Road, 1.42638,
103.75413, 1-10 Nov 2016, G.W. Yong leg.,
ZRC_HYM_0001274-1278; Pulau Ubin,
1.40308, 103.96924, 4 Dec 2016, G.W. Yong
& M.K.L. Wong leg., ZRC_HYM_0001073;
Queen’s Crescent, 14 Mar 2006, H.K. Lualeg., LHK 460, ZRC_HYM_0000185; Sime
Road, 14 Jul 2009, H.K. Lua leg., LHK
508, ZRC_HYM_0000188; Temasek Polytechnic, adjacent waste woodland, 1.34239,
103.93598, 13-15 Dec 2016, G.W. Yongleg., ZRC_HYM_0001272-1273; Tuas West
Drive, 31 Mar 2007, H.K. Lua leg., LHK 466,
ZRC_HYM_0000184; Upper Peirce Reservoir, 10 Jan 2014, Sk. Yamane leg., ZRC_
HYM_0000406-407; Western Catchment,
1.40866, 103.70185, 20 Nov 2015, M.K.L.
Wong leg., ZRC_ENT00000693.
Material not physically examined. Non-types,
J.K. Wetterer leg. (JKWC) - Kallang, Riverside Park, 29 Jul 2014, vial #372; Punggol, by
MRT station, 30 Jul 2014, vial #390; Bishan,
31 Jul 2014, vial #424; Tai Seng, by MRT
station, 31 Jul 2014, vial #422; East Coast
Park (Katong), 1 Aug 2014, vial #442; Pasir
Panjang, by MRT station, 5 Aug 2014, vial
#482; Singapore Botanic Gardens, 5 Aug
2014, vial #477.
Literature. Viehmeyer (1916); Overbeck (1924);
Tan & Corlett (2012); Wang et al. (2018a).
Localities. Bukit Timah Road; Changi; Commonwealth Avenue West; East Coast Park (Katongpart); Kranji Road; National University ofSingapore campus (multiple sites); Pasir Panjang; Pulau Ubin; Punggol; Queen’s Crescent;Riverside Park (Kallang); Sime Road; Sin-

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gapore Botanic Gardens; Tai Seng; Temasek
Polytechnic; Tuas West Drive; Upper Peirce
Reservoir; Western Catchment.
Habitat/Ecology. This species was commonly
found in young secondary forests in Singapore, also in recreational parks and other cultivated areas in urban or semi-urban settings,
and grasslands. Individuals have also beencollected from disturbed and waste woodland
secondary forests; one occurrence reported inmarshland. This species is known to nest in
soil and forage mostly on the ground, though
individuals have been collected via malaisetraps and sweep netting of vegetation. One
ground nest was described as forming ‘narrowpassages in loamy soil which go deep below
the surface.’
Meranoplus malaysianus Schödl, 1998
Material examined. Holland Woods, 1.32982,
103.78236, 16 Oct 2017, W.N. Lam leg.,
HW02_05, ZRC_ENT00047926; Kent Ridge,
1.28707, 103.7862, 9 Jan 2018, R.J.Y. Lim etal. leg., ZRC_ENT00047841; same locality
and collectors as previous, 1.28791, 103.7855,
19 May 2017, ZRC_ENT00047842.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Holland Woods; Kent Ridge.
Habitat/Ecology. This minute species was foundin mature secondary forests, frequently inlower pitcher contents of Nepenthes ampullaria or N. rafflesiana.
Meranoplus mucronatus Smith, 1857
Material examined. Mandai, 10 Sep 1980, Tay
Soon Cheng leg., sugar [sic] plantation, ZRC_
ENT00047925; locality and collectors unknown, Oct-Nov 1988, ZRC_ENT00028677.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Mandai
Habitat/Ecology. This species was once found ina sugarcane plantation in Singapore.

Remarks. This species is the largest of the genus
Meranoplus in the Oriental region, and can beeasily distinguished from its congeners in thesame region by its strikingly armed promesonotal shield. The record of M. mucronatus
in Viehmeyer (1916) was of specimens from
Gunong Angsi in Malaysia, not Singapore as
is often mistaken.
Metapone murphyi Wang, Yamada & Eguchi,
2019
Material examined. Holotype, Pulau Sakra, 7 Mar
1981, D.H. Murphy leg., DHM-SG81-Meta1, ZRC_ENT00000878; paratypes, worker,
queens and males, same locality and collecting data as holotype, ZRC_HYM_0000016.
Material not physically examined. Unknown.
Literature. Wang et al. (2019).
Localities. Pulau Sakra
Habitat/Ecology. The type series of this species
was found in a decayed coconut stump, on an
offshore island that has long since been heavily developed for industrial purposes.
Remarks. Type locality in Singapore. At the timeof writing, the only known record of this species is that of the type series from Singapore.
Considering the elusive termitophilous natureof the genus Metapone, M. murphyi might bethriving elsewhere undiscovered on the mainisland or other offshore islands in association
with termite nests.
Monomorium atomum procax Forel, 1911
Material examined. None available.
Material not physically examined. Syntype –CASENT0908766, ANTC31116 (MHNG).
Literature. Type – Forel (1911) [as Monomorium (Martia) atomus var. procax]. Viehmeyer(1916), Overbeck (1924) [both as M. (Martia)
atomus var. procax].
Localities. Bukit Timah Hill; Bukit Timah Road.
Habitat/Ecology. This species was found in primary or mature/old secondary forest, also ina garden, where colonies were collected fromdead, hollowed-out (tree) branches.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Remarks. Type locality in Singapore. We do nothave more recent physical material for reliable
systematic assessment. Differences between
M. atomum and its subspecies procax are also
subtle and difficult to discern from type images on AntWeb. Nevertheless, we observed
one striking difference: the propodeum in M.
atomum is distinctly much longer than wide in
dorsal view, whereas in M. atomum procax the
propodeum is only slightly longer than wide.
We maintain the subspecies procax for wantof more compelling empirical evidence andmaterial to support a raise in status to species.
Monomorium butteli demochrum Viehmeyer,
1916
Material examined. None available.
Material not physically examined. Types –CASENT0908694, ANTC31044 (MHNG);
dealate queen and workers, FOCOL1693-1696(ZMHB).
Literature. Type – Viehmeyer (1916); Overbeck(1924).
Localities. Bukit Timah Road
Habitat/Ecology. The type series was found in agarden under a portico, specifically under aflower-pot on a table.
Remarks. Type locality in Singapore. At the timeof writing, the only record of this species isthat of the type series from Singapore.
Monomorium floricola (Jerdon, 1851)
Material examined. Bukit Batok East, 1.34304,
103.76235, Sep-Oct 2016, G.W. Yong leg.,trunk pitfall, ZRC_HYM_0001282-1283;
Commonwealth Road, 1.31115, 103.78171,
22-24 Nov 2016, G.W. Yong leg., ZRC_
HYM_0001286-1287; Kranji Road, 1.42638,
103.75413, 1-3 Nov 2016, G.W. Yong leg.,
ZRC_HYM_0001288-1289; queens, males
and workers, National University of Singapore campus (Clementi/Kent Ridge, multiple sites), Apr-Sep 2015, M.S. Foo & W.
Wang leg., malaise trap, ZRC_BDP (multiple); Pulau Ubin, 7 Jan 2014, Sk. Yamaneleg., ZRC_HYM_0000380-381; queens,males and workers, Sungei Buloh Wetland
Reserve, 28 Mar 2018, W. Wang leg., WW-

SG18-Mono1, ZRC_ENT00000959; queens
and workers, same locality as previous, 8
Jan 2014, Sk. Yamane leg., SG14-SKY-34,
ZRC_HYM_0000440; Sunset Way, 1.32609,
103.77187, 21-23 Sep 2016, G.W. Yong leg.,
ZRC_HYM_0001280-1281; Temasek Polytechnic, adjacent waste woodland, 1.34239,
103.93598, 13-15 Dec 2016, G.W. Yong leg.,
ZRC_HYM_0001284-1285; Upper Thomson
Nature Park, 1.38311, 103.79839, Oct 2016,
G.W. Yong leg., ZRC_HYM_0001290-1292.
Material not physically examined. Non-types,
J.K. Wetterer leg. (USNM) – Amber Road
(Katong), 28 Jul 2014, vials #331, 345, 351;
Riverside Park (Kallang), 29 Jul 2014, vial
#372; City Hall, by church, 29 Jul 2014, vial
#374; Outram Park, by MRT station, 29 Jul
2014, vial #379; Queenstown, 29 Jul 2014,
vial #383; Promenade, by MRT station, 4 Aug
2014, vial #468.
Literature. Viehmeyer (1916); Overbeck (1924);
Wang et al. (2018a).
Localities. Bukit Batok East; Commonwealth
Road; Kranji Road; National University ofSingapore campus (Clementi/Kent Ridge);
Pulau Ubin; Sungei Buloh Wetland Reserve;
Sunset Way; beside Temasek Polytechnic; Upper Thomson Nature Park.
Habitat/Ecology. This species appears very common in Singapore and was often found in a
broad variety of different habitats across thedisturbance spectrum, including: abandoned
plantation and waste woodland secondary forests, disturbed secondary forest fragments inurban or semi-urban settings, gardens, mangroves, sometimes even in urban infrastructure including buildings. In mangroves, nests
were found in bark of dead tree trunks, andalso in dead twigs on trees, usually with multiple queens per colony. Colonies have also
been found in hollow twigs of mangosteen
trees and in withered bamboo.
Remarks. This species is a recognized common
tramp ant widespread throughout the wholeworld, mostly in warm tropical regions.

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Monomorium hospitum Viehmeyer, 1916

Monomorium monomorium Bolton, 1987

Material examined. None available.
Material not physically examined. Syntypes –4 queens, 4 males (MNHU). Types – queen,
male, workers, FOCOL1727-1730 (ZMHB).
Literature. Type – Viehmeyer (1916), Overbeck(1924) [both as Monomorium (Corynomyrmex) hospitum]. Bolton (1987).
Localities. Bukit Timah Road
Habitat/Ecology. The type series of this species
was found with M. floricola workers in a garden, specifically inside the narrow passagesof a branch stump.
Remarks. Type locality in Singapore. Based onimages on AntWeb, the worker of M. hospitum resembles the more widespread M. floricola, but with subtle differences (M. floricola
traits in parentheses): 1) posterior margin ofhead broadly convex, transitions to lateral
margin in a smooth curve without apparent
angle (posterior margin almost flat with weak
median depression, posterior corners roundly
angulate), 2) propodeal junction distinctly
angulate (propodeal junction indistinct andobtusely rounded).
		Monomorium hospitum was originallydescribed based only on queens and males.
Viehmeyer (1916) surmised that M. hospitum
appeared ‘parasitic’ on M. floricola, as alatesof the former species were collected from thelatter’s nest. The author also felt that the alatesof M. hospitum were distinct enough to warrant the establishment of a new subgenus —the now-defunct ‘Corynomyrmex’. The queenof M. hospitum is easily distinguished fromthat of M. floricola by its relatively convex
mesosomal dorsum, such that mesosoma appears ovate in profile (in M. floricola queens,mesosomal dorsum is almost entirely flat andstraight, mesosoma more cylindrical in profile).

Material examined. None available.
Material not physically examined. Unknown.
Literature. Tan & Corlett (2012).
Localities. National University of Singapore campus (Kent Ridge); West Coast Park.
Habitat/Ecology. Specimens identified as thisspecies were collected from young secondary
forests, recreational parks (mown grasslandwith trees and shrubs), and open grasslands.
Remarks. At the time of writing, this presumably non-native species is only known fromSingapore based on one relatively recent literature record that could not be verified fortaxonomic accuracy. Some specimens available in the ZRC were morphologically similar
to M. monomorium sensu lato but could not be
identified as the latter with confidence – thesewere at best designated as undescribed speciesof the M. monomorium group.
Monomorium pharaonis (Linnaeus, 1758)
Material examined. National University of Singapore (NUS) building, 10 Jul 2007, D. Lohman
leg. (SKYC).
Material not physically examined. ANIC32062894-062896 (ANIC); CASENT0102330,
ANTC4573 (BMNH) [former lectotype of‘Myrmica fragilis’ – now invalid name];
CASENT0008625, ANTC2204 (OUMNH)[former lectotype of ‘Monomorium/Myrmica
vastator’ – now invalid name]. Non-types,
J.K. Wetterer leg. (JKWC) – Harbourfront,
29 Jul 2014, vial #385; Katong, Amber Road,
31 Jul 2014, vial #434; Mountbatten, by MRT
station, 4 Aug 2014, vial #465; Queenstown,
29 Jul 2014, vial #383.
Literature. Smith (1857) [as Myrmica vastator];
Smith (1858) [as Myrmica fragilis]; Viehmeyer (1916); Overbeck (1924); Heterick (2006);
Tan & Corlett (2012).
Localities. Bukit Timah Road; Bukit Timah Nature
Reserve (BTNR); Central Catchment Nature
Reserve (CCNR); NUS campus.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Habitat/Ecology. These ants were commonly
found both foraging and nesting indoors in
artificial human-constructed infrastructure,such as in households and offices, also in backgardens of residential housing. This species
was also reportedly collected at baited traps
set up in primary and/or old secondary forests.
Remarks. As with many other congeners, Monomorium pharaonis is considered a globallywidespread tramp ant, invasive to many areas,
and is also infamous as a common household/indoor pest where it occurs. Records of thespecies from primary or old secondary forestsin the CCNR and BTNR were only in literature and could not be taxonomically verified
at the time of writing. It is possible that theserecords were cases of misidentification.
Myrmecina bandarensis Forel, 1913
Material examined. Alate queen, Nee Soon
Swamp Forest, NS1, 1°23’00.3”N,
103°48’46.5”E, 28 Mar-4 Apr 2012, J. Puniamoorthy et al. leg., malaise trap, Reg. 29157,
ZRC_BDP0014622.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Nee Soon Swamp Forest
Habitat/Ecology. A single alate queen of this species was found in freshwater swamp forest,
collected in a malaise trap sample.
Myrmecina magnificens Wong & Guénard, 2016
Material examined. Holotype, Central Catchment Nature Reserve, Seletar Trail, 1.39514,
103.80260, 47 m, 2 Sep 2015, M.K.L. Wongleg.,Winkler extraction, ZRC_ENT00000665;
Bukit Timah Nature Reserve, 1.34883,
103.77869, 9 May 2017, W. Wang leg., ZRC_
HYM_0000982; same locality as previous,
30 May 2016, D.J. Court leg., pitfall trap,
ZRC_ENT00048382; Mandai Road, 1.41258,
103.79839, 5-7 Dec 2016, G.W. Yong leg.,
M1D7GPb2495, ZRC_HYM_0001293.
Material not physically examined. Four paratypeworkers (HKUC).
Literature. Type - Wong & Guénard (2016).
Localities. Bukit Timah Nature Reserve; Central
Catchment Nature Reserve; Mandai Road.

Habitat/Ecology. This species is associated withprimary and/or mature secondary forests in
Singapore, typically in soil and leaf litter.
Remarks. Type locality in Singapore.
Myrmecina maryatiae Okido, Ogata & Hosoishi,
2020
Material examined. Bukit Timah Nature Reserve,
1.35599, 103.77397, 2 Aug 2017, W. Wangleg., ZRC_HYM0001743; Chestnut Nature
Park, 1.37451, 103.78137, 27 Feb 2022,
T.K.C. Leung leg., ZRC_ENT00054741.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Chestnut
Nature Park.
Habitat/Ecology. Individuals of this species were
found in primary and Dillenia suffruticosadominated young secondary forest, on fallen
logs.
Myrmicaria adpressipilosa Santschi, 1928 stat. n.
Myrmicaria (Heptacondylus) arachnoidea [sic]var. adpressipilosa Santschi, 1928: 130 (w.)
Subspecies of Myrmicaria arachnoides: Bolton,
1995: 285; Pfeiffer et al., 2011: 16.
Material examined. Queen and workers, NeeSoon forest, 11 Feb 1974, D.H. Murphy leg.,
ZRC_ENT00006313.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Nee Soon forest
Habitat/Ecology. This species was found inswamp forest and/or its adjacent drier nativedominated secondary forest, also in old or
mature secondary forest further inland. The
ants build carton nests on leaves and foliage,
usually on undersides of large tree leaves.
Remarks. Myrmicaria adpressipilosa was originally described as a subspecies of Myrmicariaarachnoides (Smith, 1857). Emery (1900)
stated that M. adpressipilosa resembled
Myrmicaria arachnoides lutea Emery, 1900
in terms of body colour and general habitus,
but could be differentiated from the latter, by
the mostly adpressed or decumbent hairs on
its body. Based on colonies of both taxa fromother parts of Southeast Asia, it was later determined that the adpressed hairs are really

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an artifact and not diagnostic of the species(Bakhtiar & Yamane 2022, unpublished)
Further examination of those colonies revealed more compelling differences between
workers of both species (Bakhtiar & Yamane
2022, unpublished). Myrmicaria adpressipilosa can be distinguished from M. arachnoides based on the following (M. arachnoides
traits in parentheses): 1) propodeal spiracle
large, its maximum diameter slightly shorter or longer than distance between spiracleand dorsal margin of propodeum (propodeal
spiracle small, its maximum diameter much
shorter than distance between spiracle anddorsal margin of propodeum), 2) pronotal andpropodeal sides irregularly striate (pronotal
and propodeal sides extensively smooth andshining), 3) propodeal declivity with laterallongitudinal carinae (propodeal declivity lacking lateral carinae) [Bakhtiar & Yamane 2022,
unpublished].
Myrmicaria arachnoides (Smith, 1857)
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah Road
Habitat/Ecology. Alate queens and males were
found at a lit-lamp in a garden; in the samegarden, a dealate queen was also found in asmall pasteboard cell on the underside of aleaf.
Remarks. At the time of writing, this species isonly known from Singapore based on past
literature records. A number of the alates collected at the same time by Overbeck were lateron identified as the subspecies lutea and luteiventris respectively (see Viehmeyer 1922).
Myrmicaria arachnoides lutea Emery, 1900
Material examined. None available.
Material not physically examined. Types– queens and males, FOCOL1602-1605(ZMHB).
Literature. Viehmeyer (1922) [type queen andmale].

Localities. Bukit Timah Road
Habitat/Ecology. Alates were collected from a
lit-lamp in a garden.
Remarks. The queen and male of this subspecies were first described by Viehmeyer (1922)
based on specimens collected from Singapore,that were in turn initially identified as M.
arachnoides in Viehmeyer’s earlier publication (see Viehmeyer 1916).
Myrmicaria luteiventris Emery, 1900
Material examined. Queens, males and workers, Pulau Ubin, 1.42022, 103.94598, 31 Aug
2016, M.S. Foo, W. Wang & M. Tuan leg.,
WW-SG16-001, ZRC_HYM_0000247; same
locality as previous, 1.41662, 103.99443, 4
Dec 2016, G.W. Yong & M.K.L. Wong leg.,
ZRC_HYM_0001079.
Material not physically examined. Type queen– FOCOL1597 (ZMHB).
Literature. Viehmeyer (1922) [type queen].
Localities. Bukit Timah Road; Pulau Ubin.
Habitat/Ecology. This subspecies was found in
relatively disturbed old or mature secondary
forest on an offshore island, where colonies
were found in carton nests on the undersidesof large leaves. Alates were collected from a
lit-lamp in a garden.
Remarks. Myrmicaria luteiventris was originallydescribed as a subspecies of M. arachnoides.
It was later raised to species status in Pfeiffer
et al. (2011), but the authors did not give any
justification for this – it might have been likely
a typographical error.
		The queen of this species was first described by Viehmeyer (1922) based on specimens collected from Singapore, which were
initially identified as M. arachnoides in Viehmeyer’s earlier publication (see Viehmeyer1916).
		The workers of M. luteiventris resemble
that of M. arachnoides, except for their gaster,which is (for at least its posterior half) muchpaler in colour than the head and mesosoma.
In M. arachnoides, the body is not distinctly
bicolorous.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy		Further examination of colonies fromother parts of Southeast Asia revealed othermore compelling differences between workers
of both species (Bakhtiar & Yamane 2022,
unpublished). Myrmicaria luteiventris can be
distinguished from M. arachnoides based on
the same characters described under Remarksfor M. adpressipilosa. As with M. arachnoides, M. luteiventris can also be differentiatedfrom M. adpressipilosa by the latter’s generally uniform body colour.

Literature. Bolton (1988).
Localities. Sungei Buloh Wetland Reserve
Habitat/Ecology. A single queen was found in amalaise trap sample from mangroves.
Remarks. Queens and males of this species canbe distinguished from other congeners by thepresence of long, acutely-pointed, curved finehairs projecting from the tibiae, also similar
long and fine curved standing hairs on entire body dorsum, including the head (Bolton1988).

Myrmicaria melanogaster Emery, 1900

Pheidole aglae Forel, 1913

Material examined. Peirce Catchment, 28 Dec
1973, D.H. Murphy leg., A340-6, ZRC_
ENT00006312; same locality and collection
data as previous, ZRC_HYM_0000771.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Peirce Catchment
Habitat/Ecology. This species appears associatedwith either primary or old/mature secondary
forest in Singapore, as inferred from its collection locality. Some specimens were alsocollected from the forest edge.

Material examined. Sunset Way, 1.32609,
103.77187, 21-23 Sep 2016, G.W. Yongleg., pitfall trap, SW5GPc324, ZRC_
HYM_0001779.
Material not physically examined. FOCOL1356(ZMHB) [type of P. exasperata var. polita –now invalid name].
Literature. Viehmeyer (1916), Overbeck (1924) [both as Pheidole exasperata var. polita]; Eguchi (2001); Tan & Corlett (2012).
Localities. Bukit Timah Hill; MacRitchie Reservoir; Sunset Way.
Habitat/Ecology. This species was found mostly
in primary or mature secondary forests in
Singapore, sometimes young secondary habitats such as abandoned plantation secondary
forests. Colonies have been found in rotten
branches and rotting wood on the forest floor.

Paratopula demeta Bolton, 1988
Material examined. MacRitchie, 1.34291,
103.82689, 15 Jun 2016, M.K.L. Wong leg.,
MKLW000049, ZRC_ENT00027953.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. MacRitchie
Habitat/Ecology. A single worker was found on
a vine in primary or old/mature secondary
forest. This genus is known to be largely arboreal.
Paratopula oculata (Smith, 1857)
Material examined. Dealate queen, Sungei Buloh Wetland Reserve, SB2, 1°26’47.7”N,
103°43’49.9”E, 24-31 Oct 2012, J. Puniamoorthy et al. leg., malaise trap, Reg. 29544,
ZRC_BDP0016446.
Material not physically examined. One female
specimen (queen) in BMNH – identified by
Bolton (1988).

Pheidole aristotelis Forel, 1911
Material examined. Bukit Timah Nature Reserve,
1.35499, 103.78167, 19 Jul 2017, W. Wangleg., ZRC_ENT00047932; same locality as
previous, 5 Jun 1968, D.H. Murphy leg., D610, ZRC_HYM_0000733; Mandai Road,
1.41155, 103.79015, 21 Nov 2016, G.W. Yongleg., M2D1GH1938, ZRC_HYM_0001782.
Material not physically examined. Unknown.
Literature. Eguchi (2001); Tan & Corlett (2012).
Localities. Bukit Timah Nature Reserve; Central
Catchment Nature Reserve; Mandai Road.
Habitat/Ecology. This species was found in primary or old/mature secondary forests in Singapore, including native-dominated secondary forest.

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Pheidole binghamii Forel, 1902
Material examined. Seletar Link, 1.40518,
103.88555, 25 Oct 2016, G.W. Yong leg.,
SL6GH1060, ZRC_HYM_0001780.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Seletar Link
Habitat/Ecology. Individual workers were collected from waste woodland secondary forest.
Remarks. At the time of writing, this species isonly known from a one locality in Singapore.
Pheidole cariniceps Eguchi, 2001
Material examined. Bukit Timah Nature Reserve, 24 Aug 2015, J. Koh leg., ZRC_
ENT00013867; queen and workers, same locality as previous, 1.35386, 103.77988, 4 Jan
2017, W. Wang leg.,WW-SG17-004, ZRC_
HYM_0000568; same locality as previous,
plot III-2, 12 Oct 2015, D.J. Court leg., pitfall trap, ZRC_ENT00048443; Mandai Road,
1.41258, 103.79839, 16 Nov 2016, G.W. Yongleg., M1D4GH2163, ZRC_HYM_0001777.
Material not physically examined. Unknown.
Literature. Tan & Corlett (2012).
Localities. Bukit Timah Nature Reserve; Central
Catchment Nature Reserve; Mandai Road.
Habitat/Ecology. This species was found in primary and old/mature secondary forests in
Singapore, including native-dominated secondary forest. The ants were found in rotting
wood; individuals were usually collected fromleaf litter.
Pheidole clypeocornis Eguchi, 2001
Material examined. Bukit Timah Nature Reserve, BT05, 1.35386, 103.77988, 4 Jan
2017, W. Wang leg., WW-SG17-001, ZRC_
HYM_0000564; same locality and collector as previous, BT02, 15 Mar 2017, WWSG17-040, ZRC_ENT00007803; queen andworkers, Mandai Lake Road, Night Safari,N01°24’22.0’’, E103°47’04.7’’, 48 m, 8
Sep 2020, J.S. Tan & M. Bte. Hussin leg.,
JTM-MIS-017, ZRC_ENT00047775; males,

queens and workers, National University ofSingapore campus (Clementi/Kent Ridge,
multiple sites), Apr-Aug 2015, M.S. Foo &W. Wang leg., malaise trap, ZRC_BDP (multiple).
Material not physically examined. Unknown.
Literature. Tan & Corlett (2012); Wang et al.
(2018a).
Localities. Bukit Timah Nature Reserve; Central
Catchment Nature Reserve; Mandai Lake
Road; National University of Singapore campus (Clementi/Kent Ridge).
Habitat/Ecology. This species was found in bothprimary and old/mature secondary forests in
Singapore, including disturbed secondary
forest fragments in urban or semi-urban settings. In primary forest, colonies were foundin fallen twigs or in soil near bases of agedrotting wood. In old secondary forest, the ants
were found nesting under bark of fallen rotting
branches or logs, also on leaf litter. Numerous
alates were collected from cultivated grassy
patches and disturbed secondary forest fragments in urban and semi-urban settings, usingmalaise traps.
Remarks. The minor workers of P. clypeocornis
are almost indistinguishable from those of P.
hortensis. Species occurrence was therefore
only confirmed where major workers wereavailable, or when DNA barcodes (from alates
and minors) matching the verified species
were obtained (see Wang et al. 2018a).
Pheidole elisae Emery, 1900
Material examined. Bukit Timah Nature Reserve,plot III-2, 19 Apr 2016, D.J. Court leg., pitfall trap, ZRC_ENT00048453; Mandai Lake
Road, 5 Nov 2016, G.W. Yong leg., ZRC_
ENT00048424.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Mandai
Lake Road.
Habitat/Ecology. This species was found in old or
mature secondary forests in Singapore.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Pheidole fervens Smith, 1858

Pheidole magrettii Emery, 1887

Material examined. Labrador beach, 29 Apr 1985,
D.H. Murphy leg., ZRC_HYM_0000728.
Material not physically examined. Syntypes –CASENT0901519-0901520, ANTC21037(BMNH).
Literature. Type – Smith (1858). Viehmeyer(1916), Overbeck (1924) [both as Pheidole
javana]. Eguchi (2001, 2004); Sarnat et al.
(2015).
Localities. Bukit Timah Road; Labrador beach.
Habitat/Ecology. This species is associated witha wide variety of habitats in Singapore, especially in heavily disturbed areas in urban and/or semi-urban settings such as public beaches,roadsides, and forest edges. Nests were typically found in soil or under stones, also inrotten wood, and in dead tree stumps filledwith ‘humus’ or other debris. In one instance,a colony was observed underneath a decaying
fallen tree, ‘partly in the tree’ and ‘partly in(sic) the ground.’Remarks. Type locality in Singapore. Substantial
numbers of specimens of this species were observed amongst existing unsorted and uncatalogued museum material. Thus it could be inferred that the species is much more common
than was represented by official catalogued
records of the ZRC, at the time of writing. It
is known as a globally widespread and potentially invasive species, and could be native toa large part of the Indomalayan bioregion or
the Oceanic region (Sarnat et al. 2015).

Material examined. Sungei Buloh Wetland Reserve, 1.44614, 103.73048, 14 Mar 2018,
M.S. Foo leg., WW-SG18-Phei1, ZRC_
ENT00000901.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Sungei Buloh Wetland Reserve
Habitat/Ecology. This species was found nesting in fallen rotting wood in mangrove backforest.
Remarks. A few hours after collection, majors
and minors of this species were observed toeach possess a pair of conspicuous opaquebright yellow spots/patches at base of gaster(first gastral tergite), spots dissipated 2 days
after collection — possibly midgut or crop
contents, fatty mass. The major worker confers most with P. magrettii sensu lato, withlateral face of occipital lobe and gena mostly
smooth and shiny.

Pheidole hortensis Forel, 1913
Material examined. Bukit Timah Nature Reserve, BT08, 1.35435, 103.78225, 14 Dec
2016, W. Wang leg., WW-SG16-004, ZRC_
HYM_0000294.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve
Habitat/Ecology. A colony was found in soil neartree roots in mature secondary forest.
Remarks. The minor workers of P. hortensis arealmost indistinguishable from those of P. clypeocornis. Species identity was therefore only
confirmed for material where major workers
were available.

Pheidole megacephala (Fabricius, 1793)
Material examined. Bukit Batok East Avenue 6,
1.34304, 103.76235, 30 Aug 2016, G.W. Yongleg., BB4GH101, ZRC_HYM_0001776;
Bukit Timah forest, Oct 1968, D.H. Murphyleg., ZRC_HYM_0000739; University campus (Bukit Timah), 23 Jun 1968, D.H. Murphyleg., ZRC_HYM_0000738.
Material not physically examined. In Wetterer (2012) – site unknown, 1879, F. Smith(BMNH). Non-types, ANIC32-022899(ANIC). Non-types, J.K. Wetterer leg.
(USNM) – Bishan, by MRT station, 31 Jul
2014, vial #424; Harbourfront, 4 Aug 2014,
vial #469; Little India, 27 Jul 2014, vial #330;
Mountbatten, by MRT station, 4 Aug 2014,
vial #465; Promenade, by MRT station, 4 Aug
2014, vial #466.
Literature. Forel (1901); Wetterer (2007, 2012);
Tan & Corlett (2012).
Localities. Bishan; Bukit Batok East; Bukit Timah
forest; Harbourfront; Little India; Mountbatten; University campus (Bukit Timah); National University of Singapore campus (Clementi/Kent Ridge); Promenade.

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Habitat/Ecology. This species was found mostly
in disturbed young secondary forest fragments
in urban or semi-urban settings in Singapore,including waste woodlands and abandoned
plantation forests, also along exposed forest fringes. Sometimes, ant trails were also
found within urban man-made infrastructuresuch as concrete buildings, typically close toadjoining patches of disturbed spontaneousvegetation.
Remarks. Commonly known as the ‘African bigheaded ant’, P. megacephala is a recognizedglobally widespread and particularly aggressive invasive species. It has been reported tooften have severe negative impacts on native
invertebrate fauna where it occurs (Wetterer
2012a). The native range of P. megacephalais presumably the Afrotropical region (Wetterer 2015),
Pheidole nodgii Forel, 1905
Material examined. None available.
Material not physically examined. Type males– FOCOL1362-1364 (ZMHB).
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah Road
Habitat/Ecology. Colonies of this species were
found in a garden, under a flower-pot and ona felled tree on the ground respectively.
Remarks. The male of P. nodgii was first described based on specimens collected fromSingapore (see Viehmeyer 1916).
Pheidole nodgii verlatenensis Wheeler, 1937
Material examined. Bukit Timah Nature Reserve,
1.35284, 103.77995, 11 Jan 2017, W. Wangleg., WW-SG17-008, ZRC_HYM_0000579;
Mandai Road, 1.41258, 103.79839, 21 Dec
2016, G.W. Yong leg., M1D20GH2080, ZRC_
HYM_0001531.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Mandai
Road.
Habitat/Ecology. This subspecies was found in
old or mature native-dominated secondary
forest in Singapore. A colony was found insoil near rotting wood.

Remarks. Specimens available at the time of writing could not be definitely and confidently
identified to species based on morphology,
in the absence of type images or material forcomparison. For the purposes of this checklist,
we tentatively treat these specimens as Pheidole nodgii verlatenensis sensu lato, with the
caveat that this record may change in future if
more information or material becomes available for a more accurate species diagnosis.
Pheidole parva Mayr, 1865
Material examined. Bukit Timah Nature Reserve, along path, 22 Sep 2016, W. Wang leg.,
ZRC_ENT00000894; Kranji Road, 1.42638,
103.75413, 8-10 Nov 2016, G.W. Yong leg.,
KR9GPa1492, ZRC_HYM_0001774; males
and workers, Prince George’s Park Residences (NUS), 1.292389, 103.774583, AprAug 2015, M.S. Foo & W. Wang leg., malaise trap, ZRC_BDP (multiple); males andworkers, University Town (NUS), 1.30622,
103.77458, Apr-Sep 2015, M.S. Foo & W.
Wang leg., malaise trap, ZRC_BDP (multiple); male, Pulau Semakau New Fragment,
SMN1, 1°12’03.1”N, 103°45’45.4”E, 25 Oct1 Nov 2012, J. Puniamoorthy et al. leg., malaise trap, Reg. 29547, ZRC_BDP0016131;
alate queen, Pulau Semakau Old Fragment,
SMO3, 1°12’23.9”N, 103°45’37.6”E, 13-20
Aug 2012, J. Puniamoorthy et al. leg., malaise
trap, Reg. 29435 ZRC_BDP0016540.
Material not physically examined. Unknown.
Literature. Man & Lee (2012); Tan & Corlett(2012); Wang et al. (2018a).
Localities. Bukit Timah Nature Reserve; Kranji
Road; National University of Singapore campus (Clementi/Kent Ridge, multiple sites);
Pulau Semakau.
Habitat/Ecology. This tiny and inconspicuous
species was found in a wide variety of habitats, mostly highly disturbed areas including
young waste woodland secondary forest fragments and cultivated parks/gardens in urban
or semi-urban settings. These ants have also
been reported in mangroves, open grasslands,
trailside along forest edges, and even on impervious surfaces in residential areas andother man-made infrastructure (see Tan &Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Corlett 2012). Individuals were often foundin soil and leaf litter, in rotting wood, understones or root mats.
Remarks. Pheidole parva is a relatively widespread species presumed native to the Indomalayan bioregion and introduced outside
thereof (Sarnat et al. 2015). It appears welladapted to human-dominated environments,
and is considered invasive in parts of the Old
World (Fischer & Fisher 2013). The species
has been suggested to have potential to becoming indoor nuisance pests where they occur in large numbers (Man & Lee 2012; Sarnat
et al. 2015)
Pheidole plagiaria Smith, 1860
Material examined. Bukit Batok East Avenue 6,
1.34304, 103.76235, 5 Oct 2016, G.W. Yongleg., BB6GH412, ZRC_HYM_0001778;
Bukit Timah, Hindhede Drive, 25 Nov 1989,
H.K. Lua leg., ZRC_ENT00000247; Bukit
Timah Nature Reserve, 1.35284, 103.77995,
11 Jan 2017, W. Wang leg., WW-SG17-006,
ZRC_HYM_0000574; Lorong Banir, 16 Jun
1995, Yang & Lua leg., ZRC_ENT00000377;
Mandai Lake Road, 5 Nov 2016, G.W. Yongleg., ZRC_ENT00048426; Mandai Road,
1.41258, 103.79839, 21 Dec 2016, G.W. Yongleg., M1D3GH2059, ZRC_HYM_0001781;
I-cube building (NUS), 29 Jul-5 Aug 2015,
M.S. Foo & W. Wang leg., malaise trap, ZRC_
BDP0042659; Pulau Ubin, PU1, 1°24’36.3”N,
103°59’25.5”E, 17-24 Nov 2012, J. Puniamoorthy et al. leg., malaise trap, Reg. 29592,
ZRC_BDP0015318; same locality and collectors as previous, 12-19 Oct 2013, malaise trap,
Reg. 30212, ZRC_BDP0016192.
Material not physically examined. Unknown.
Literature. Tan & Corlett (2012); Wang et al.
(2018a).
Localities. Bukit Batok; Bukit Timah Nature
Reserve; Lorong Banir; Mandai Lake Road;
Mandai Road; National University of Singapore campus (Clementi/Kent Ridge); Pulau
Ubin.

Habitat/Ecology. This species was found in a
broad range of habitats spanning across different levels of the disturbance spectrum, including primary and mature secondary forests,
young secondary forest fragments in semiurban settings, mangrove back forests, scrub
and open grasslands. Sometimes, the ants
were even observed foraging on impervious
surfaces in man-made infrastructure. Nests
were found in rotting wood, also in soil atbases of large trees.
Pheidole plinii Forel, 1911
Material examined. Mandai Road, 1.41155,
103.79015, 21-23 Nov 2016, G.W. Yongleg., pitfall trap, M2D2GPd1956, ZRC_
HYM_0001789.
Material not physically examined. Syntype – CASENT0904258, ANTC24164(MSNG). Lectotype and paralectotype –CASENT0907938-0907939, ANTC2906929070 (MHNG).
Literature. Type – Forel (1911). Eguchi (2001)[designation of lectotype, paralectotypes].
Localities. Mandai Road
Habitat/Ecology. While no detailed habitat information was available for the type series,
more recently-collected material were mainly
from old or mature native-dominated secondary forests.
Remarks. Type locality in Singapore.
Pheidole rinae Emery, 1900
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah Road
Habitat/Ecology. Individuals of this species were
found in a garden, some on fallen Jambu (Syzygium samarangense) fruits on the ground.
Remarks. At the time of writing, this species isonly known from Singapore based on past
literature records.

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Pheidole rufigera Eguchi, 2001

Pheidole singaporensis Özdikmen, 2010Material examined. Bukit Batok East Avenue 6,
1.34304, 103.76235, 27-29 Sep 2016, G.W.
Yong leg., pitfall trap, BB9GPc472, ZRC_
HYM_0001788.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Batok
Habitat/Ecology. Individuals of this species werecollected from an abandoned plantation secondary forest fragment in semi-urban settings.

Material examined. Bukit Timah Nature Reserve, 15 Aug 2011, J. Koh leg., Winkler
extraction, ZRC_ENT00000893; same locality as previous, 2011, M.S. Foo leg.,
ZRC_ENT00007794-7796; same locality as previous, 1.35386, 103.77988, 4 Jan
2017, W. Wang leg., WW-SG17-002, ZRC_
HYM_0000565; same locality as previous,
28 Mar, collection year unknown, D.H.
Murphy leg., ZRC_HYM_0000740; Lower
Peirce Reservoir, Aug 1990, collectors unknown, ZRC_ENT (multiple); same locality
as previous, 7 May 2017, G.W. Yong leg.,
ZRC_ENT00048417; Mandai Road, 1.41258,
103.79839, ZRC_HYM_0001538-1543;
Mandai Lake Road, 5 Nov 2016, G.W. Yongleg., ZRC_ENT00048425; Upper Peirce Reservoir, 10 Jan 2014, Sk. Yamane leg., SG14SKY-48, ZRC_HYM_0000367.
Material not physically examined. Types –CASENT0901398, ANTC20921 (OUMNH);
CASENT0901505, ANTC21025 (BMNH) [assyntypes of Myrmica longipes].
Literature. Type – Smith (1857) [as Myrmica
longipes]. Eguchi (1999, 2001, 2003), Tan &Corlett (2012) [all as Pheidole longipes].
Localities. Bukit Timah Nature Reserve; Lower
and Upper Peirce Reservoir; Mandai Lake
Road; Mandai Road.
Habitat/Ecology. This species is associated mostly with primary and old/mature secondary forests in Singapore, including native-dominated secondary forests. Nests were frequently
found under fallen logs, or in soil at bases ofliving trees or dead tree stumps. Individual
workers were often found foraging in leaf litter. The ants are often found at high densities
or abundances in and around nesting sites.
Remarks. Type locality in Singapore. Previously
known as ‘Pheidole longipes (Smith, 1857)’,
before being rendered the current replacement
name by Özdikmen (2010) as the original
name was a junior secondary homonym of‘Pheidole longipes (Latreille, 1802)’.

Pheidole sexspinosa Mayr, 1870
Material examined. Kranji mangroves, 9 May
1985, D.H. Murphy leg., DHM-SG85-Pheiss1, ZRC_ENT00000768; Lim Chu Kang
mangrove, 10 Mar 1987, D.H. Murphy leg.,
DHM-SG87-Phei-ss1, ZRC_ENT00000769;
same locality as previous, 27 Feb 2002,
BL1101 Helpers leg., ZRC_ENT00007572;
queen and workers, Mandai mangroves, 27
Oct 1977, D.H. Murphy leg., DHM-SG77Phei-ss2, ZRC_ENT00000770; same locality
and collector as previous, 18 Oct 1977, DHMSG77-Phei-ss3, ZRC_ENT00000772; same
locality and collector as previous, 2 Oct 1978,
ZRC_ENT00027931; queen and workers, same
locality as previous, 8 May 2018, W. Wangleg., WW-SG18-Phei4, ZRC_ENT00007290;
Sungei Buloh Wetland Reserve, 1.4462,
103.73058, 14 Mar 2018, W. Wang leg., WWSG18-Phei3, ZRC_ENT00000908-909; same
locality as previous, 1.4472, 103.72537, 20 Dec
2018, W. Wang & M.S. Foo leg., WW-SG18Phei5, ZRC_ENT00007644.
Material not physically examined. Unknown.
Literature. Wang et al. (2018b).
Localities. Kranji mangroves; Lim Chu Kang
mangrove; Mandai mangroves; Sungei Buloh
Wetland Reserve.
Habitat/Ecology. This species is associated exclusively with mangrove habitats in Singapore.
Nests can be found in abandoned Thalassina(a.k.a mud lobsters) mounds — sometimes
shared with other invertebrates including themangrove trap-jaw ant Odontomachus litoralis, cable roots of mangrove trees such as
Excoecaria agallocha L., or in decayed orliving stems of Rhizophora above tide inundation levels.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Pheidole singaporensis conicollis (Emery, 1900)
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[both as Pheidole (Ischnomyrmex) longipes
var. conicicollis (sic)].
Localities. Bukit Timah Hill
Habitat/Ecology. This subspecies was found ineither primary or old secondary forest, in a rotten, half-decayed tree branch on the ground.
Remarks. The subspecies conicollis may be distinguished from P. singaporensis by a slightly
wider head, and relatively shorter/less elongate occipital collar or ‘neck’; other more
subtle differences include relatively shorter
propodeal spines and a weakly pointed petiolar apex (as opposed to an almost flat apicodorsal petiolar face in P. singaporensis) At thetime of writing, this subspecies is known fromSingapore only based on literature records.
Pheidole singaporensis continentis (Forel, 1911)
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[both as Pheidole (Ischnomyrmex) longipes
var. continentis].
Localities. Bukit Timah Hill
Habitat/Ecology. This subspecies was found ineither primary or old secondary forest, in arotten tree branch on the ground.
Remarks. Said to be of intermediate form between
P. singaporensis and its other subspecies conicollis in Forel’s (1911) original description, the subspecies continentis is in general
smaller and lighter-coloured than P. singaporensis. At the time of writing, this subspecies is known from Singapore only based onliterature records.
Pheidole tandjongensis Forel, 1913
Material examined. Mandai Road, 1.41155,
103.79015, 21-23 Nov 2016, G.W. Yongleg., pitfall trap, M2D2GPd1957, ZRC_
HYM_0001785.2; same locality and collector as previous, 19 Dec 2016, G.W. Yongleg., ZRC_HYM_0001033; Upper Thomson

Nature Park, 1.38311, 103.79839, 17-19 Oct
2016, G.W. Yong leg., pitfall trap, UT5GPd1634, ZRC_HYM_0001785.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Mandai Road; Upper Thomson Nature
Park.
Habitat/Ecology. This species was found in bothold/mature and young secondary forests in
Singapore. Specimens have been collectedusing pitfall traps from native-dominated andabandoned plantation secondary forests.
Pheidole tjibodana Forel, 1905
Material examined. Mandai Road, 1.41119,
103.80513, 29 Nov-1 Dec 2016, G.W. Yongleg., pitfall trap, M3D6GPa890, ZRC_
HYM_0001784; same locality and collector
as previous, 1.41258, 103.79839, 6-8 Dec
2016, G.W. Yong leg., pitfall trap, M1D6GPa2312, ZRC_HYM_0001784.2.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Mandai Road
Habitat/Ecology. This species was found in old/
mature native-dominated secondary forest in
Singapore. Individuals were observed in or
under leaf litter.
Pheidole umbonata Mayr, 1870
Material examined. Pulau Semakau Old Fragment, SMO2, 1°12'20.2"N, 103°45'35.8"E,
16-23 Aug 2012, J. Puniamoorthy et al. leg.,malaise trap, Reg. 29421, ZRC_BDP0016529;
same locality and collectors as previous,
SMO3, 1°12’23.9”N, 103°45’37.6”E, 11-18
Oct 2012, malaise trap, Reg. 29526, ZRC_
BDP0014518; Pulau Ubin, PU3, 1-8 Sep
2012, J. Puniamoorthy et al. leg., malaise
trap, Reg. 29451, ZRC_BDP0016449; Sungei
Buloh Wetland Reserve, SB1, 1°26’46.3”N,
103°43’49.9”E, 17-24 Oct 2012, J. Puniamoorthy et al. leg., malaise trap, Reg. 29530,
ZRC_BDP0017348; same locality and collectors as previous, 8-15 Aug 2012, malaise trap,
Reg. 29400, ZRC_BDP0017287; same locality and collectors as previous, 24-31 Oct 2012,
malaise trap, Reg. 29543, ZRC_BDP0017276.

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Material not physically examined. Unknown.
Literature. None. New record.
Localities. Pulau Semakau; Pulau Ubin; Sungei
Buloh Wetland Reserve.
Habitat/Ecology. Male alates were collected frommangroves in Singapore.
Remarks. Specimens available for examination
were all winged males collected using malaisetraps, thus were not taxonomically verifiableby morphology, which requires other castes
especially major workers. The males from theZRC were identified to P. umbonata solely
based on COI barcodes (313 bp), which werematched against sequences of known species
on GenBank using BLAST. We cannot completely rule out the possibility of misidentification of source organisms to gene sequenceson GenBank. This record will need to be revised should more material be made available
in future to disprove this tentative COI-based
species diagnosis.

as previous, 1.41343, 103.79386, 21-23 Nov
2016, G.W. Yong et al. leg., M1D16TPc1996,
ZRC_ENT00047917; Pulau Ubin, 4 Dec 2016,
G.W. Yong leg., ZRC_ENT00027981; Upper
Thomson Nature Park, 1.38311, 103.79839,
19 Oct 2016, G.W. Yong leg., UT2GH1620,
ZRC_HYM_0001705.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Kent
Ridge Park; Mandai Lake Road; Mandai
Road; Pulau Ubin; Upper Thomson Nature
Park.
Habitat/Ecology. This species was mainly foundin disturbed secondary forests across a rangeof ages (young – old), including abandoned
plantation and native-dominated secondary
forests. Individuals were typically collectedfrom soil and root substrate matrices at basesof trees, rotting fallen tree trunks, and sometimes on tree trunks.

Pristomyrmex bicolor Emery, 1900

Pristomyrmex costatus Wang, 2003

Material examined. Nee Soon Swamp Forest,
17 Dec 1987, D.H. Murphy leg., ZRC_
ENT00000913; same locality and collector
as previous, near Nageia track, 26 Oct 1995,
ZRC_ENT00007573.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Nee Soon Swamp Forest
Habitat/Ecology. Found in swamp forest, in one
instance on a fallen tree trunk.

Material examined. Mandai, 1.40144, 103.77702,
7 Apr 2016, M.K.L. Wong leg., Winkler
extraction, ZRC_ENT00000696; Prince
George’s Park Residences (NUS), 1.29239,
103.7787, 3 May 2017, W. Wang leg., ZRC_
HYM_0001171; same locality as previous, 6
Dec 2017, Sk. Yamane leg. (SKYC); Seletar
Reservoir, 16 May 1994, collector unknown,
NS 134B, ZRC_ENT00000266; Upper
Thomson Road, 2 Oct 2016, G.W. Yong leg.,
ZRC_HYM_0001046; Western Catchment,
1.40866, 103.70185, 20 Nov 2015, M.K.L.
Wong leg., ZRC_ENT00000695.
Material not physically examined. In Wang
(2003) but specimen repositories not specified – Nee Soon Swamp forest, R.W. Taylor
leg., collection date unavailable; Bukit Timah
Nature Reserve, D.H. Murphy leg., collection
date unavailable.
Literature. Wang (2003).
Localities. Bukit Timah Nature Reserve; Mandai; Nee Soon Swamp Forest; Prince George’s
Park Residences (NUS); Seletar Reservoir;
Upper Thomson Road; Western Catchment.

Pristomyrmex brevispinosus Emery, 1887
Material examined. Bukit Timah Nature Reserve, 1.35127, 103.78161, 28 Jun 2017, W.
Wang leg., ZRC_HYM_0001733; Kent Ridge
Park, YM site 2, 7 Dec 2017, W. Wang leg.,
ZRC_ENT00027982; workers, alate queen
and male, same locality and collection dateas previous, F. Ito leg. (SKYC); Mandai
Lake Road, 5 Nov 2016, G.W. Yong leg.,
ZRC_ENT00048421; Mandai Road, 1.41258,
103.79839, 21-23 Nov 2016, G.W. Yong leg.,
ZRC_HYM_0001706-1707; same locality

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Habitat/Ecology. This species is associated mostly with old/mature secondary forests in Singapore, including native-dominated forest andswamp forest. It was sometimes also found inyoung abandoned plantation secondary forest
fragments. Levels of disturbance vary widely
between forest sites. Individuals of this species were usually found in leaf litter, sometimes on tree branches. Nests were collectedfrom rotten logs.
Pristomyrmex picteti Emery, 1893
Material examined. None available.
Material not physically examined. In Wang
(2003) but specimen repositories not specified – locality and collection date unavailable,
Baker coll.
Literature. Wang (2003).
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore context.
Pristomyrmex punctatus (Smith, 1860)
Material examined. Bukit Batok East Avenue 6,
1.34304, 103.76235, 1 Sep 2016, G.W. Yongleg., ZRC_HM_0001704; Bukit Batok East,
1.34218,103.76316, Aug-Sep 2016, G.W.
Yong et al. leg., ZRC_ENT00047915-47916;
Upper Thomson Road, 2 Oct 2016, G.W. Yongleg., ZRC_ENT00000981.
Material not physically examined. In Wang
(2003) but specimen repositories not specified – locality and collection date unavailable,
H. Overbeck coll.
Literature. Viehmeyer (1916), Overbeck (1924)[both as ‘Prystomyrmex (sic) pungens]. Wang
(2003).
Localities. Bukit Batok East; Bukit Timah Road;
Upper Thomson Road.
Habitat/Ecology. This species is associated mostly with young secondary forests in Singapore,including abandoned plantation secondary
forest, and has also been found in gardens.
These ants are known to frequently occur inopen and disturbed habitats, and were usually found nesting in soil, leaf litter, or rottingwood.

Proatta butteli Forel, 1912
Material examined. Bukit Timah Nature Reserve, 8 Dec 1995, H.K. Lua et al. leg.,
NS 208A, ZRC_ENT00000362; same locality as previous, 6 Jul 2015, J. Koh leg.,
ZRC_ENT00000890; Lower Peirce Reservoir Road, 28 Jul 1990, H.K. Lua leg., LHK
108, ZRC_ENT00000403; MacRitchie Reservoir, 9 Jan 2014, Sk. Yamane leg., ZRC_
HYM_0000425-426; Mandai Road, 1.41258,
103.79839, 16-18 Nov 2016, G.W. Yongleg., ZRC_HYM_0001629-1632; Nee Soon
forest, 16 Aug 2017, W. Wang leg., ZRC_
ENT00047966; male, Prince George’s Park
Residences (NUS), 1.29239, 103.77869, 29
Apr-6 May 2015, M.S. Foo & W. Wang leg.,malaise trap, NUS0021, ZRC_BDP0041414;
Pulau Ubin, 1.40822, 103.97051, 15 Oct 2016,
G.W. Yong et al. leg., ZRC_HYM_0001074;
same locality as previous, 1.41662, 103.99443,
4 Dec 2016, G.W. Yong & M.K.L. Wongleg., ZRC_HYM_0001075; Seletar Reservoir Park, 17 Mar 1998, C.M. Yang leg.,
ZRC_HYM_0000267; Upper Thomson Nature Park, 1.38311, 103.79839, Sep-Oct 2016,
G.W. Yong leg., ZRC_HYM_0001626-1628.
Material not physically examined. Non-types,
CASENT0178527, ANTC8736 (MCZ); Harbourfront, Marang Trail, 4 Aug 2014, vial
#472 (JKWC).
Literature. Viehmeyer (1916); Overbeck (1924);
Weber (1958); Moffett (1986); Wheeler &Wheeler (1986); Tan & Corlett (2012).
Localities. Singapore Botanic Gardens; Bukit
Timah Nature Reserve; Bukit Timah Road;
Harbourfront; Lower Peirce Reservoir Road;
MacRitchie Reservoir; Mandai Road; Nee
Soon; Prince George’s Park Residence (NUS);
Pulau Ubin; Seletar Reservoir Park; Upper
Thomson Nature Park.
Habitat/Ecology. This species was found mainlyin old/mature or young secondary forests in
Singapore, including abandoned plantationand native-dominated secondary forests, andsometimes in semi-urban parklands or gardens. These ants were collected from bamboo
shoots, soil, leaf litter, even decaying logs.

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Rhopalomastix glabricephala Wang, Yong &Jaitrong, 2018Material examined. Holotype – Mandai Road,
1.41379, 103.7982, 16 Nov 2016, G.W. Yong
& K.H. Pwa leg., GY-SG16-RhoE, ZRC_
ENT00000873; paratypes – same collection
data as holotype, ZRC_HYM_0000508.
Material not physically examined. Unknown.
Literature. Type – Wang et al. (2018c).
Localities. Mandai Road [type]
Habitat/Ecology. The type series was found inmature native-dominated secondary forest,
where the ants were nesting in bark of a
Tembusu tree. As with other congeners, thisspecies lives in apparent mutualistic association with scale insects (Diaspididae). The
ants consume the waxy protein-rich cuticularsecretions of the diaspidids, in exchange thelatter gets protection from external elements
or enemies.
Remarks. Type locality in Singapore. At the timeof writing, this species is known from Singapore only by the type series collected from asingle locality.
Rhopalomastix javana Wheeler, 1929
Material examined. Bukit Timah Nature Reserve,
1.35449, 103.78211, 21 Jun 2017, C. Peetersleg., WW-SG17-015, ZRC_HYM_0001732;
Mandai forest, 1.40931, 103.78334, 13 Nov
2018, W. Wang leg., WW-SG18-Rho3, ZRC_
ENT00007584; Mandai Track, 4 Jan 2017,
G.W. Yong leg., GY-SG17-RhoN, ZRC_
HYM_0000576.
Material not physically examined. Unknown.
Literature. Wang et al. (2018c); Wang et al.
(2021).
Localities. Bukit Timah Nature Reserve; Mandai
forest; Mandai Track.
Habitat/Ecology. Found mostly in old or maturesecondary forests in Singapore, including
native-dominated secondary forest. The ants
were found nesting in bark of large dipterocarp trees, such as Campnosperma auriculatum (locally-known as ‘Terentang’).

Rhopalomastix johorensis Wheeler, 1929
Material examined. Syntypes of ‘R. rothneyi
subsp. johorensis’ – no specific locality given, MCZ-ENT00023066 (MCZ). Non-types– Bukit Timah Nature Reserve, 1.35127,
103.78161, 1 Nov 2017, W. Wang leg., WWSG17-022, ZRC_HYM_0001794; Hougang
Avenue 3, 1.34769, 103.88838, 12 Dec 2016,
G.W. Yong & S.X. Chui leg., GY-SG16RhoL, ZRC_HYM_0000511; male, I-cubebuilding (NUS), 1.293472, 103.77633, 24
Mar-1 Apr 2015, M.S. Foo & W. Wang leg.,malaise trap, NUS0002, ZRC_BDP0044708;
Lee Kong Chian Natural History Museum(NUS), side yard, 24 Aug 2018, G.W. Yongleg., WW-SG18-Rho1, ZRC_ENT00007463;
queens and workers, Mandai Road, 1.41445,
103.79831, 4 Oct 2018, G.W. Yong & W.
Wang leg., WW-SG18-RhoMA, ZRC_
ENT00007519; males and workers, Mandai
Road, 1.41333, 103.79839, 18 Nov 2016,
G.W. Yong & S.X. Chui leg., GY-SG16-RhoF,
ZRC_HYM_0000289; queen and workers,
Mandai Road, 1.41318, 103.79405, 21 Nov
2016, G.W. Yong and S.X. Chui leg., GYSG16-RhoH, ZRC_HYM_0000291; same
locality as previous, 13 Jan 2017, G.W. Yongleg., GY-SG17-RhoO, ZRC_HYM_0000577;
queens and workers, Mandai forest, near Zoo,
15 Jan 2020, W. Wang leg., WW-SG20-005– 006, ZRC_ENT00027891-27892; NeeSoon Swamp Forest, 24 Sep 2019, W. Wangleg., WW-SG19-005, ZRC_ENT00013876;
queen and workers, Pulau Tekukor, 1.23081,
103.83774, 15 Jan 2017, G.W. Yong leg.,
GY-SG17-RhoP, ZRC_HYM_0000578;
male, Pulau Ubin, PU1, 1°24’36.3”N,
103°59’25.5”E, 1-8 Sep 2012, J. Puniamoorthy et al. leg., mangroves, malaise trap, Reg.
29449, ZRC_BDP0014267; male, Pulau Semakau Old Fragment, SMO3, 1°12’23.9”N,
103°45’37.6”E, 17-25 Oct 2012, J. Puniamoorthy et al. leg., mangroves, malaise trap,
Reg. 29539, ZRC_BDP0014651; queens andworkers, Upper Seletar Reservoir Park, 14
Feb 2018, G.W. Yong leg., GY-SG18-Rho1,
ZRC_ENT00000897.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Material not physically examined.
CASENT0911229, ANTC33476 (MHNG)[headless worker].
Literature. Viehmeyer (1916), Overbeck (1924)[both as Rhopalomastix rothneyi Forel, 1900 –verified as R. rothneyi johorensis by Wheeler(1929)]. Wheeler (1929) [as R. rothneyi johorensis]. Wang et al. (2018c); Wang et al.
(2021).
Localities. Bukit Timah Nature Reserve; Bukit
Timah Road; Hougang; National Universityof Singapore campus (Clementi/Kent Ridge,
multiple sites); Mandai Road; Mandai forest
near Singapore Zoo; Nee Soon Swamp Forest;
Pulau Tekukor; Pulau Semakau; Pulau Ubin;
Upper Seletar Reservoir Park.
Habitat/Ecology. Rhopalomastix johorensis appears ubiquitious locally and could be foundin a broad range of different habitat types,
including urban or semi-urban habitats (suchas roadside vegetation), parks and gardens,
old/mature and native-dominated secondary
forests, and swamp forests. The ants nest in
bark of various trees, both native species such
as: Aquilaria malaccensis, Artocarpus integer (cempedak), Campnosperma auriculatum,
Macaranga gigantea, Horsfieldia sparsa, andalso introduced tree species such as Mangifera
indica (mango). Nests have also been found indead ends of twigs of cultivated mangosteen
trees, and in bark of durian trees. Multiple
nests (one nest per tree) can occur at relatively
high densities per site where the species occurs — these are probably independent colonies and not satellites of a larger super-colony.
The species appears tolerant to high levelsof habitat disturbance. Males were collectedfrom malaise traps set up in mangroves.
Remarks. Type locality in Singapore. As withmany congeners, this species is known to
exhibit very broad intranidal, generally isometric size variation.

Rhopalomastix murphyi Wang, Yong & Jaitrong,
2018
Material examined. Holotype – Upper Thomson
Nature Park, 1.38653, 103.81938, 24 Oct 2016,
G.W. Yong & S.X. Chui leg., durian tree, GYSG16-RhoA, ZRC_ENT00000874; paratypemales, queens and workers, same collection
data as holotype, ZRC_HYM_0000287. Nontypes – queens and workers, Bukit Timah Nature Reserve, 2 Jan 1981, D.H. Murphy leg.,
DHM-SG81-Rho1, ZRC_HYM_0000009;
Mandai Lake Road, 1.40915, 103.78234,
3 Nov 2016, G.W. Yong & C. Peeters leg.,
GY-SG16-RhoD, ZRC_HYM_0000507;
males and workers, Mandai Road, 1.4144,
103.79845, 4 Oct 2018, G.W. Yong & W. Wangleg., WW-SG18-Rho2, ZRC_ENT00007518;
Upper Thomson Road, 29 Oct 2016, G.W.
Yong leg., ZRC_HYM_0001008-1014.
Material not physically examined. Unknown.
Literature. Type – Wang et al. (2018c). Yong etal. (2019).
Localities. Bukit Timah Nature Reserve; Mandai
Lake Road; Mandai Road; Upper Thomson
Nature Park [type]; Upper Thomson Road.
Habitat/Ecology. This minute, yellowish-brownspecies was found mostly in old/mature or
young secondary forests, including abandoned plantation forest. Colonies could befound nesting in bark of live durian trees orother large dipterocarps, e.g. Myristicaceae,
where nests were often observed to be rather
extensive, sprawled across an indeterminate
expanse of the host tree trunk. Colonies wereobserved to be polygynous, i.e., each colony
houses multiple queens. The ants also livewith diaspidid scale insects in their nests, withwhich they share a seeming obligate relationship. The diaspidids allow the ants to consume
their waxy outer scale secretions in exchange
for apparent protection from predators. Silk
spinning behaviour has been documented inthis species (Yong et al. 2019), where the ants
produce silk from head glands in order to secure nest tunnels.
Remarks. Type locality in Singapore.

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Rhopalomastix striata Wang, Yong & Jaitrong,
2018
Material examined. Holotype – Mandai Road,
1.4144, 103.79845, 18 Nov 2016, G.W.
Yong & S.X. Chui leg., GY-SG16-RhoG,
ZRC_ENT00000876; paratype males andworkers, same collection data as holotype,
ZRC_HYM_0000290. Non-types – queens
and workers, Bukit Timah Nature Reserve, BT05, 1.35386, 103.77988, 30 Aug
2017, W. Wang leg., WW-SG17-Rho1,
ZRC_HYM_0001752; queen and workers,
same locality as previous, BT03, 1.34904,
103.77892, 27 Sep 2017, W. Wang leg., WWSG17-021, ZRC_HYM_0001793; same locality as previous, BT02, 1.35599, 103.77397,
12 Jul 2017, W. Wang leg., WW-SG17-RhoR,
ZRC_ENT00027915; queens and workers,
Mandai forest, near Singapore Zoo, 15 Jan
2020, W. Wang leg., WW-SG20-004, ZRC_
ENT00027890; Mandai Road, 1.40824,
103.802, 21 Dec 2016, G.W. Yong leg.,
GY-SG16-RhoM, ZRC_HYM_0000512;
queen and workers, Rifle Range Road, 6 Dec
2017, W. Wang leg., WW-SG17-046, ZRC_
ENT00027893.
Material not physically examined. Unknown.
Literature. Type – Wang et al. (2018c).
Localities. Bukit Timah Nature Reserve; Mandai forest; Mandai Road [type]; Rifle Range
Road.
Habitat/Ecology. This species is associatedmostly with primary and old/mature secondary forests in Singapore. Nests were found inbark of various large native dipterocarp trees,e.g. Endospermum diadenum, Prunus sp., andShorea curtisii. In one instance, the ants were
found nesting in bark of large buttress rootnear tree base.
Remarks. Type locality in Singapore.
Rhopalomastix tenebra Wang, Yong & Jaitrong,
2018
Material examined. Holotype – Upper Thomson Nature Park, 1.38653, 103.81938, 24 Oct
2016, G.W. Yong & S.X. Chui leg., durian
tree, GY-SG16-RhoB, ZRC_ENT00000877;
paratype male, queens and workers,

same collection data as holotype, ZRC_
HYM_0000288. Non-types – Bukit Timah
Nature Reserve, 9 Mar 1983, D.H. Murphyleg., ZRC_HYM_0000686; Mandai forest,
1.40843, 103.78326, 13 Nov 2018, W. Wangleg., WW-SG18-Rho4, ZRC_ENT00007585;
Namly Avenue, 28 Apr 1985, D.H. Murphyleg., DHM-85-Rho1, ZRC_ENT00000885;
Upper Thomson Road, 11 Oct 2016, G.W.
Yong leg., ZRC_HYM_0001015-1022.
Material not physically examined. Unknown.
Literature. Type – Wang et al. (2018c).
Localities. Bukit Timah Nature Reserve; Mandai
forest; Namly Avenue; Upper Thomson Nature Park [type]; Upper Thomson Road.
Habitat/Ecology. This species was found mainlyin old/mature and young secondary forests in
Singapore, including abandoned plantation
forest. Nests were found in bark of large dipterocarp trees such as durian.
Remarks. Type locality in Singapore.
Rostromyrmex pasohensis Rosciszewski, 1994
Material examined. None available.
Material not physically examined.
CASENT0106221 – 106221-D01,
PSW09576-15 (UCDC).
Literature. None. New record.
Localities. Bukit Timah
Habitat/Ecology. Individuals were usually foundin sifted forest litter (comprising leaf moldand rotten wood) from primary or old/maturesecondary forests.
Rotastruma recava Bolton, 1991
Material examined. None available.
Material not physically examined. Paratypes–3 workers, CASENT0178523, ANTC8732(MCZ); 4 queens and 17 workers [BMNH,
MHNG – according to Bolton (1991)].
Non-types – CASENT0010807 – 10808,
PSW10254 (UCDC); 2 queens and 1 male,
CASENT0010926 – 10928, PSW10254(UCDC); CASENT0102499, ANTC4706(MHNG); CASENT0106250 – 106250-D01/
D11, PSW09601 (UCDC).
Literature. Paratype – Bolton (1991).
Localities. Bukit Timah [type]; Peirce Reservoir.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Habitat/Ecology. This species is associated withprimary and old/mature secondary rainforest.
Apparently arboreal, these ants were foundin dead twigs, vines, or dead twigs of vines.
Sometimes, individuals were collected fromsurfaces of tree trunks and/or branches.
Remarks. The secondary type locality for thisspecies is in Singapore, where paratypes
were collected. The holotype specimen wascollected in Mulu National Park in Sarawak(Malaysia), the primary type locality.
Solenopsis geminata (Fabricius, 1804)
Material examined. Kranji mangroves, 24 Sep
1964, D.H. Murphy leg., ZRC_ENT00047993;
Lim Chu Kang, Oct 1969, D.H. Murphy leg.,
DHM-SG69-Sol1, ZRC_ENT00047992; University campus, 17 Dec 1980, D.H. Murphyleg., DHM-SG80-Sol1, ZRC_ENT00047994;
Phillip Street, 24 Jun 2011, ex. sugar container, R. Lohman leg. (SKYC).
Material not physically examined. Unknown.
Literature. Mayr (1897).
Localities. Kranji; Lim Chu Kang; Phillip Street;
University campus (Bukit Timah).
Habitat/Ecology. This species was found in arange of habitat types, including disturbed
secondary forest fragments in semi-urban
settings, mangrove back forests, and even incultivated farmland – where the ants were foraging on Solanum melongena L. (eggplant).
Ant trails were also observed in close proximity to Termes rostratus mounds in disturbed
forest patches.
Remarks. Commonly known as the ‘tropical fireant’, S. geminata is a cosmopolitan pest species believed to have originated from the Neotropics and spread to much of the Old World,
mainly tropical Asia and Oceania, via humancommerce (Wetterer 2011, 2015). Its notoriety as a worldwide invasive has in recent
times been somewhat downplayed in contrast
to the growing infamy of the closely-related
congener S. invicta – the ‘imported fire ant’.
The latter species continues to be the dominant focus of most pest invasions research andpress reports till present.

Solenopsis overbecki Viehmeyer, 1916
Material examined. Mandai forest, 7 Jun 2018,
W. Wang leg., WW-SG18-001, ZRC_
ENT00027959; same locality and collector as previous, 7 Jun 2018, W. Wang leg.,
WW-SG18-003, ZRC_ENT00047997; NeeSoon forest, 26 Apr 2018, W. Wang leg.,
WW-SG18-007, ZRC_ENT00047962; Pulau Ubin, 1.41320, 103.97720, 17 Nov 2021,
W.N. Lam leg., ZRC_ENT00057811; Upper
Peirce Reservoir Park, 1.35229, 103.82450,
21 Oct 2021, same collector as previous,
ZRC_ENT00057812.
Material not physically examined. Types –CASENT0908878, ANTC31310 (MHNG);
3 queens and 3 workers, FOCOL1870 – 1875
(ZMHB).
Literature. Type – Viehmeyer (1916); Overbeck(1924).
Localities. Bukit Timah Road [type]; Mandai;
Nee Soon forest; Pulau Ubin; Upper Peirce
Reservoir.
Habitat/Ecology. The type series was collectedfrom a garden, where the colony was foundin the narrow passages within a broken branch
piece on the ground. The type colony is described to be polygynous with multiple dealate
queens. In more recent times, the species was
also found in old or mature secondary forests,
either in leaf litter, or nesting inside root cavities embedded in soil mounds next to termite
nests. In one instance, the ants were also foundnesting in bark of Rhodamnia cinera.
Remarks. Type locality in Singapore.
Strumigenys chapmani Brown, 1954
Material examined. Bukit Timah Nature Reserve,
8 Mar 2017, W. Wang leg., WW-SG17-032,
ZRC_ENT00001000; queens and workers,
same locality and collector as previous, 22 Feb
2017, WW-SG17-033, ZRC_ENT00007240;
males and workers, same locality and collector as previous, 22 Feb 2017, WW-SG17-034,
ZRC_ENT00007241; same locality and collector as previous, 1.35435, 103.78225, 14 Dec
2016, WW-SG16-002, ZRC_HYM_0000281.

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Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve
Habitat/Ecology. This species is associated withold/mature secondary forests in Singapore.
Nests were observed to occur at rather highdensity (separated only by a few metres fromeach other) within a limited area of forest,
though these could possibly be merely satellite colonies. The ants were usually foundnesting in soil-roots-substrate matrices onbark/trunk surfaces near bases of trees.
Strumigenys eggersi Emery, 1890
Material examined. Queens and workers, Prince
George’s Park Residences (NUS), 1.292389,
103.77869, 5 Apr 2017, W. Wang leg., ZRC_
HYM_0000608; alate queens, same locality
as previous, May-Jun 2015, M.S. Foo & W.
Wang leg., malaise trap, ZRC_BDP (multiple); queens and workers, Pulau Ubin, 22
Oct 2016, W. Wang leg., leaf litter, Winkler
extraction, ZRC_ENT00007274; Sunset Way,1.32608, 103.77187, 21-23 Sep 2016, G.W.
Yong leg., pitfall trap, SW1GPb238, ZRC_
HYM_0001636.
Material not physically examined. Non-types,
J.K. Wetterer leg. (USNM) – East Coast Park
(Katong part), 1 Aug 2014, vial #442; Marine
Parade, 1 Aug 2014, vial #435.
Literature. Wang & Yamane (2017).
Localities. Prince George’s Park Residences(NUS); Pulau Ubin; Sunset Way.
Habitat/Ecology. This species was found in disturbed mature or young secondary forest fragments in semi-urban settings, including abandoned plantation secondary forest. Workers
and queens were collected mainly from leaf
litter or ground debris; multiple alate queens
were also collected via malaise traps.
Remarks. An introduced species native to the
Neotropical region, S. eggersi has been reported to be an unusually resilient species
tolerant of relatively dry environments, andadaptable to a broad range of habitats including highly disturbed forests (Deyrup 1997;
Wang & Yamane 2017).

Strumigenys emmae (Emery, 1890)
Material examined. Queens and males, I-cubebuilding (NUS), 1.293472, 103.77633, JunSep 2015, M.S. Foo & W. Wang leg., malaise trap, ZRC_BDP (multiple); alate queen,
Kranji Road, 1.42638, 103.75413, 8-10 Nov
2016, G.W. Yong leg., ZRC_HYM_0001637;
Labrador beach, collection date unknown,
D.H. Murphy leg., ZRC_HYM_0000780;
Pulau Ubin, 7 Jan 2014, Sk. Yamane leg.,
ZRC_ENT00000970; male, University Town
(NUS), 1.30622, 103.77458, 29 Jul – 5 Aug
2015, M.S. Foo & W. Wang leg., malaise trap,
ZRC_BDP0042899.
Material not physically examined. Non-types,
J.K. Wetterer leg. (USNM) – East Coast Park
(Katong part), 1 Aug 2014, vial #442; Hougang, 30 Jul 2014, vial #402; Kranji, by MRT
station, 2 Aug 2014, vial #452; Pasir Panjang,
by MRT station, 5 Aug 2014, vial #483; Serangoon, 30 Jul 2014, vial #389.
Literature. Szabó (1909), Donisthorpe (1916),
Bolton (2000) [all as Epitritus clypeatus].
Localities. I-cube building (NUS); Kranji Road;
Labrador beach; Pulau Ubin; University Town
(NUS).
Habitat/Ecology. This species was found in asandy beach, also disturbed secondary forest
fragments or open grassy patches in semi-urban settings, including waste woodland forest.
On one occasion, individuals were found on
decayed wood. Multiple alates were collectedfrom secondary and/or open habitats via malaise traps.
Remarks. A recognized and apparently successful pantropical tramp species, S. emmae isdeemed to have originated from Australia(Bolton 2000) or Australasia (Wetterer 2015),
and spread nearly worldwide by human commerce and natural dispersal. The species isknown to frequently thrive in highly disturbed
and anthropogenically-modified habitats suchas beaches and gardens (Wetterer 2012b).

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Strumigenys epyna Bolton, 2000
Material examined. Queens and workers, Mandai Road, 21 Dec 2016, G.W. Yong leg., GYSG16-Strum2, ZRC_ENT00007266.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Mandai Road
Habitat/Ecology. A nest series was found in either
mature or young secondary forest in Singapore.
Strumigenys extemena (Taylor, 1968)
Material examined. Bukit Timah Nature Reserve, 23 Aug 2017, W. Wang leg., ZRC_
ENT00007252; same locality and collector
as previous, 13 Sep 2017, W. Wang leg.,
ZRC_ENT00007253.
Material not physically examined. Holotype– ANIC32-001163, berl.no.B45 (Berlesate)(ANIC).
Literature. Type – Taylor (1968) [as Dysedrognathus extemenus]. Bolton (2000) [as Pyramica
extemena].
Localities. Bukit Timah Nature Reserve [type]
Habitat/Ecology. This species is associated withprimary and old/mature secondary forests in
Singapore. The ants were usually found in soil
at bases of trees.
Remarks. Type locality in Singapore. “Degradedcoastal hill forest on granite” printed on theoriginal type label is probably an inaccurate
description of the actual forest types in Bukit
Timah Nature Reserve (BTNR), based on
known land use histories of BTNR and its
surroundings.
Strumigenys godeffroyi Mayr, 1866
Material examined. Kent Ridge, 13 Feb 1985,
D.H. Murphy leg., ZRC_HYM_0000790;
Mandai forest, at border with Central Catchment Nature Reserve, 1.40933, 103.78339, 13
Nov 2018, W. Wang leg., ZRC_ENT00007587.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924);
Bolton (2000).
Localities. Bukit Timah Road; Kent Ridge; Mandai forest.

Habitat/Ecology. This species was found in agarden inland, nesting in fibrous tissue about
the base of leaves of a coconut palm (‘tree’).
Individuals were also found under brick in asemi-urban setting, vegetation-type unknown.
The ants were also collected from old/maturesecondary forest.
Remarks. Strumigenys godeffroyi is known to
be a tramp species widely distributed acrossislands in the Pacific and Indian Ocean. At
the time of writing, substantial material in the
ZRC were at best identified to the eponymous
godeffroyi species complex, but could not be
reliably verified as S. godeffroyi mostly due
to poor specimen condition such as abradedhairs. Pilosity is an important state used todifferentiate between members of the godeffroyi complex.
Strumigenys juliae Forel, 1905
Material examined. Bukit Timah Nature Reserve,near BT02, 17 Nov 2016, W. Wang leg.,
ZRC_ENT00007283; same locality and collector as previous, near BT07, 28 Sep 2016,
W. Wang leg., ZRC_ENT00007286; queens
and workers, same locality and collector as
previous, Aug-Dec 2016, leaf litter, Winkler
extraction, ZRC_ENT00007287-7288; same
locality and collector as previous, near BT05,
14 Sep 2016, leaf litter, Winkler extraction,
ZRC_ENT00007289; queens and workers,
Mandai, Northern Node, 1.40842, 103.78319,
54m a.s.l., 25 Jul 2019, J.Tan & S. Tang leg.,
JTST-MIS-010, ZRC_ENT00013873.
Material not physically examined. Unknown.
Literature. Bolton (2000).
Localities. Bukit Timah Nature Reserve; Mandai.
Habitat/Ecology. This species was found mostly
in primary and old/mature secondary forests,
sometimes in young secondary forest in Singapore. These ants were collected mainly
from leaf litter; nests were found in fallen
rotting wood.

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Strumigenys koningsbergeri Forel, 1905
Material examined. Bukit Timah Nature Reserve,near BT09, 5 Oct 2016, W. Wang leg., leaf litter, Winkler extraction, ZRC_ENT00007246;
same locality and collector as previous, near
BT03, 11 Aug 2016, ZRC_ENT00007259;
same locality and collector as previous, near
BT07, 28 Sep 2016, leaf litter, Winkler extraction, ZRC_ENT00007260; Lower Peirce
Reservoir, Forest B, 25 Aug 1990, collector
unknown, ZRC_ENT00000104; Nee Soon
forest, 2 Oct 2017, W. Wang leg., WT_FL1/
NS2, ZRC_ENT00007258.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Lower
Peirce Reservoir; Nee Soon forest.
Habitat/Ecology. This species was found mostly
in old/mature secondary forests in Singapore.
Nests were found in fallen rotting wood; individual specimens were often collected fromleaf litter and ground debris via Winkler extraction.
Strumigenys kraepelini Forel, 1905
Material examined. Queen and workers, Bukit
Timah Nature Reserve, near BT09, 5 Oct
2016, W. Wang leg., ZRC_ENT00007245;
queen and workers, same locality and collector as previous, near BT01, 1 Feb 2017,
ZRC_ENT00007272; same locality, site, andcollector as previous, 8 Dec 2016, leaf litter, Winkler extraction, ZRC_ENT00007275;
same locality and collector as previous, near
BT07, 28 Sep 2016, leaf litter, Winkler extraction, ZRC_ENT00007276; same locality
as previous, 6 Jun 1968, D.H. Murphy leg.,D7-4, ZRC_ENT00000795; Nee Soon, 18
Jun 1969, D.H. Murphy leg., D10-7, ZRC_
ENT00000785.
Material not physically examined. Unknown.
Literature. Bolton (2000).
Localities. Bukit Timah Nature Reserve; Nee
Soon.
Habitat/Ecology. This species was found in bothold and mature secondary forests in Singapore. Individuals were usually collected fromleaf litter, occasionally in rotten wood.

Remarks. At the time of writing, this species isknown from only one locality in Singapore.
Strumigenys membranifera Emery, 1869
Material examined. None available.
Material not physically examined. City Hall, by
church, 5 Aug 2014, J.K. Wetterer leg., vial
#488 (USNM).
Literature. None. New record.
Localities. City Hall
Habitat/Ecology. This species was found in soil
or debris at bases of trees near urban infrastructure, i.e., a church in the central business
district.
Remarks. This is a pantropical tramp species(Bolton 2000) with a broad cosmopolitan
distribution (Wetterer 2011b); its presumed
native range is the Afrotropical region (Wetterer 2015).
Strumigenys mitis (Brown, 2000)
Material examined. Queen and workers, Bukit Timah Nature Reserve, near BT07, 28 Sep 2016,
W. Wang leg., ZRC_ENT00007242; same locality and collector as previous, near BT01, 8
Dec 2016, ZRC_ENT00007243; same locality
and collector as previous, near BT09, 5 Oct
2016, ZRC_ENT00007244; queen and workers, Nee Soon forest, 1.38245, 103.80206,
46 m, NS_W2, 26 Apr 2018, W. Wang leg.,
ZRC_ENT00047851.
Material not physically examined. In Bolton(2000) – 1 non-type queen, locality unspecified (MCZ).
Literature. Bolton (2000) [as Pyramica mitis].
Localities. Bukit Timah Nature Reserve; Nee Soon
forest.
Habitat/Ecology. This species was found mainlyin old or mature secondary forest in Singapore, including Adinandra belukar-dominantforest, where individuals were usually collected from leaf litter.
Remarks. Elsewhere in the world, this species
is known to be able to thrive in a wide rangeof habitats including heavily disturbed anddegraded grasslands and plantations (see Tanget al. 2019).

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Strumigenys mutica Brown, 1949
Material examined. None available.
Material not physically examined. Unknown.
Literature. Bolton (2000).
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore context.
Remarks. At the time of writing, this rare species
is only known from Singapore based on a literature record, with no specific locality given.
Strumigenys nanzanensis Lin & Wu, 1996
Material examined. Queen and workers, Bukit
Timah Nature Reserve, near BT02, 2 Aug
2017, W. Wang leg., ZRC_ENT00007271;
Mandai Lake Road, Mandai Bird Park Buffer,
N01°24’19.3’’, E103°46’56.2’’, 73 m a.s.l.,
26 Mar 2020, W. Zhang leg., WZ-MIS-018,
ZRC_ENT00047777; Mandai Road, 21 Dec
2016, G.W. Yong leg., GY-SG16-Strum1,
ZRC_ENT00007267; queen, same locality as
previous, 27 Jun 2019, J. Tan & N. Chin leg.,
ZRC_ENT00013535; Mandai Track, 4 Jan
2017, G.W. Yong leg., ZRC_ENT00007270;
queen and workers, Upper Seletar Reservoir
Park, 14 Feb 2018, G.W. Yong leg., GYSG18-Strum1, ZRC_ENT00007268; Upper
Thomson Road, 2 Oct 2016, G.W. Yong leg.,
ZRC_ENT00007269.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Mandai
Lake Road (Bird Park Buffer); Mandai Road;
Mandai Track; Upper Seletar Reservoir Park;
Upper Thomson Road.
Habitat/Ecology. This species was found mostly
in primary or mature secondary forests ofvarying levels of disturbance in Singapore,as well as young secondary forest such as
abandoned plantation and parkland forest
fragments in semi-urban settings. Individuals
were found in leaf litter, also rotting wood andsoil at bases of trees. On one occasion, someindividuals were found in an Anoplolepis gracilipes nest in a fallen rotting tree branch.

Strumigenys nepalensis Baroni Urbani & De
Andrade, 1994
Material examined. Alate queen, Prince George’s
Park Residences (NUS), 1.292389, 103.77869,
17-24 Jun 2015, M.S. Foo & W. Wang leg.,malaise trap, ZRC_BDP0044435.
Material not physically examined. Unknown.
Literature. Bolton (2000) [as Pyramica nepalensis].
Localities. Prince George’s Park Residences(NUS)
Habitat/Ecology. An alate queen was collectedfrom a malaise trap set up at a disturbed secondary forest fragment in semi-urban settings.
At the time of writing, mounting evidence
supports the apparent tramp nature of thisspecies – it is currently known to be widelydistributed in Asia and beyond (Tang et al.
2019).
Strumigenys rofocala Bolton, 2000
Material examined. Queens and workers, Bukit
Timah Nature Reserve, near BT01, 8 Dec
2016, W. Wang leg., leaf litter, Winkler extraction, ZRC_ENT00007280; same locality and collector as previous, near BT02,
17 Nov 2016, leaf litter, Winkler extraction,
ZRC_ENT00007282; same locality and collector as previous, near BT07, 28 Sep 2016,
ZRC_ENT00007284; queens and workers,
same locality and collector as previous, near
BT06, 23 Aug 2016, leaf litter, Winkler extraction, ZRC_ENT00007285; same locality
and collector as previous, near BT09, 5 Jan
2016, leaf litter, Winkler extraction, ZRC_
ENT00048729; same locality as previous, plot
III-2,12 Oct 2015, D.J. Court leg., pitfall trap,
ZRC_ENT00048445; Pulau Ubin, 22 Oct
2016, W. Wang leg., ZRC_ENT00007277.
Material not physically examined. Unknown.
Literature. Bolton (2000).
Localities. Bukit Timah Nature Reserve; Pulau
Ubin.
Habitat/Ecology. This species was found mainlyin old/mature secondary forests in Singapore.
Individuals were usually collected from leaf
litter.

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Strumigenys roganas Bolton, 2000

Strumigenys sublaminata Brown, 1959

Material examined. None available.
Material not physically examined. Unknown.
Literature. Bolton (2000).
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore context.
Remarks. At the time of writing, this species isonly known from Singapore based on a single
literature record.

Material examined. Bukit Timah Nature Reserve, near BT05, 13 Sep 2017, W. Wang leg.,
WW-SG17-035, ZRC_ENT00007254; same
locality and collector as previous, 1.35386,
103.77988, 1 Nov 2017, W. Wang leg., ZRC_
ENT00047953; queen and workers, same locality and collector as previous, near BT02,
19 Apr 2017, W. Wang leg., WW-SG17-036,
ZRC_ENT00007255; queens and workers,
same locality, collection date and collector
as previous, near BT02, WW-SG17-037,
ZRC_ENT00007256; males and workers,
same locality and collector as previous, near
BT09, 28 Jun 2017, WW-SG17-038, ZRC_
ENT00007257.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve
Habitat/Ecology. This species is associated mostly
with primary forest, and less frequently withold/mature secondary forest in Singapore. In
primary forest, nests were found in tree bark,under concavities in tree bark filled with soil,
and under bark of dead fallen logs. In maturesecondary forest, nests were found under barkof fallen trees or large rotting logs.
Remarks. At the time of writing, S. sublaminata
is only known from one locality in Singapore.
This species is quite similar to S. gamegyn,
but in the former species, hairs on dorsolateral
margin of head posterior to apicoscrobal hair
are not projecting laterally (2-3 hairs obviously projected laterally in full face view for
S. gamegyn), pleurae and sides of propodeum
also mostly smooth and shiny (in S. gamegyn,
only the katepisternum is smooth and shining,rest of lateral mesosomal surface reticulatepunctate).

Strumigenys rogeri Emery, 1890
Material examined. Queen and workers, Pulau
Ubin, 22 Oct 2016, W. Wang leg., leaf litter, Winkler extraction, ZRC_ENT00007273;
queen and workers, Mandai Road, near Ulu
Sembawang Road, 14 Apr 1993, collector unknown, NS40, ZRC_HYM_0000278.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Pulau Ubin
Habitat/Ecology. This species appears to be associated mainly with disturbed young or maturesecondary forest, where the ants were foundin leaf litter.
Remarks. Deemed to be African in origin (Bolton
2000), S. rogeri was probably introduced toSingapore via human commerce – which has
been attributed to for the species’ spread toother parts of the world (Wetterer 2012c).
Strumigenys signeae Forel, 1905
Material examined. None available.
Material not physically examined. Unknown.
Literature. Bolton (2000).
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore context.
Remarks. At the time of writing, this species isonly known from Singapore based on a single
literature record.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Strumigenys sydorata Bolton, 2000
Material examined. Nee Soon forest, 2 Oct 2017,
W. Wang leg., leaf litter, Winkler extraction,
WT_FL1/NSW2, ZRC_ENT00000999.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Nee Soon forest
Habitat/Ecology. A single worker of this species
was found in leaf litter from old or mature secondary forest, at a site close to swamp forest.
Remarks. This species is a member of the lyroessa complex. At the time of writing, thesingle specimen identified as S. sydorata wassomewhat abraded, with many hairs missing
or damaged, including standing pronotal humeral hairs that are considered diagnostic of
this species.
Strumigenys szalayi Emery, 1897
Material examined. Punggol Road mangroves,
5 Aug 1979, D.H. Murphy leg., ZRC_
HYM_0000777.
Material not physically examined. Unknown.
Literature. Bolton (2000).
Localities. Punggol Road mangroves
Habitat/Ecology. This species was found in mangroves in Singapore, possibly mangrove backforest.
Syllophopsis australica (Forel, 1907)
Material examined. Queen and workers, Bukit
Timah Nature Reserve, near BT02, 2 Aug
2017, W. Wang leg., WW-SG17-030, ZRC_
ENT00000986; MacRitchie Reservoir, 9 Jan
2014, Sk. Yamane leg., ZRC_HYM_0000383384; Upper Peirce Reservoir, 10 Jan 2014, Sk.
Yamane leg., ZRC_HYM_0000385.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; MacRitchie Reservoir; Upper Peirce Reservoir.
Habitat/Ecology. This species was found in primary forest, and close to walking trails in old/
mature secondary forest. A nest was foundin soil at the base of a large tree in primary
forest. Individuals were often collected fromleaf litter.

Remarks. Most specimens examined for thischecklist were originally identified as ‘Monomorium talpa’ – now a junior synonym of S.
australica.
Syllophopsis sechellensis (Emery, 1894)
Material examined. None available.
Material not physically examined. Amber Road,
Katong, 28 Jul 2014, J.K. Wetterer leg., vial
#337 (MSC).
Literature. Wetterer & Sharaf (2017).
Localities. Amber Road (Katong)
Habitat/Ecology. Specimens were collected froma wooded area in an urban residential district.
Remarks. Workers of this species can be distinguished from most other congeners by amatte and reticulate punctate mesopleuron,and short propodeal spines. These ants areapparently native to the Old World tropics(Wetterer & Sharaf 2017), but have spreadto other regions, including the New World,
mainly by human commerce.
Syllophopsis subcoeca (Emery, 1894)
Material examined. None available.
Material not physically examined. Amber Road,
Katong, 28 Jul 2014, J.K. Wetterer leg., vials#337, #345 (MSC, USNM); Marine Parade,
28 Jul 2014, J.K. Wetterer leg., vial #368
(MSC, USNM); East Coast Park, 1 Aug 2014,
J.K. Wetterer leg., vial #442 (MSC, USNM);
Punggol Point, 30 Jul 2014, J.K. Wetterer
leg., vial #397 (MSC); Harbourfront, Marang Trail, 4 Aug 2014, J.K. Wetterer leg.,vial #472 (MSC).
Literature. Wetterer & Sharaf (2021).
Localities. Amber Road (Katong); East Coast
Park; Harbour Front; Punggol Point.
Habitat/Ecology. This species appears common inhighly-disturbed woodland and secondary forest fragments in close proximity to man-madeinfrastructure such as residential buildings andhighways. Specimens were also found at basesof trees in parks.

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Tetramorium aptum Bolton, 1977
Material examined. Bukit Timah forest, collection date unknown, D.H. Murphy leg., F36,
ZRC_HYM_0000753; Pulau Ubin, 1.41529,
103.94155, 15 Nov 2021, W.N. Lam leg.,
ZRC_ENT00057805.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah forest; Pulau Ubin.
Habitat/Ecology. Individuals of this species werecollected from either primary or old/maturesecondary forest in Singapore.
Tetramorium bicarinatum (Nylander, 1846)
Material examined. I-cube building (NUS),
1.293472, 103.776333, 24 Jun-1 Jul 2015,
M.S. Foo & W. Wang leg., malaise trap, ZRC_
BDP0044220; Temasek Polytechnic, adjacentwaste woodland, 1.34239, 103.93598, 13-15
Dec 2016, G.W. Yong leg., TT2GPa578,
ZRC_HYM_0001760; Lee Kong Chian Natural History Museum (LKCNHM), NUS, 13
Dec 2017, Sk. Yamane leg., SG17-SKY-42(SKYC).
Material not physically examined. Non-types,
J.K. Wetterer leg. (USNM) – Hougang, 30
Jul 2014, vial #401; Kallang, Riverside Park,
29 Jul 2014, vial #372; Katong, Amber Road,
28 Jul 2014, vials #333, 349; Kranji, by MRT
station, 2 Aug 2014, vial #452; Marine Parade,
Jul-Aug 2014, vials #352, 435; Tai Seng, by
MRT station, 31 Jul 2014, vial #422.
Literature. Viehmeyer (1916), Overbeck(1924) [both misclassified as ‘Tetramorium
guineense’]. Wang et al. (2018a).
Localities. Amber Road (Katong); Hougang;I-cube building (NUS); LKCNHM (NUS);
Katong; Kranji; Marine Parade; Riverside
Park (Kallang); Tai Seng; (beside) Temasek
Polytechnic.
Habitat/Ecology. This species is associated mainly with disturbed young secondary habitats insemi-urban settings in Singapore including
waste woodland forest, and sometimes also

scrubland. Colonies have been found in rotting wood and bamboo in a garden. A nest wasalso found in an electrical circuit box beside abus stop outside a university campus building.
Remarks. Tetramorium bicarinatum is considered a highly successful and globally widespread tramp species, occurring in almost all
tropical and subtropical habitats worldwide.
It has been strongly postulated to be native to
the Southeast Asian region, and introduced
to the rest of the world (Bolton 1977). Morphologically similar to another pantropical
tramp species – T. insolens (also listed here),
T. bicarinatum can be distinguished from thelatter by its sculptured mandibles, rectangularnodiform petiole with antero- and posterodorsal angles at identical heights, and finally its
bicolored body.
Tetramorium curtulum Emery, 1895
Material examined. Pulau Ubin, 7 Jan 2014, Sk.
Yamane leg., ZRC_HYM_0000457 (also inSKYC).
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Pulau Ubin
Remarks. This species has only been recorded
from an offshore island at the time of writing. A member of the scabrosum group, T.
curtulum can be distinguished by its relatively
smaller eyes (Bolton 1977).
Tetramorium eleates Forel, 1913
Material examined. Mandai Track, 4 Jan 2017,
G.W. Yong leg., ZRC_ENT00027989.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Mandai Track
Habitat/Ecology. This species was found in native-dominated old/mature secondary forest
in Singapore.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Tetramorium insolens (Smith, 1861)
Material examined. Kranji Road, 1 Nov 2016,
G.W. Yong leg., ZRC_ENT00048415; alates
and workers, National University of Singapore
campus (Clementi/Kent Ridge, multiple sites),
Apr-Aug 2015, M.S. Foo & W. Wang leg.,malaise trap, ZRC_BDP (multiple); queen andworkers, Sungei Buloh Wetland Reserve, 14
Mar 2018, W. Wang & M.S. Foo leg., WWSG18-Tetram1, ZRC_ENT00000907.
Material not physically examined. Unknown.
Literature. Wang et al. (2018a) [as Tetramorium
cf. insolens – ZRC material determined as T.
insolens post publication].
Localities. Kranji Road; National University ofSingapore campus (Clementi/Kent Ridge);
Sungei Buloh Wetland Reserve.
Habitat/Ecology. Tetramorium insolens is associated with a wide variety of different habitats
in Singapore, from mangrove back forest tourban cultivated patches, also disturbed secondary forest fragments in urban or semiurban settings. In back mangroves, the ants
have been found in a thin carton nest on theunderside of a thick branch of a living tree.
Remarks. This species is considered a relatively
widespread ‘pan-global’ tramp species originating from Indonesia and probably introduced elsewhere (Bolton 1977). It is morphologically similar to and often misidentified
as T. bicarinatum, but may be differentiatedby characters detailed in ‘Remarks’ under thelatter species section above.
Tetramorium kheperra (Bolton, 1976)
Material examined. Upper Peirce Reservoir, 10 Jan 2014, Sk. Yamane leg., ZRC_
HYM_0000415-417.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Upper Peirce Reservoir
Habitat/Ecology. This species was found in either
primary or old/mature secondary forest, neara walking trail.

Remarks. Tetramorium kheperra is known to be awidespread tramp species, where individuals
are mostly collected from disturbed habitats.
In view of this, its actual prevalence in Singapore may be underrepresented in sorted ZRC
material examined for this checklist.
Tetramorium lanuginosum Mayr, 1870
Material examined. Kent Ridge Park, 11 Jan
2014, Sk. Yamane leg., ex. leaf litter, SG14SKY-56 (SKYC); Kranji Road, 1 Nov 2016,
G.W. Yong leg., ZRC_ENT00048416; queen
and worker, Pulau Tekukor, 24 Jun 2017,
G.W. Yong leg., ZRC_ENT00048413; Sisters’
Island, 9 Dec 2016, G.W. Yong leg., ZRC_
ENT00027977; Yishun wasteland, 1.42078,
103.84934, 21 Feb 2016, M.K.L. Wong leg.,
ZRC_ENT00000698.
Material not physically examined. Type alate
queen, FOCOL2063 (ZMHB) [as Triglyphothrix lanuginosa]. Non-types, J.K. Wetterer leg.
(USNM) – Buona Vista, by MRT station, 5 Aug
2014, vial #486; City Hall, by church, 29 Jul
2014, vial #374; Dhoby Ghaut, Istana Park,
30 Jul 2014, vial #403; Harbourfront, 29 Jul
2014, vial #385; Hougang, 30 Jul 2014, vial
#401; Kallang, Riverside Park, 29 Jul 2014,
vial #372; Katong, Jul-Aug 2014, vials #348,
442; Kent Ridge, 31 Jul 2014, vials #426, 431;
Marine Parade, 28 Jul 2014, vials #358, 368;
Outram Park, by MRT station, 29 Jul 2014, vial
#379; Pasir Panjang, by MRT station, 5 Aug
2014, vial #482; Promenade, by MRT station,
4 Aug 2014, vial #468; Punggol Point, 30 Jul
2014, vial #397; Queenstown, 29 Jul 2014, vial
#383; Serangoon, 30 Jul 2014, vial #389.
Literature. Viehmeyer (1916), Overbeck (1924)[both as Triglyphothrix lanuginosa and T.
striatidens]. Wheeler (1929) [as type of Triglyphothrix striatidens flavescens]; Bolton(1976) [as Triglyphothrix lanuginosa].
Localities. Bukit Timah Road; Buona Vista; City
Hall; Dhoby Ghaut; Harbourfront; Hougang;
Katong; Kent Ridge; Kranji Road; Marine
Parade; Outram Park; Pasir Panjang; Promenade; Pulau Tekukor; Punggol Point; Queenstown; Riverside Park (Kallang); Serangoon;
Sisters’ Island; Yishun wasteland.

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Habitat/Ecology. This species was collected
mainly from young secondary habitats, including waste woodland forest fragments inurban or semi-urban settings, and scrublandon offshore islands. The ants were also foundnesting in a hollow spot under the bottom of aflower stand in a garden. Foragers were often
also found in plantings, bases of trees, or gardens close to man-made infrastructure.
Remarks. Tetramorium lanuginosum is a recognized globally widespread tramp species commonly known as the wooly ant, suspected tohave been mainly dispersed through humancommerce (Wetterer 2010). Its native range
remains a point of contention, though somebelieve it is native to tropical Asia, or more
specifically tropical and subtropical East Asia.
The type queen of the species was originallydescribed from Singaporeas ‘Triglyphothrix
lanuginosa’ (see Viehmeyer 1916).
Tetramorium meshena (Bolton, 1976)
Material examined. Lower Peirce Reservoir,
Forest A, 7 Aug 1990, collector unknown,
ZRC_ENT00000176; same locality, site andcollection date as previous, collector unknown, ZRC_ENT00000194; same locality
as previous, Forest B, 25 Aug 1990, collector
unknown, ZRC_ENT00000115; same locality and site as previous, 10 Aug 1990, collector unknown, ZRC_ENT00000227; Upper
Thomson Nature Park,1.38311, 103.79839,
17-19 Oct 2016, pitfall trap, UT5GPc1629,
ZRC_HYM_0001761; Pulau Ubin, 7 Jan
2014, Sk. Yamane leg. (SKYC).
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Lower Peirce Reservoir; Pulau Ubin;
Upper Thomson Nature Park.
Habitat/Ecology. This species is associatedmostly with primary or old/mature secondary
forests in Singapore. Some specimens werecollected from young abandoned plantation
secondary forests. Individuals were usuallyfound in leaf litter.

Tetramorium obtusidens Viehmeyer, 1916
Material examined. Bukit Timah forest, collection date unknown, D.H. Murphy leg.,A124-2, ZRC_HYM_0000749; University
Hall (NUS), 1.29711, 103.77658, 1-8 Apr
2015, M.S. Foo & W. Wang leg., malaise trap,
NUS0008, ZRC_BDP0045351; same locality
and collectors as previous, 10-17 Jun 2015,
NUS0048, ZRC_BDP0044430.
Material not physically examined. Type queens– FOCOL2072-2073 (ZMHB).
Literature. Type – Viehmeyer (1916); Overbeck(1924). Bolton (1977); Wang et al. (2018a).
Localities. Bukit Timah forest; Jurong Road
[type]; University Hall (NUS).
Habitat/Ecology. This species appears associatedwith primary and/or mature secondary forests
in Singapore, including disturbed forest fragments in semi-urban settings. Queens havebeen found alongside other ant species suchas Cataulacus praetextus and Technomoyrmex
albipes in rotten branches.
Remarks. Type locality in Singapore. The species
was first described based on syntype queens(see Viehmeyer 1916), while the worker was
described much later (see Bolton 1977) basedon specimens from a locality different fromthat of the queen types, and also specimensfrom other countries.
Tetramorium pacificum Mayr, 1870
Material examined. Commonwealth Road,
1.31115, 103.78171, 22-24 Nov 2016, G.W.
Yong leg., ZRC_HYM_0001758-1759; Icube building (NUS), 1.29347, 103.77633,
Jul-Sep 2015, M.S. Foo & W. Wang leg.,malaise trap, ZRC_BDP (multiple); Kent
Ridge, 20 Apr 1985, D.H. Murphy leg.,
ZRC_HYM_0000757; Mandai, 1.4083,
103.77803, 19 Jun 2016, M.K.L. Wongleg., ZRC_ENT00000700; Namly forest,
10 May 1985, D.H. Murphy leg., ZRC_
HYM_0000746; Prince George’s Park Residences (NUS), 1.29239, 103.77869, May-Sep
2015, M.S. Foo & W. Wang leg., ZRC_BDP(multiple); Yishun wasteland, 1.42078,

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

103.84934, 21 Feb 2016, M.K.L. Wong leg.,
ZRC_ENT00000699; Bukit Timah, 14 Apr
1993, Sk. Yamane leg. (SKYC); worker andfounding queen, Upper Thomson Nature Park,
9 Dec 2017, Sk. Yamane leg., SG17-SKY-38(SKYC).
Material not physically examined. In SchlickSteiner et al. (2006) - queen, Singapore Zoo,
12 Jan 2002, H. Zettel leg.; queen, National
University of Singapore campus, 26-28 Jan
2002, H. Zettel leg.; 15 queens, same locality
as previous, 21 Oct - 7 Nov 2003, H. Zettel
leg. (all in ZCW).
Literature. Schlick-Steiner et al. (2006); Wang etal. (2018a); Yamane et al. (2022).
Localities. Bukit Timah; Commonwealth Road;
Kent Ridge; Mandai; Namly forest; National
University of Singapore campus (Clementi/
Kent Ridge, multiple sites); Singapore Zoo;
Yishun wasteland.
Habitat/Ecology. This species is associated mostly with young secondary forests of varying
disturbance levels in Singapore, including
waste woodland secondary forest. Individuals
have been collected on bamboo, tree trunks,
in decayed wood, and on foliage of Wormia(currently known as ‘Dillenia’) shrubs/trees.
Remarks. The species has a very broad distribution range throughout most of the Oriental andIndo-Australian realms.
Tetramorium pulchellum Emery, 1897
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah Road
Habitat/Ecology. This species was found in agarden.
Remarks. At the time of writing, this species isonly known from Singapore based on literature records.
Tetramorium scabrum Mayr, 1879
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah Road

Habitat/Ecology. Individual workers were collected from a garden.
Remarks. At the time of writing, this species isonly known from Singapore based on literature records.
Tetramorium smithi Mayr, 1879
Material examined. Kent Ridge, 25 Apr 1985,
D.H. Murphy leg., ZRC_HYM_0000756;
Mandai Lake Road, N 01°24’19.3’’, E
103°46’56.2’’, 73 m a.s.l., 26 Mar 2020, W.
Zhang leg., ZRC_ENT00047776.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[both as Xiphomyrmex smithi var. kanarense].
Tan & Corlett (2012).
Localities. Bukit Timah Road; Kent Ridge.
Habitat/Ecology. This tiny species was foundmostly in disturbed young or mature secondary forests, in clay mounds and fallen rotting
wood or logs. A colony was also found in the
ground of a garden.
Tetramorium tonganum Mayr, 1870
Material examined. Prince George’s Park Residences (NUS), 1.29239, 103.77869, Apr-May
2015, M.S. Foo & W. Wang leg., malaise trap,
ZRC_BDP0045634, -46101, -46770; Pulau
Ubin, 7 Jan 2014, Sk. Yamane leg., ZRC_
HYM_0000455-456.
Material not physically examined. Unknown.
Literature. Wang et al. (2018a).
Localities. Prince George’s Park Residences(NUS); Pulau Ubin.
Habitat/Ecology. This species is associated withyoung secondary habitat in Singapore, mostly
disturbed secondary forest fragments withinsemi-urban settings, or close to man-madeinfrastructure.
Remarks. It has been surmised that Java andSumba (both part of Indonesia) represent thewest-most limits of the species native range,
and that populations in Malaysia and Singapore are possibly introduced (Bolton 1977).

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Tetramorium walshi (Forel, 1890)

Trichomyrmex mayri (Forel, 1902)

Material examined. Sisters’ Island, 9 Dec 2016,
G.W. Yong leg., ZRC_ENT00027991; locality unknown, 23 Jul 1996, Sk. Yamane leg.
(SKYC).
Material not physically examined.
CASENT0909099, ANTC31549 (MHNG) [assyntype of Triglyphothrix walshi var. spuria].
Literature. Forel (1912) [as type of Triglyphothrix
walshi var. spuria].
Localities. Sisters’ Island
Habitat/Ecology. This species was found on an
offshore island, possibly in young or maturesecondary forest.

Material examined. None available.
Material not physically examined. Unknown.
Literature. Forel (1911) [as Monomorium destructor subsp. mayri].
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore context.
Remarks. At the time of writing, this species isknown from Singapore based only on a single
literature record.

Trichomyrmex destructor (Jerdon, 1851)
Material examined. None available (but see Remarks).
Material not physically examined. In Wetterer(2009) – 1899, Haviland (BMNH). ANIC32061821, Shattuck1345379064 (ANIC).
Literature. Wetterer (2009).
Localities. Unknown.
Habitat/Ecology. Uncertain in the Singapore context, but presumably in and around man-madeinfrastructure in urban and semi-urban areas,
based on anecdotal observations.
Remarks. Commonly known as ‘the Singapore
ant’, T. destructor is an infamous worldwide
cosmopolitan pest species that appears to not
be as locally ubiquitous as is suggested by its
common name. It is presumably native to the
Palearctic region (Wetterer 2015). The species is probably underrepresented in the ZRC– wholly in uncatalogued, unsorted material(W. Wang, pers. obs.). There have been pastoccasional anecdotal reports of the pest ant
causing damage to man-made infrastructure,such as chewing through electrical wiringin urban buildings, but species identities of
the reported ants have not been validated. In
Bolton (1987), the locality of material supposedly collected from Singapore was indicatedas ‘Sabang’ – which may actually be a city or
town located in Indonesia or Malaysia, butdoes not exist in Singapore.

Tyrannomyrmex rex Fernández, 2003
Material examined. Mandai, 1.40144, 103.77702,
10 Mar 2016, M.K.L. Wong leg., MKWSG16-Tyr1, ZRC_ENT00000703-705.
Material not physically examined. In Jacquemin
et al. (2015) – SPM_ID4904401, general inventory 33045 (RBINS).
Literature. Jacquemin et al. (2015); Wong & Yong(2017).
Localities. MacRitchie Reservoir; Mandai.
Habitat/Ecology. This species was found inprimary or old/mature secondary forests in
Singapore, typically in leaf litter. Nests were
found at bases of trees.
Remarks. In one literature record, habitat was described as ‘selectively logged and regenerated
primary forest’ – this can be simply and moreaccurately inferred as ‘secondary forest’ byour definition (see Introduction).
Vollenhovia brevicornis (Emery, 1893)
Vollenhovia fridae Forel, 1923, worker, Sumatra,syn. nov.
Material examined. Mandai Road, 1.41258,
103.79839, 16-18 Nov 2016, G.W. Yongleg., ZRC_HYM_0001711-1712; Mandai
Lake Road, Northern Node, 1°24’36.5”N,
103°47’04.0”, 53 m, 18 Jul 2019, J.S. Tan &N.L. Chin leg., leaf litter, Winkler extraction,
ZRC_ENT00028311; Bukit Timah forest,
3 Sep 1964, D.H. Murphy leg., A-292-10,
DHM-SG64-Voll1, ZRC_ENT00027927;
locality unknown, 14 Apr 1993, Sk. Yamaneleg. (SKYC).

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Mandai
Lake Road; Mandai Road.
Habitat/Ecology. This species was found in primary and/or native-dominated old or maturesecondary forests, sometimes on tree trunks,
but mainly from leaf litter and upper topsoil
substrate.
Remarks. We have carefully compared the Sumatran specimens (workers) of ‘Vollenhovia fridae’ with the images and original descriptionof V. brevicornis (described in Monomorium),and concluded that these are the same species.
Singaporean specimens examined well agreewith V. brevicornis and ‘V. fridae’. Althoughthese are slightly smaller than the Sumatran
specimens, in this species the worker caste ishighly variable in size. This species appears
almost identical to the Sulawesi species Vollenhovia pertinax (Smith, 1861) that wasmuch earlier described than V. brevicornis,
but we need more information and material todecide the status of these two forms (see also
Remarks for Vollenhovia penetrans Smith,1857).
Vollenhovia escherichi Forel, 1911
Material examined. Pulau Ubin, 1.41662,
103.99443, 4 Dec 2016, G.W. Yong & M.K.L.
Wong leg., ZRC_HYM_0001077; Pulau
Ubin, Jalan Sam Heng, 15 Oct 2016, G.W.
Yong et al. leg., ZRC_HYM_0001146.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Pulau Ubin
Habitat/Ecology. This species was found in young
and/or mature secondary forest; an individual
was collected from leaf litter by Winkler extraction.
Remarks. At the time of writing, V. escherichi was
identified only from an off-shore island northeast of the mainland. The species resembles
Vollenhovia subtilis, but can be distinguishedby the reticulate sculpture on head dorsum
with larger interspaces or areola, larger eyes(also bulging more from sides of head in fullface view), and postpetiole in dorsal view
distinctly wider than long and as wide as the
propodeum.

Vollenhovia minuta Viehmeyer, 1916 stat. n.
Vollenhovia brevicornis var. minuta Viehmeyer,
1916: 129 (w.)
Subspecies of Vollenhovia brevicornis: Emery,
1922: 164; Ettershank, 1966: 149; Bolton,
1995: 423.
Material examined. Males and workers, Mandai forest, 7 Jun 2018, W. Wang leg., WWSG18-002, ZRC_ENT00027960; Nee Soon
forest, 7 Jan 2015, W. Wang leg., ZRC_
ENT00000998; Upper Thomson Road, 9
Dec 2017, W. Wang leg., WW-SG17-Voll1,
ZRC_ENT00000963; same locality as previous, 11 Oct 2016, G.W. Yong leg., ZRC_
ENT00048027; queen, Upper Thomson Nature Park, 1.38159, 103.81921, 11 Oct 2016,
G.W. Yong et al. leg., ZRC_ENT00048028.
Material not physically examined. Unknown.
Literature. None (see Remarks). New record.
Localities. Mandai forest
Habitat/Ecology. This species was found in old/
mature secondary forest, including abandoned
plantation forests, typically nesting under barkof large fallen tree trunks or near the bases oflive trees.
Remarks. The type locality for this species is in
Gunong Angsi in Malaysia (see Overbeck
1924), not Singapore as erroneously indicatedin most existing publications and online resources (e.g. Bolton 2021, AntWeb).
		This species was originally described asa subspecies of V. brevicornis, however, it ismorphologically completely different fromthe latter and should be treated as a distinct
species. Vollenhovia minuta can be distinguished by the following characters: 1) minutesize (TL 1.8 – 2 mm), 2) head much longer
than wide, 3) head in profile with flattened
appearance, 4) antennal scapes very short,
not reaching posterior head margin, 5) mesosoma elongate with almost flat dorsum, 6) no
propodeal spines, 7) head, mesosoma, petiole
and postpetiole densely reticulate-punctate.
The species resembles V. piroskae taipingensis Forel, 1913, but can be differentiated byits much longer head in full face view (in V.
piroskae taipingensis, head is scarcely longer
than wide).

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Vollenhovia overbecki Viehmeyer, 1916
Material examined. Queens, Bukit Timah forest,
A124-11, 17 Dec 1972, D.H. Murphy leg.
Material not physically examined. Types –CASENT0908662, ANTC031013 (MHNG);
male and worker, FOCOL0313-314 (DEIC);
2 males and 3 workers, FOCOL1664-1668(ZMHB).
Literature. Type – Viehmeyer (1916); Overbeck(1924).
Localities. Bukit Timah forest; Bukit Timah Road;
Jurong Road.
Habitat/Ecology. The type colony was found in arotten trunk of a tree, macrohabitat unspecified. Individuals were also found in a garden,
and in either primary or old/mature secondary
forest.
Remarks. Type locality/localities in Singapore.
Species identity of material examined waslargely deduced based on original descriptionsof the worker, since no images or descriptionsof the queen were available for reference atthe time of writing.

from Borneo, which had in turn been directlycompared with type specimens of V. fridae
at the MHNG by one of us (SKY). All thesestrongly imply the conspecifity of the four
forms: brevicornis, fridae, penetrans, pertinax(see Remarks for V. brevicornis).
		
If that is the case, the oldest name should
be V. penetrans, with all the others being its
junior synonyms.
		However, we are reluctant to synonymize
these three forms under V. penetrans, as it remains possible that some of these constitutea cryptic species complex. For example, we
detected a slight difference among the queens
examined, namely the space between frontal
carinae (posterior portion of clypeus) – thisarea is densely striate-reticulate in the Singapore queen and the type specimen image, butsmooth in the queens from Borneo, the Malay
Peninsula and Sulawesi. We believe that allthese forms are closely related, but final species statuses should be decided with assessment of more information – including male
genitalia and DNA – amassed in future.

Vollenhovia penetrans (Smith, 1857)

Vollenhovia rufiventris Forel, 1901

Material examined. Queen, Mandai Road, 18 Nov
2016, G.W. Yong leg., ZRC_ENT00027986.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Mandai Road
Habitat/Ecology. A single queen of this species
was collected from native-dominated old/mature secondary forest in Singapore.
Remarks. This species was first described as ‘Atta
penetrans’ based on a single queen from Borneo (Smith 1857). At the time of writing, we
verified the identity of ZRC material withthe original description and holotype images
available online.
		We also examined a few queen specimens associated with workers from Borneo
and the Malay Peninsula. A single isolated
queen from Sulawesi well agreed with thesequeens. Furthermore, queen-associated workers mentioned above well agreed with separate samples of workers (without queens)

Material examined. Bukit Timah forest,
16 Feb 1975, D.H. Murphy leg., ZRC_
HYM_0000901.
Material not physically examined. Types – maleand dealate queen, FOCOL2920-FOCOL2921(ZMHB).
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah forest; Bukit Timah Road.
Habitat/Ecology. This species is associated withprimary and mature secondary habitats in Singapore. Individuals can sometimes be seen on
fallen wood or logs. Colonies can be found in
the soft wood of rotting fallen trees.
Remarks. The male and queen of this species werefirst described from Singapore (see Viehmeyer1916), based on a colony series collected froma garden, many years after the species itself
was established in 1901 based on the worker.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

PONERINAE (20 genera, 61 species)
Anochetus agilis Emery, 1901
Material examined. None available.
Material not physically examined. ANIC32016869, Shattuck1345241385 (ANIC).
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah Nature Reserve; Bukit
Timah Road.
Habitat/Ecology. Individuals including the type
queens were collected from a garden. This
species was found in either primary or old/
mature secondary forest in more recent times.
Remarks. The queen of this species was firstdescribed by Viehmeyer (1916) based on
specimens collected by H. Overbeck fromSingapore.
Anochetus graeffei Mayr, 1870
Material examined. Kent Ridge Park, 7 Dec
2017, W. Wang leg.,WW-SG17-027, ZRC_
ENT00000979; workers and dealate queen,same locality as previous, 7 Dec 2017, Sk. Yamane leg., SG17-SKY-21 (SKYC); Nee Soon
forest, 27 Jan 2015, W. Wang leg., WW-SG15Anoc1, ZRC_ENT00000978; male, Pulau Semakau Old Fragment, SMO3, 1°12’23.9”N,
103°45’37.6”E, 5-12 Jul 2012, J. Puniamoorthy et al. leg., mangroves, malaise trap,
Reg. 29342, ZRC_BDP0014528; queens
and males, Pulau Ubin, PU1, 1°24’36.3”N,
103°59’25.5”E, Jul-Nov 2012, J. Puniamoorthy et al. leg., mangroves, malaise trap, ZRC_
BDP (multiple); male, Sungei Buloh Wetland
Reserve, SB1, 1°26›46.3»N, 103°43›49.9»E,
21-28 Nov 2012, J. Puniamoorthy et al. leg.,mangroves, malaise trap, Reg. 29595, ZRC_
BDP0017502; Upper Thomson Road, 24 Oct
2016, G.W. Yong leg., ZRC_ENT00048414.
Material not physically examined. ANIC32015989, Shattuck1345236804 (ANIC).
Literature. Viehmeyer (1916), Overbeck (1924)[both as Anochetus punctiventris]. Shattuck& Slipinska (2012).
Localities. Kent Ridge Park; Labrador Nature Reserve; Nee Soon forest; Pulau Semakau; Pulau
Ubin; Sungei Buloh Wetland Reserve; Upper
Thomson Road.

Habitat/Ecology. Workers of this species were
found in a garden, semi-urban parklands,
young secondary forest and also Imperata
cylindrica-dominated grassy scrub. Nests
have been collected from dead twigs in leaf
litter. Alates were collected from mangroves
using malaise traps.
Remarks. This species is known to exhibit broad
morphological variation between different
populations, and may actually comprise acomplex of multiple sibling species. In thischecklist, we tentatively treat all morphological variants of this potential species complex
in the ZRC as conspecific, bearing in mind thepossibility that these might be separated into
different species should more convincing empirical evidence be made available in future.
Anochetus incultus Brown, 1978
Material examined. None available.
Material not physically examined. ANIC32016921, Shattuck1345241729 (ANIC).
Literature. None. New record.
Localities. Bukit Timah Nature Reserve
Habitat/Ecology. This species is associated withprimary or old/mature secondary forest in
Singapore. Individuals were found in rotting
logs or leaf litter.
Remarks. At the time of writing, this species isknown from only one specific locality in Singapore.
Anochetus muzziolii Menozzi, 1932
Material examined. Mandai, 1.40469, 103.77653,
6 Mar 2016, M.K.L. Wong leg., ZRC_
ENT00000706; Upper Seletar Reservoir, 4
Oct 2020, G.W. Yong leg., GY-SG20-001,
ZRC_ENT00028566.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Mandai; Upper Seletar Reservoir.
Habitat/Ecology. This species was collected fromeither mature or young secondary forest, often
close to trailside, on one occasion on deadwood.
Remarks. A member of the A. rugosus species
group, A. muzziolii is largely distinguished byits smooth and shiny pronotum, and relatively
short mandible.

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Anochetus rugosus (Smith, 1857)
Material examined. None available.
Material not physically examined. Holotype,Singapore, A.R. Wallace leg. (OUMNH).
Literature. Type – Smith (1857) [as Odontomachus rugosus].
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore context.
Remarks. Type locality in Singapore, specifics
unknown. The only Smith collection specimen of Anochetus rugosus in OUMNH – assumed to be the holotype – bears two locality labels: ‘SAR’ and ‘Gilolo’, which refer tolocalities outside Singapore. It is speculated
that these labels may have been switched withthe original type label some time after 1857
(Bolton 2021).
Anochetus tua Brown, 1978
Material examined. None available.
Material not physically examined. ANIC32016967, Shattuck1345241965 (ANIC).
Literature. None. New record.
Localities. Nee Soon Swamp Forest
Habitat/Ecology. This species was found inswamp forest, collected from leaf mould
berlesate.
Brachyponera luteipes (Mayr, 1862)
Material examined. Bukit Timah Nature Reserve,plot III-2, 28 Apr 2016, D.J. Court leg., pitfall
trap, ZRC_ENT00048448; Mandai forest, 7
Jun 2018, W. Wang leg., ZRC_ENT00047995;
queen and workers, Mandai, Northern Node,1.40842, 103.78319, 54 m a.s.l., 25 Jul
2019, J. Tan & S. Tang leg., JTST-MIS-009,
ZRC_ENT00013871; same locality as previous, 4 Sep 2019, J. Tan & N. Chin leg., WWSG19-008, ZRC_ENT00027941.
Material not physically examined. Harbourfront,
Jul-Aug 2014, J.K. Wetterer leg., vials #386,
469 471 (JKWC).
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Harbourfront; Mandai.

Habitat/Ecology. This species was mainly foundin old or mature secondary forests in Singapore, sometimes at bases of trees at forest or
trail edges. Nests were collected from fallen
rotting wood or large branches, sometimes inroot-soil-substrate matrices at bases of trees.
Remarks. Morphologically very similar to thepresumably widespread invasive B. chinensis, B. luteipes may be differentiated fromthe former based on characters illustrated in
Yashiro et al. (2010). An affiliated unnamed
species, currently identified as Brachyponera
sp.1.of.SKY (= Pachycondyla sp.28.of.SKY),was found to be common across secondary
forests and seemingly more widespread thanB. luteipes.
Brachyponera obscurans (Walker, 1859)
Material examined. Alate queen and male, University Town (NUS), 1.30622, 103.77458,
12-19 Aug 2015, M.S. Foo & W. Wang leg.,malaise trap, ZRC_BDP0042558, 42561.
Material not physically examined. Unknown.
Literature. Wang et al. (2018a).
Localities. University Town (NUS)
Habitat/Ecology. Alates of this species were collected from a malaise trap in cultivated grassy
scrub, in an urban setting.
Remarks. At the time of writing, the only recordsof this species in Singapore are that of winged
alates collected using malaise traps.
Buniapone amblyops (Emery, 1887)
Material examined. Bukit Timah Nature Reserve, plot III-2, 19 Apr 2016, D.J. Court
leg., pitfall trap, ZRC_ENT00048455;
same locality as previous, below Kruing(sic) Hut, 4 Oct 1973, D.H. Murphy leg.,
A340-1, ZRC_ENT00047941; Kent Ridge
scrub, 15 Jan 1981, D.H. Murphy leg.,
ZRC_ENT00047942; Pulau Ubin, 1.41662,
103.99443, 4 Dec 2016, G.W. Yong & M.K.L.
Wong leg., ZRC_HYM_0001090; Seletar Trail, 1.3946, 103.80113, 28 Aug 2015,
M.K.L. Wong leg., ZRC_ENT00000708.
Material not physically examined.
CASENT0172431, A64-9 (ANIC).

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Literature. Viehmeyer (1916), Overbeck (1924)[both as Pseudoponera amblyops]; Yong etal. (2017).
Localities. Bukit Timah Nature Reserve; Kent
Ridge; Pulau Ubin; Seletar Trail.
Habitat/Ecology. This species is associated mostly
with primary or old/mature secondary forests
in Singapore, including disturbed secondary
forest fragments or scrubland in semi-urban
settings. Individuals were sometimes foundunder stones, planked steps, and imbedded
bricks on a rock or sandy paths in mature secondary forest or scrub.
Centromyrmex feae greeni Forel, 1901
Material examined. None available.
Material not physically examined. Syntype –CASENT0907211, ANTC27417 (MHNG).
Literature. Type – Forel (1901).
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore context.
Remarks. At the time of writing, this subspecies isknown from Singapore only by its type series.
Type locality in Singapore.
Centromyrmex hamulatus (Karavaiev, 1925)
Material examined. Bukit Timah Nature Reserve,
2016, W. Wang leg., ZRC_ENT00028294;
Nee Soon, 23 May 1973, D.H. Murphy leg.,
ZRC_HYM_0000762-763; alate queen, Pulau
Ubin West, 21-28 May 2019, D. Yap et al. leg.,mangroves, malaise trap, Reg. 5242, ZRC_
ENT00047844; Sime Road forest, 4 Apr 1975,
D.H. Murphy leg., ZRC_HYM_0000893-894.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Nee
Soon; Pulau Ubin West; Sime Road forest.
Habitat/Ecology. This species seems mostly associated with primary or old/mature secondary
forests in Singapore, including dry secondary
forest adjacent to swamp forest. It is considered subterranean, and individuals could befound in soil at bases of trees, under logs, inside rotting logs and termite mounds, suggesting a possible termitophagous nature of thisspecies. An alate queen was collected from amalaise trap set up in mangroves.

Cryptopone butteli Forel, 1913
Material examined. Mandai Lake Road, buffer area (Northern Node), 1°24’36.5”N,
103°47’04.0”, 53m, 18 Jul 2019, J.S. Tan& N.L. Chin leg.,Winkler extraction, ZRC_
ENT00028295; Nee Soon forest, 24 Dec
2014, W. Wang leg., ZRC_ENT00027965;
Seletar Trail, 1.39097, 103.79952, 7 Dec
2015, M.K.L. Wong leg., Winkler extraction,
ZRC_ENT00000709.
Material not physically examined. ANIC32017898-17899, Berlno3; ANIC32-017067,
Shattuck1345242298; ANIC32-017084, Shattuck1345242360 (all ANIC).
Literature. None. New record.
Localities. Bukit Timah forest; Mandai Lake
Road; Nee Soon forest; (near) Peirce Reservoir; Seletar Trail.
Habitat/Ecology. This species was found in primaryor old/mature secondary forests in Singapore,including swamp forest. Typically collectedfrom forest leaf litter via Winkler or Berlese
extractions, also in soil and rotting logs.
Cryptopone testacea Emery, 1893
Material examined. Mandai Lake Road, buffer area (Northern Node), 1°24’36.5”N,
103°47’04.0”, 53m, 18 Jul 2019, J.S. Tan &N.L. Chin leg., Winkler extraction, ZRC_
ENT00028296.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Mandai Lake Road
Habitat/Ecology. Individuals of this species were
found in leaf litter and upper soil substratefrom mature secondary forest.
Remarks. At the time of writing, this species isonly known from one locality in Singapore.
		Cryptopone testacea was first describedfrom Sri Lanka, and currently appears tohave a broad geographic distribution from
the Malagasy region to Micronesia. Multiple
species endemic to different regions have beensynonymized under C. testacea, albeit withsubtle morphological differences observed
between the congeners. It remains possiblethat ‘C. testacea’ could comprise a complex ofmorphologically-similar species, but this issue
may only be resolved with DNA evidence.

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Diacamma geometricum (Smith, 1857)

Diacamma pallidum (Smith, 1858)

Material examined. Mandai Lake Road, buffer area (Northern Node), 1°24’36.5”N,
103°47’04.0”, 53 m, 18 Jul 2019, J.S. Tan &N.L. Chin leg., ZRC_ENT00047849; Mandai
Road, plot 1, 1.41263, 103.79237, 21 Nov
2016, G.W. Yong et al. leg., M1D17GH2031,
ZRC_ENT00047840; Sime Road forest, 2 Jan 1978, D.H. Murphy leg., ZRC_
HYM_0000812; locality unknown, 14 Apr
1993, Sk. Yamane leg., ZRC_HYM_0000421422; MacRitchie Reservoir, 6 Jan 2014, Sk.
Yamane leg. (SKYC).
Material not physically examined. Holotype CASENT0901342, ANTC20781 (OUMNH)[as Ponera geometrica]. Non-types CASENT0916122-916123, ANTC43588ANTC43589 (CASC); Harbourfront, Marang
Trail, 4 Aug 2014, J.K. Wetterer leg., vial #473
(JKWC); Punggol Point, 30 Jul 2014, J.K.
Wetterer leg., vial #399 (JKWC). In Laciny etal. (2015) – McRitchie (sic) Reservoir, reservearea, 23 Oct 2003, H. Zettel leg. (ZCW).
Literature. Type – Smith (1857) [as Ponera geometrica]. Viehmeyer (1916), Overbeck (1924)[both as D. tritschleri]; Laciny et al. (2015);
Zettel et al. (2016).
Localities. Bukit Timah Hill; Harbourfront; MacRitchie Reservoir; Mandai Lake Road; Mandai Road; Punggol Point; Seletar Reservoir;
Sime Road forest; Thompson Road.
Habitat/Ecology. This species is associated mostly with old/mature secondary forests in Singapore, also sometimes in young secondary
forest. Individuals have been collected fromleaf litter and upper soil substrate.
Remarks. Type locality in Singapore. Diacamma
geometricum was initially considered a synonym of D. rugosum (see Wilson 1958), but
was revived/restored to species by Laciny etal. (2015).

Material examined. Bukit Timah Nature Reserve,
13 Dec 1995, H.K. Lua et al. leg., NS 212C,
ZRC_ENT00000352; same locality as previous, Hindhede Drive, 25 Dec 1989, H.K. Lualeg., ZRC_ENT00000241; males and workers, National University of Singapore campus (Clementi/Kent Ridge, multiple sites),
Apr-Sep 2015, M.S. Foo & W. Wang leg.,malaise trap, ZRC_BDP (multiple); Upper
Peirce Reservoir, 10 Jan 2014, Sk. Yamaneleg., SG14-SKY-44, ZRC_HYM_0000484.
Material not physically examined. Unknown.
Literature. Wang et al. (2018a).
Localities. Bukit Timah Nature Reserve; National
University of Singapore campus (Clementi/
Kent Ridge); Upper Peirce Reservoir.
Habitat/Ecology. This species was found in bothmature and young secondary forests broadly
variable levels of disturbance and degradation, including disturbed secondary forest
fragments in urban or semi-urban settings.
In contiguous mature secondary forest, nests
were found in rotting wood. Male alates werealso collected from malaise traps set up in
cultivated grassy scrub in urban settings.
Remarks. Most verified records of this species
comprise of males collected via malaisetraps; the original description of the species
by Smith (1858) was also only of the male.
This species is often assumed to be D. rugosum at first glance, but differs from the latter
both morphologically (differences may only
be visible under the microscope) and in termsof overall size — D. pallidum is generally
smaller in size relative to D. rugosum.
Diacamma rugosum (Le Guillou, 1842)
Material examined. Lower Peirce Reservoir
Road, 21 Jul 1990, H.K. Lua leg., ZRC_
HYM_0000072; Pulau Ubin, 1.41662,
103.99443, 4 Dec 2016, G.W. Yong & M.K.L.
Wong leg., ZRC_HYM_0001093; same locality, collectors, and collection date as previous, Jalan Sam Heng, 1.412604, 103.99317,
ZRC_HYM_0001094.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Material not physically examined. Unknown.
Literature. Forel (1907, 1911) [as Diacamma
rugosum birmanum; see Remarks]. Tan &Corlett (2012).
Localities. Central Catchment Nature Reserve;
Lower Peirce Reservoir Road; Pulau Ubin.
Habitat/Ecology. This species is associatedmostly with old/mature secondary forests in
Singapore, often disturbed forest fragments
in semi-urban settings, less frequently in primary forest.
Remarks. At the time of writing, there is much
uncertainty surrounding the species D. rugosum and the taxonomic validity of its 21 valid
subspecies. Given the taxonomic uncertainties
at the time of writing, we therefore considered
all subspecies of D. rugosum (e.g. D. rugosum birmanum) as conspecific to the species
for the purposes of this checklist. We recognize the possibility that at least some of thesesubspecies might be raised to species status
in future, which should entail a revision of
Diacamma species in this checklist.
Diacamma vagans (Smith, 1860)
Material examined. Kent Ridge, 1 Jun 1985, D.H.
Murphy leg., ZRC_HYM_0000814-815.
Material not physically examined. Unknown.
Literature. Emery (1893).
Localities. Kent Ridge
Habitat/Ecology. Individuals of this species were
found in grassland or scrub at the fringes ofsecondary forest in Singapore.
Ectomomyrmex leeuwenhoeki (Forel, 1896)
Material examined. Worker and queen, Bukit
Timah Nature Reserve, 1.34772, 103.77766,
30 Nov 2015, M.K.L. Wong leg., ZRC_
ENT00000711-712; same locality as previous,
plot II-I, 15 Aug 2011, J. Koh leg., Winkler
extraction, ZRC_ENT00000888; same locality as previous, BT04, 1.35342, 103.7782,
19 Jan 2017, W. Wang leg., WW-SG17-010,
ZRC_HYM_0000583; queen, same locality and collector as previous, near BT02,
1.35599, 103.77397, 8 Aug 2017, ZRC_
HYM_0001746; queen and worker, same lo-cality and collector as previous, near BT01, 7
Jun 2017, ZRC_ENT00014153; same locality and collector as previous, BT09, 1.35127,
103.78161, 5 Jul 2017, ZRC_ENT00014154;
Central Catchment Nature Reserve, 16 Feb
2016, G.W. Yong leg., ZRC_ENT00047846;
same locality as previous, 10 Aug 2011, E.J.Y.
Soh leg., ZRC_ENT00048001; Lower Peirce
Reservoir, 7 May 2017, G.W. Yong leg., ZRC_
ENT00048420; same locality as previous,
Forest B, 10 Aug 1990, collector unknown,
ZRC_ENT00000205; same locality as previous, Forest A, 7 Aug 1990, collector unknown,
ZRC_ENT00000127; Mandai Lake Road,
Northern Node, 1°24’36.5”N, 103°47’04.0”,
18 Jul 2019, J.S. Tan & N.L. Chin leg.,Winkler
extraction, ZRC_ENT00028287-28288; Mandai Road, 1.41054, 103.79809, 5 Dec 2016,
G.W. Yong et al. leg., M1D5GH2474, ZRC_
ENT00047845; queen and worker, Nee Soon
Swamp Forest, 1.38252, 103.80208, 25 Sep
2017, W. Wang leg., NS_W1, Winkler extraction, ZRC_ENT00027909; same locality as
previous, dry plot 301, 1.38043, 103.8033,
22 Aug 2019, W. Wang leg.,WW-SG19-002,
ZRC_ENT00013872; alate queens and males,same location as previous, 2012-2013, J. Puniamoorthy et al. leg., malaise trap, ZRC_BDP(multiple).
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[both as Pachycondyla (Ectomomyrmex) leeuwenhoeki].
Localities. Bukit Timah Nature Reserve; Bukit
Timah Road; Central Catchment Nature Reserve; Lower Peirce Reservoir; Mandai Lake
Road; Mandai Road; Nee Soon Swamp Forest.
Habitat/Ecology. This species is associated mostly
with primary or old/mature secondary forests
in Singapore, including swamp forest (drier
sections). The ants have also been found inabandoned plantation and native-dominatedsecondary forests. Nests were found in soil
substrate matrices under dead wood, also inrotting wood or branches on forest floors. Individuals were often collected from leaf litterand in soil at bases of living trees.

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Remarks. Amongst material examined for E. leeuwenhoeki, we observed substantial variation
across different samples of workers. Thesevariable features include: petiole shape (wider
and shorter in dorsal view for some individuals), strength of sculpture on petiolar node
(fovea generally weaker in some individuals),
and relative dimensions of head. Variation,
however, is subtle and can occur between intranidal workers, which sometimes present
a gradation of intermediate morphologicalstates.
		Some of the variable states we observed
are apparently diagnostic, according to original descriptions, for the two subspecies of E.
leeuwenhoeki – E. leeuwenhoeki jacobsoni(Forel, 1915) and E. leeuwenhoeki sumatrensis (Forel, 1901). But these traits seem too
unstable, at least based on local samples, toreliably differentiate between E. leeuwenhoeki
and its subspecies.
		At the time of writing, we are unable toassess the validity of both subspecies – no access to type material including type images ofjacobsoni – against E. leeuwenhoeki. For now,
we choose to err on the side of caution andrefrain from making definitive conclusions on
species status for the two forms. It remains
possible that the two subspecies might be synonymized under E. leeuwenhoeki, provided
more compelling evidence over a wider geographic range arises in future.

Material examined. Bukit Timah Nature Reserve, Jun-Oct 1968, D.H. Murphy leg.,
ZRC_ENT00027946-27948; same locality as previous, BT07, 1.35499, 103.78167,
ZRC_HYM_0000350; queen and workers,
Nee Soon Swamp Forest, 2 Oct 2017, W.
Wang leg., Winkler extraction, WT_FL1/
NSW2, ZRC_ENT00000976; same locality as previous, 1.39932, 103.80954, 13 Sep
2015, M.K.L. Wong leg., Berlese extraction,
ZRC_ENT00000713.
Material not physically examined. Unknown.
Literature. Baroni Urbani (1975) [as Belonopelta
buttelreepeni].
Localities. Bukit Timah Nature Reserve; Nee Soon
Swamp Forest.
Habitat/Ecology. This species was found inprimary or old/mature secondary forests in
Singapore, including swamp forest. Workers
were collected from soil and leaf/forest litter.

Ectomomyrmex overbecki (Viehmeyer, 1916)

Euponera malayana (Wheeler, 1929)

Material examined. Queen, locality unknown,
1994, collector unknown, NS 136A,ZRC_ENT00000277; Pulau Ubin (OBS
area), 28 May 2019, W. Wang leg., ZRC_
ENT00007921; Mandai Road, 1.40869,
103.80138, 19-21 Dec 2016, G.W. Yong et al.
leg., M1D3GPc20149, ZRC_ENT00047847
Material not physically examined. Types –CASENT0915272, ANTC40433 (NHMB);FOCOL0958-961 (ZMHB). Non-type – FOCOL0364 (DEIC) [as cotype of unpublished‘Bothroponera helleri’].
Literature. Type – Viehmeyer (1916); Overbeck(1924) [both as Pachycondyla (Ectomomyrmex) overbecki].

Material examined. Queen and worker, Mandai
Besar S3 Mangrove, 5 Aug 1987, S. Teo leg.,
ZRC_ENT00048803-48804; males, Prince
George’s Park Residences (NUS), May-Sep
2015, M.S. Foo & W. Wang leg., ZRC_BDP
multiple.
Material not physically examined. Unknown.
Literature. Wang et al. (2018a) [as Euponera
sp.1.of.WW – but see Remarks].
Localities. Mandai Besar Mangrove; Prince
George’s Park Residences (NUS).
Habitat/Ecology. A queen and worker of this species were found in an abandoned mud lobster(Thalassina sp.) mound in mangroves. Males
were identified from a malaise trap sample
from a disturbed secondary forest fragment,
in a semi-urban setting.

Localities. Bukit Timah Road [type]; Mandai
Road; Pulau Ubin.
Habitat/Ecology. The type series of this species
was found nesting between the roots of a tree
in loose earth in a garden. Individuals werealso collected from disturbed native-dominated secondary forest, on one occasion in arotting log on the ground.
Remarks. Type locality in Singapore.
Emeryopone buttelreepeni (Forel, 1912)

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Remarks. This genus appears similar to Ectomomyrmex, but can be most easily distinguishedfrom the latter by the presence of basal mandibular pits (absent in Ectomomyrmex).
		Euponera malayana can be distinguishedfrom most other congeners in Southeast Asia
by its relative smaller size (TL 4.3-4.4 mm),indistinct longitudinal subocular groove, andnear-trapezoidal petiole in profile view withanterior face weakly convex (the latter condition more evident in dorsal view).
		Based on observations of males of other
Euponera species and their worker nestmates,
we deduced that Euponera sp.1.of.WW mentioned in Wang et al. (2018a) should be E.
malayana. The male specimens mentioned
in the study correspond to those encoded‘ZRC_BDP’ under ‘Material examined’. We
observed that petioles of Euponera males
tend to resemble those of conspecific workers, albeit smaller in size with weaker edges.
Males of Euponera sp.1.of.WW seem relatively smaller than other congeneric males,
and have short, subtriangular petioles withrounded apices, much like a shrunken versionof the worker’s petiole.
		
Thus, we tentatively treat these males as
E. malayana, but this assessment may change
in future if full colony series of the species
are collected, and our initial deductions areproven otherwise.
Euponera sharpi Forel, 1901
Material examined. Queen and workers, Namly
Avenue, 28 Apr 1985, D.H. Murphy leg.,
DHM-SG85-Eup1, ZRC_ENT00027961; Pulau Ubin, 7 Jan 2014, Sk. Yamane leg., ZRC_
HYM_0000450; Upper Seletar Reservoir, 4
Oct 2020, G.W. Yong leg., GY-SG20-002,
ZRC_ENT00028567.
Material not physically examined. Syntype –CASENT0907284, ANTC27598 (MHNG).
Literature. Type – Forel (1901)[as Euponera(Pseudoponera) sharpi]. Viehmeyer (1916),
Overbeck (1924) [both as Euponera (Trachymesopus) sharpi].

Localities. Bukit Timah Road [type]; Namly Avenue; Pulau Ubin; Upper Seletar Reservoir.
Habitat/Ecology. This species was found inboth young and mature secondary forests ofvarying levels of disturbance in Singapore,including disturbed wasteland secondary forest fragments or scrub in semi-urban settings.
Individuals could be found in soil around rootsof old fallen trees, or under rocks. Colonies
were sometimes observed living together withtermites.
Remarks. Type locality in Singapore.
Harpegnathos venator (Smith, 1858)
Material examined. Sime Road, 30 Dec 1974,
D.H. Murphy leg., ZRC_ENT00007406.
Material not physically examined. Unknown.
Literature. Davidson et al. (2008).
Localities. MacRitchie; Sime Road.
Habitat/Ecology. This species is associated withprimary or mature secondary forests in Singapore, in fringe forest or near forest edges.
Individuals were found foraging on the forest
floor; nests were observed to be undergroundin soil with funnel entrances.
Hypoponera confinis (Roger, 1860)
Material examined. None available.
Material not physically examined. Unknown.
Literature. Forel (1912) [as Ponera confinis].
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore context.
Remarks. At the time of writing, this species
is known from Singapore only based on asingle literature record, in contrast with its
three original subspecies where both literatureand physical records are available. We cannot
overrule the possibility that this single record
may actually correspond to any one of thethree subspecies (two raised to species in thispaper) later described from Singapore.

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Hypoponera confinis epinotalis (Viehmeyer,
1916)
Material examined. None available.
Material not physically examined. Types –CASENT0907306, ANTC27772 (MHNG);
FOCOL0358-359 (DEIC); FOCOL0989-991(ZMHB).
Literature. Type – Viehmeyer (1916); Overbeck(1924) [both as Ponera confinis epinotalis].
Localities. Bukit Timah Road [type].
Habitat/Ecology. Workers of the type series were
found under a flower pot in a garden.
Remarks. Type locality in Singapore.
Hypoponera javana (Forel, 1905) stat. n.
Ponera confinis var. javana Forel, 1905: 6 (w.q.)
Combination in Hypoponera: Bolton, 1995: 215.
Subspecies of Hypoponera confinis: Forel, 1907a:
17; Forel, 1911a: 194; Emery,con 1911: 90;
Wheeler, 1919: 57; Wheeler, 1924: 242;
Wheeler & Chapman, 1925: 69; Wheeler,
1937: 21; Chapman & Capco, 1951: 69; Baltazar, 1966: 246; Bolton, 1995: 215; Pfeiffer etal., 2011: 56.
Material examined. Kent Ridge, 25 Apr 1985,
D.H. Murphy leg., DHM-SG85-Hypo4, ZRC_
ENT00027998; Mandai Lake Road, NorthernNode, 1°24’36.5”N, 103°47’04.0”, 53 m, 18
Jul 2019, J.S. Tan & N.L. Chin leg., Winkler
extraction, ZRC_ENT00028303; queen andworkers, Namly Avenue (kampong), 28 Apr
1985, D.H. Murphy leg., DHM-SG85-Hypo3,
ZRC_ENT00027997.
Material not physically examined. Unknown.
Literature. Forel (1907) [as Ponera confinis javana].
Localities. Kent Ridge; Mandai Lake Road; Namly Avenue.
Habitat/Ecology. This species was found in youngor mature secondary forest, often disturbed
fragments in semi-urban settings close to
man-made infrastructure. In one instance, theants were found nesting under brick. Usually,
individuals were collected from leaf litter andtopsoil substrate.

Remarks. Hypoponera javana was originally described as a subspecies of H. confinis (thenknown as ‘Ponera confinis’). Workers of both
species appear very similar, but can be distinguished based on the following (H. confinischaracters in parentheses): 1) head wider withbroadly convex sides, slightly narrowed anteriorly (head less wide with almost parallelsides, not narrowed anteriorly), 2) antennal
scapes reaching posterior margin of head (antennal scapes not reaching posterior margin ofhead), 3) petiole shorter in dorsal view, thinner
in profile (petiole longer and nearly reniform
in dorsal view, thicker in profile), 4) propodeal
dorsum relatively short, not much longer thanwide, distinctly shorter than declivitous slope
in profile (propodeal dorsum long, distinctly
longer than wide, as long as or slightly longer
than propodeal declivity).
		The queen of H. javana can also be separated from that of H. confinis by its more
distinctly angulate propodeal junction andlonger, more sharply truncate propodeal slope.
		Hypoponera javana closely resembles
Hypoponera singaporensis Viehmeyer, 1916
stat. n., which was also originally described
as a subspecies of H. confinis. Differences
between the two species are discussed under
Remarks for H. singaporensis.
Hypoponera malayana (Wheeler, 1929)
Material examined. Mandai Lake Road, NorthernNode, 1°24’36.5”N, 103°47’04.0”, 53 m, 18
Jul 2019, J.S. Tan & N.L. Chin leg., Winkler
extraction, ZRC_ENT00048408.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Mandai Lake Road
Habitat/Ecology. Individuals of this species werecollected in leaf litter and upper soil substratefrom mature secondary forest.
Remarks. At the time of writing, since both typeimages and physical specimens of this species
were unavailable for comparison, identities of
ZRC material examined were deduced basedon the original species description by Wheeler(1929).

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Hypoponera pygmaea (Forel, 1907)
Material examined. Bukit Timah Nature Reserve, BT03, 1.34913, 103.77914, 17 May
2017, W. Wang leg., WW-SG17-056, ZRC_
ENT00028316; Nee Soon forest, 26 Apr 2018,
W. Wang leg., ZRC_ENT00047964.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Nee Soon
forest.
Habitat/Ecology. This species was found mostly
in old secondary forest, on one occasion insoil at the base of a large tree.
Remarks. This tiny species (< 1.5 mm) can be
distinguished from most other congeners by
its minute size and distinctly long antennal
club. The original description states that the‘last article of the antenna’, presumably the
apical segment of the club, is about as longas the combined length of its preceding ninesegments. In local specimens, we observed theapical antennal segment to be slightly shorter.
		Workers of Hypoponera pygmaea arealso similar to those of another minute species – Hypoponera butteli (Forel, 1913), but
can be differentiated by their shorter scapes
which are distant from posterior margin ofhead when lain back. In H. butteli, the scapes
nearly reach the posterior margin of head.
Hypoponera singaporensis (Viehmeyer, 1916)
stat. n.
Ponera confinis var. singaporensis Viehmeyer,
1916: 115 (w.q.)
Combination in Hypoponera: Bolton, 1995: 216.
Subspecies of Hypoponera confinis: Chapman &Capco, 1951: 69; Bolton, 1995: 216.
Material examined. Labrador beach, 29 Apr
1985, D.H. Murphy leg., DHM-SG85-Hypo1,
ZRC_ENT00027995.
Material not physically examined. Types – FOCOL0356-357 (DEIC); queen and workers,
FOCOL0985-988 (ZMHB).
Literature. Type – Viehmeyer (1916). Overbeck(1924) [both as Ponera confinis singaporensis].
Localities. Bukit Timah Road [type]; Labrador
beach.

Habitat/Ecology. Type specimens of this species
were found in a garden, specifically in the wet
pulp of a large fungus on a termite-infested
tree, also in the rotten wood of a decaying
branch. Individuals were also collected froma considerably rocky beach, in close proximity
to coastal beach forest.
Remarks. Type locality in Singapore. Originallydescribed as a subspecies of H. confinis, workers of H. singaporensis can be differentiatedfrom those of the former by points (1) – (3)mentioned in Remarks under H. javana. In addition, H. singaporensis workers are darker incolour and visibly more shiny than those of H.
confinis. The propodeal junction in H. singaporensis workers is also less sharply angularand more ill-defined than that of H. confinis,
because of a slightly bulging sclerite.
		Hypoponera singaporensis can be distinguished from H. javana based on the following (H. javana traits in parentheses): 1)
in workers, propodeum distinctly longer thanwide, propodeal dorsum as long as or slightly
longer than declivitous slope in profile (propodeal dorsum relatively shorter, not much
longer than wide, propodeal dorsum shorter
than declivitous slope), 2) in queens, propodeal junction mirrors that of workers, rounded
and ill-defined (propodeal junction distinctly
angulate and well-defined).
Hypoponera truncata (Smith, 1860)
Material examined. Nee Soon Swamp Forest,
17 Dec 2021, W.N. Lam et al. leg., ZRC_
ENT00057832 – 57833.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Nee Soon Swamp Forest
Habitat/Ecology. This species was found in freshwater swamp forest, collected from leaf litterand soil debris.
Remarks. Hypoponera truncata was first described based on a holotype queen (Smith
1860). At the time of writing, both reliablyidentified worker images and physical specimens of this species were unavailable forcomparison. Thus, the identities of ZRC material examined (all workers) were deduced
based on the original worker description by

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Forel (1905), and the holotype queen image
on AntWeb. These specimens should be considered Hypoponera truncata sensu lato.
		This blackish species resembles Hypoponera elliptica (Forel, 1900) from Australia,
but is generally smaller (worker TL 2.7-3.1mm, see Forel 1905), has longer antennal
scapes that visibly exceed the posterior marginof head, more strongly convex sides of head,and shorter propodeal dorsum.
Hypoponera zwaluwenburgi (Wheeler, 1933)
Material examined. University campus ‘floodplain’ (sic), 14 Aug 1964, D.H. Murphy leg.,
ZRC_ENT00027944.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. University campus (Bukit Timah)
Habitat/Ecology. It is not clearly understood frominformation on the original ZRC specimenlabel what type of habitat this species was
associated with. A ‘flood plain’ is technicallydefined as an area of low-lying ground adjacent to a river or stream, formed largely of silt
and sediment and subject to flooding. However, there were no known major water bodies
within said university campus grounds at thetime of specimen collection. The most we caninfer from locality data is that the species was
possibly found in a disturbed secondary forest
fragment in an urban setting/matrix.
Leptogenys borneensis Wheeler, 1919
Material examined. Bukit Timah Nature Reserve,
11 Sep 1967, D.H. Murphy leg., A82-13,
DHM-SG67-Lep2, ZRC_ENT00047690.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve
Habitat/Ecology. Individuals were collected fromprimary or old/mature secondary forest in Singapore.

Leptogenys chinensis (Mayr, 1840)
Material examined. Mandai, Bird Park Buffer, 17 Jul 2019, J. Tan & N. Chin leg.,
JTNC-MIS-003, ZRC_ENT00013544;
Prince George’s Park Residences, 1.292389,
103.7787, 3 Dec 2016, W. Wang leg., WWSG16-013, ZRC_HYM_0000506.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Mandai; Prince George’s Park Residences (NUS).
Habitat/Ecology. This species was found in disturbed secondary forest fragments in semiurban settings. The ants were found nestingin fallen rotting branches or wood, in one instance alongside a Gnamptogenys nest.
Leptogenys diminuta (Smith, 1857)
Material examined. Males and workers, Bukit
Timah Nature Reserve, 30 Sep 1964, D.H.
Murphy, A-292-11, DHM-SG64-Lep1, ZRC_
ENT00027949; same locality and collector as
previous, Fern Valley contour, 14 May 1967,
DHM-SG67-Lep1, ZRC_ENT00027950;
same locality as previous, 1965, collector
unknown, ZRC_ENT00048393.
Material not physically examined. Unknown.
Literature. None. New record (see Remarks).
Localities. Bukit Timah Nature Reserve
Habitat/Ecology. This species is associated withprimary or mature secondary forests in Singapore. Individuals were found in rotting wood,
under bark of fallen dead trees, or in leaf litterand soil debris.
Remarks. Supposed historical literature recordsof this species in Singapore (i.e., Viehmeyer
1916, Overbeck 1924) actually reported specimens from Gunong Angsi in Malaysia, notSingapore. This error has unfortunately been
perpetuated in contemporary online databases
and some secondary literature sources, e.g.
Xu & He (2015).

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Leptogenys hemioptica Forel, 1901
Material examined. Bukit Timah Nature Reserve,
1 May 1967, D.H. Murphy leg., C2-3, DHMSG67-Lep3, ZRC_ENT00047820; same locality as previous, plot II-2, 30 May 2016, D.J.
Court leg., pitfall trap, ZRC_ENT00048383;
same locality as previous, 1 Apr 1967, D.H.
Murphy leg., ZRC_HYM_0000680.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve
Habitat/Ecology. This species was found in either
primary or old/mature secondary forest, typically on the ground or in forest litter (leaves
and soil debris).
Leptogenys iridescens (Smith, 1857)
Material examined. Bukit Timah Nature Reserve,
1.34772, 103.77766, collector unknown, 30
Nov 2015, ZRC_ENT00000714; same locality as previous, 18 May 1969, D.H. Murphyleg., A24-10, ZRC_HYM_0000672; same
locality and collector as previous, 4 October
1973, ZRC_HYM_0000676; same locality
and collector as previous, 8 Jun 1967, ZRC_
HYM_0000874; Central Catchment Nature
Reserve, MacRitchie, 10 Sep 2011, E.J.Y. Sohleg., ZRC_ENT00000858; Nee Soon forest,
1.37770, 103.80645, 3 Aug 2017, W.N. Lamleg., NS02_04, ZRC_ENT00000859; Seletar Trail, 1.3946, 103.80113, 28 Aug 2015,
M.K.L. Wong leg., Berlese extraction, ZRC_
ENT00000715.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Central
Catchment Nature Reserve (MacRitchie); NeeSoon forest; Seletar Trail.
Habitat/Ecology. This species is associated withprimary or old/mature secondary forests in
Singapore, including swamp forest whereindividuals were found in the lower pitcher
contents of Nepenthes ampullaria. The ants
were also collected from leaf litter, and on
forest paths or trails.

Remarks. At the time of writing, most examinedmaterial in the ZRC were expert-identified
to the ‘L. iridescens complex’ – broad morphological variation was observed between
different samples. Some specimens had characters that would more closely confer with L.
mutabilis sensu lato (W. Wang 2021, pers.
obs.). In the absence of more compelling empirical evidence supporting different species
hypotheses, we tentatively treat these variable
specimens as conspecific for the purposes ofthis checklist.
Leptogenys kraepelini Forel, 1905
Material examined. Alate queens, males, andworkers, National University of Singapore(multiple sites), Apr-Sep 2015, M.S. Foo& W. Wang leg., malaise trap, ZRC_BDP(multiple); National University of Singapore
campus, 13 Oct 2016, P. Escoubas leg., EscoSG16-SG005, ZRC_ENT00047817-47819;
Pulau Ubin, 15 Oct 2016, P. Escoubas leg.,
SG016-18, ZRC (uncatalogued); Seletar, 18
Oct 2016, same collector as previous, SG023,
ZRC (uncatalogued).
Material not physically examined. Singapore
Botanic Gardens, 5 Aug 2014, J.K. Wetterer
leg., vial #477 (JKWC).
Literature. Wang et al. (2018a).
Localities. National University of Singapore campus (Clementi/Kent Ridge); Pulau Ubin; Seletar; Singapore Botanic Gardens.
Habitat/Ecology. This species was often foundmostly in disturbed secondary forest fragments in urban or semi-urban settings, including waste woodland forest. Numerous alates
were collected from cultivated grass patches
or scrubland in urban settings using malaisetraps. Nests were sometimes found in fallen
rotting wood.
Remarks. Broad morphological variation wasobserved between samples determined as L.
kraepelini sensu lato, suggesting possible
cryptic species or a species complex. Pending
the availability of empirical evidence proving
multiple species, for this checklist we tentatively treat all variants as conspecific.

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Leptogenys mutabilis (Smith, 1861)
Material examined. Lower Peirce Reservoir,
7 Aug 1990, collector unknown, ZRC_
ENT00000121, ZRC_ENT00000126; same
locality as previous, 7 May 2017, G.W. Yongleg., ZRC_ENT00048418; Mandai Lake
Road, 5 Nov 2016, G.W. Yong leg., ZRC_
ENT00048423; male and workers, Mandai
Road, 13 Jan 2016, G.W. Yong leg., GYSG16-Lep1, ZRC_ENT00048407; Nee Soon
Swamp Forest, 26 Apr 2018, W. Wang leg.,
WW-SG18-Lepto1, ZRC_ENT00000950.
Material not physically examined. Unknown[but see Remarks under ‘L. processionalis’].
Literature. None. New record.
Localities. Lower Peirce Reservoir; Mandai Lake
Road; Mandai Road; Nee Soon Swamp Forest.
Habitat/Ecology. This species appears associatedmostly with primary and old/mature secondary forests, including swamp forest where anest was found in a mud mound on waterlogged ground.
Remarks. This species is similar to L. iridescens,but may be distinguished from the latter by
a few characters such as: 1) head in full faceview more squarish, only slightly longer thanwide (in L. iridescens distinctly longer thanwide); 2) lateral margins of head in full faceview evenly and broadly convex (in L. iridescens the sides are almost straight); 3) eye
farther from lateral head margin in full faceview (in L. iridescens the eye is about half theformer distance from lateral head margin);4) antennal scape relatively thicker across its
entire length (more ‘flattened’ laterally) in
proportion to head width.
Leptogenys myops (Emery, 1887)
Material examined. Central Catchment Nature
Reserve (Venus Loop), 1.35550, 103.81577,
15 Jun 2015, M.K.L. Wong leg., pitfall trap,
ZRC_ENT00000717; Lim Chu Kang mangrove, 10 Mar 1987, D.H. Murphy leg.,
ZRC_ENT00000886; Seletar Trail, 1.39192,
103.80081, 18 Oct 2015, M.K.L. Wong leg.,
pitfall trap, ZRC_ENT00000716; Sunset Way,1.32609, 103.77187, 22-24 Aug 2016, G.W.
Yong leg., pitfall trap, SW2GP023, ZRC_
HYM_0001188.2.

Material not physically examined. Unknown.
Literature. None. New record.
Localities. Central Catchment Nature Reserve;
Lim Chu Kang mangrove; Seletar Trail; Sunset Way.
Habitat/Ecology. This species was found in bothyoung and mature secondary forests, suchas an abandoned plantation forest, and also
in a mangrove, where individuals were collected from atop a mud lobster (Thalassinasp.) mound.
Leptogenys pangui Xu, 2000
Material examined. Bukit Timah Nature Reserve,collection date unknown, D.H. Murphy leg.,C1-19, ZRC_HYM_0000681; Sime Road,
15 Dec 1975, D.H. Murphy leg., ZRC_
HYM_0000907.
Material not physically examined. Unknown.
Literature. None. New record (but see Remarks).
Localities. Bukit Timah Nature Reserve; SimeRoad.
Habitat/Ecology. This species was found in either
primary or old/mature secondary forest.
Remarks. The Singapore specimens may in fact
be a closely-related but different species withcharacters intermediate between L. pangui
and the smaller L. assamensis. General characters of the ZRC material, however, confer
more with the original description of theformer species, thus we tentatively treat thismaterial as L. pangui. This treatment may possibly change in future when more empiricalevidence is made available.
Leptogenys peuqueti (André, 1887)
Material examined. Bukit Timah Nature Reserve, 8 Sep 2005, collector unknown, ZRC_
HYM_0000276; same locality as previous,
collection date unknown, D.H. Murphy leg.,C2-1, ZRC_HYM_0000683; same locality
as previous, 25 Dec 1989, H.K. Lua leg.,
ZRC_ENT00000242; Labrador beach, 29 Apr
1985, D.H. Murphy leg., DHM-SG85-Lep1,
ZRC_ENT00047823; Lower Peirce Reservoir Road, Aug 1990, collector unknown,
ZRC_ENT00000153; Mandai Lake Road, N
01°24’19.3’’, E 103°46’56.2’’, 73 m, 26 Mar

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

2020, W. Zhang leg.,WZ_MIS_020, ZRC_
ENT00047799; Rifle Range Road, pipeline
track, 26 Jan 1986, D.H. Murphy leg., DHMSG86-Lep1, ZRC_ENT00047822; beside
Temasek Polytechnic, 1.34355, 103.93572,
13-15 Dec 2016, G.W. Yong et al. leg., TT2GPd561, ZRC_ENT00047862; Prince George’s
Park Residences (NUS), 6 Dec 2017, Sk. Yamane leg. (SKYC).
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924)[both as Leptogenys (Lobopelta) peuqueti].
Localities. Bukit Timah Nature Reserve; Bukit
Timah Road; Labrador beach; Lower Peirce
Reservoir Road; Prince George’s Park Residences (NUS); Mandai Lake Road; Rifle
Range Road; Temasek Polytechnic (adjacentwaste woodland).
Habitat/Ecology. This species is associated mostly
with primary or old/mature secondary forests,
sometimes also in young secondary forest including coastal beach forest, and cultivatedgardens in Singapore. Individuals and nests
have been sampled from inside fallen rotting
or decayed wood, and logs.
Leptogenys pompiloides (Smith, 1857)
Material examined. None available.
Material not physically examined. Male,CASENT0281941, ANTC20700 (BMNH).
Literature. Type – Smith (1857) [as Ponera pompiloides; see Remarks].
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore context.
Remarks. This species was first described basedon males apparently collected from Singapore,though type locality was indicated as ‘Borneo(Sarawak)’ in Smith (1857). The worker forthis species remains a mystery till present day.
Donisthorpe (1932) concluded that the locality labels on the type male specimens were
probably miswritten, since no males in the
A.R. Wallace collections were associated with
Sarawak, but two males linked to Singapore
matched the original description of L. pompiloides [as Ponera pompiloides].

Leptogenys processionalis (Jerdon, 1851)
Material examined. None available.
Material not physically examined.
CASENT0217534, ANTC17790 (CASC).
Literature. None. New record.
Localities. Bukit Timah Nature Reserve
Habitat/Ecology. This species was found in either primary or old/mature secondary forest
in Singapore.
Remarks. The unexamined specimen at CASC
might be misidentified L. mutabilis – uponobservation of the full face image on AntWeb,
it appears that there are no striations on theanterior one-third of the head dorsum, a feature diagnostic of L. processionalis.
Mesoponera rubra (Smith, 1857)
Material examined. Bukit Timah Nature Reserve, 1.35499, 103.78167, 19 Jul 2017,
W. Wang leg., WW-SG17-050, ZRC_
ENT00027968; same locality as previous,
1.35130, 103.77859, 31 Jul 2015, M.K.L.
Wong leg., SPT15, ZRC_ENT00000720;
same locality as previous, 28 Apr 2016, D.J.
Court leg., ZRC_ENT00048450; locality andcollection date unknown, D.H. Murphy leg.,
ZRC_HYM_0000798; locality and collection date unknown, D.H. Murphy leg., ZRC_
HYM_00000832.
Material not physically examined. Holotype
queen – CASENT0901349, ANTC20788(OUMNH).
Literature. Type – Smith (1857) [as Ponera rubra]. Emery (1893, 1911) [as Ponera rubra].
Localities. Bukit Timah Nature Reserve
Habitat/Ecology. This species is associated withprimary or old/mature secondary forests in
Singapore. Individuals were sometimes foundin soil at base of large trees, or collected using
pitfall traps.
Remarks. Type locality in Singapore, but specificlocation unknown. This species was first described based on a dealate queen from Singapore.

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Mesoponera javana (Forel, 1905) stat. n.
Euponera (Mesoponera) rubra var. javana Forel,
1905: 6 (q.m)
Combination in Pachycondyla: Brown, 1995: 306.
Combination in Mesoponera: Schmidt & Shattuck,
2014: 111.
Subspecies of Mesoponera rubra: Emery, 1911:
81; Wheeler, 1924: 242; Karavaiev, 1926:
419; Chapman & Capco, 1951; Bolton, 1995:
306.
Senior synonym of Mesoponera rubra minirubra:Özdikmen, 2010 (unnecessary replacement
name)
Material examined. Queen and worker, Lower Peirce, 7 May 2017, G.W. Yong leg.,
ZRC_ENT00027967; Bukit Timah forest,
26 Nov 1969, D.H. Murphy leg., ZRC_
HYM_0000832; Upper Thomson Nature
Park, 19 Oct 2016, G.W. Yong et al. leg.,
ZRC_ENT00057834.
Material not physically examined. Unknown
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Lower
Peirce; Upper Thomson Nature Park.
Habitat/Ecology. Individuals were found in either
primary or old/mature secondary forests in
Singapore, usually in leaf litter or in soil atbases of trees. In one instance, workers werecollected under bark of a rotting log, in wet
sand.
Remarks. Mesoponera javana resembles M. rubra, but is generally smaller – the queen of M.
javana is either similar in size to or smaller
than the worker of M. rubra. The workers of
M. javana can also be differentiated from M.
rubra based on the following (M. rubra traits
in parentheses): 1) median part of clypeus
produced anteriorly as a broad, subtriangular lobe with median apical point and distinct
rounded, translucent lamellate edge (medianpart of clypeus weakly produced anteriorly
as a shallow, broadly convex lobe with indistinct lamellate edge), 2) eye relatively larger
and closer to anterior margin of head, shortest
distance of eye to mandibular insertions lessthan 2x eye length (eye relatively smaller andfurther away from anterior margin of head,
shortest distance of eye to mandibular insertion more than or equal to 2x eye length).

		Condition of the clypeus, i.e., character(1) previously mentioned, can be used to differentiate between queens of the two species
as well, besides body size.
Myopias mayri (Donisthorpe, 1932)
Material examined. Mandai forest, 7 Jun
2018, W. Wang leg., ZRC_ENT00007263.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Mandai
Habitat/Ecology. A worker was collected fromfallen rotting wood in secondary forest.
Odontomachus litoralis Wang, Yamada & Yamane, 2020
Material examined. Types – holotype, Sungei
Buloh Wetland Reserve, 28 Mar 2018, W.
Wang & M.S. Foo leg.,WW-SG18-Odonto1,
ZRC_ENT00013883; paratype male, queens
and workers, same collection data as holotype, ZRC_ENT0000917. Non-types – Admiralty, 6 Apr 2019, Benjamin Yeap leg., ZRC_
ENT00048015; Lim Chu Kang mangrove,
1987, Serena Teo leg., ZRC_ENT00000778;
Loyang mangroves, 29 Nov 2002, N. Sivasothileg., ZRC_ENT00027921; Mandai mangroves,
1 Apr 2009, S.P. Goh & D. Pitta de Araujo leg.,
ZRC_ENT00000774; workers and males, same
locality as previous, 15 Oct 1978, D.H. Murphyleg., DHM-SG78-Odo1, ZRC_ENT00027918;
workers and males, same locality and collector
as previous, 18 Oct 1977, DHM-SG77-Odo1,
ZRC_ENT00027920; Pulau Ubin, Celestial
Resort Ubin, 6 Mar 2012, I. Iesa leg., CMBSM01, ZRC_ENT00007278; males, queens
and workers, Sungei Buloh Wetland Reserve,
5 Dec 2018, W. Wang et al. leg.,WW-SG18Odonto3/4, ZRC_ENT00007635-7636; males,
queens and workers, same locality as previous,
20 Dec 2018, W. Wang & M.S. Foo leg., WWSG18-Odonto6/7, ZRC_ENT00007645-7646.
Material not physically examined. Unknown.
Literature. Type – Wang et al. (2020).
Localities. Admiralty; Lim Chu Kang mangroves;
Loyang mangroves; Mandai mangroves; Pulau
Ubin; Sungei Buloh Wetland Reserve [type].

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Habitat/Ecology. This species is found exclusively in mangroves in Singapore. Nests
were usually found in abandoned Thalassina(mud lobster) mounds, sometimes well above
maximum tide levels. Nests have also beenreported from soil mounds among roots of
uprooted trees. Workers were often observed
foraging on mud and debris in close proximity to mangrove back forests during low tide.
Remarks. Type locality in Singapore.
Odontomachus malignus Smith, 1859
Material examined. Lim Chu Kang mangrove, 23
Sep (year unknown), D.H. Murphy leg., ZRC_
HYM_0000902; males, Pulau Semakau Old
Fragment, 1°12’23.9”N, 103°45’37.6”E, JulNov 2012, J. Puniamoorthy et al. leg., malaise
trap, ZRC_BDP (multiple); male, Pulau Semakau New Fragment, SMN2, 1°12’04.5”N,
103°45’46.1”E, 6-13 Dec 2012, same collectors as previous, malaise trap, Reg. 29626,
ZRC_BDP0016086.
Material not physically examined. Unknown.
Literature. Wang et al. (2020).
Localities. Lim Chu Kang mangrove; Pulau Semakau.
Habitat/Ecology. A worker of this species wasfound foraging in mangrove at low tide, close
to mudflats opening to the sea. Nests havenever been found in mangroves in Singapore.
Most specimens were males collected in malaise traps set up in mangroves at an offshore
landfill. It is possible that these males could
have originated from actual nests in extensive
coral rubble surrounding the mangrove fragments on this landfill.
Remarks. This species is similar to O. litoralis,but can be differentiated from the latter by theabsence of a strongly carinate posterior propodeal edge in the worker, and also a generally
less coarse or shagreened, somewhat shinier
mesosoma. Odontomachus malignus has beenreported to nest in coastal coral rubble andlimestone rockfaces facing the sea in othercountries – this apparent nesting preference
may also distinguish it from O. litoralis which
has only been found nesting in mangroves.

Odontomachus rixosus Smith, 1857
Material examined. Bukit Batok East, 1.34304,
103.76235, Sep 2016, G.W. Yong leg., ZRC_
HYM_0001294-1295; Bukit Timah Nature
Reserve, 19 Feb 2016, collector unknown,
ZRC_ENT00000723; same locality as previous, 1.35386, 103.77988, 23 Aug 2017, W.
Wang leg., ZRC_ENT00007559; same locality as previous, 15 Aug 1983, D.H. Murphyleg., ZRC_HYM_0000669; same locality as
previous, plot III-2, 19 Apr 2016, D.J. Court
leg., pitfall trap, ZRC_ENT00048454; males,
Bukit Timah Summit, 13-22 Apr 2009, Z.
Chen leg., malaise trap, ZRC_ENT0004802048022; Catchment forest, 15 Sep 1964, D.H.
Murphy leg., ZRC_HYM_0000663; Lower
Peirce Reservoir, 11 Aug 1990, collector unknown, ZRC_ENT00000109; same locality
as previous, 7 Aug 1990, collector unknown,
ZRC_ENT00000122; same locality as previous, 31 Jul 1990, collector unknown, ZRC_
ENT00000209; MacRitchie Reservoir, 9 Jan
2014, Sk. Yamane leg., ZRC_HYM_0000424;
Mandai, 1.40661, 103.78848, 13 Nov 2018,
W. Wang leg., ZRC_ENT00007586; Mandai Road, Nov-Dec 2016, G.W. Yong leg.,
ZRC_HYM_0001303-1322; male, Pulau
Ubin, PU1, 1°24’36.3”N, 103°59’25.5”E, 2330 Jun 2012, J. Puniamoorthy et al. leg., malaise trap, Reg. 29319, ZRC_BDP0016035;
Seletar Trail, 1.39097, 103.79952, 7 Dec
2015, M.K.L. Wong leg., Winkler extraction,
ZRC_ENT00000722; Upper Thomson Nature
Park, 1.38311, 103.79839, Oct 2016, G.W.
Yong leg., ZRC_HYM_0001298-1302.
Material not physically examined. Syntype – CASENT0900656, ANTC19758(BMNH). Non-types – ANIC32-031703,
RW Ta c c . 6 8 . 11 ( A N I C ) ; A N I C 3 2 031673, Shattuck134500711 (ANIC);
CASENT0217544, ANTC17803 (CASC);FMNHINS0002821936, FMHD_81-282(FMNH).
Literature. Type – Smith (1857). Viehmeyer(1916); Overbeck (1924); Tan & Corlett(2012); Satria et al. (2015).

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Localities. Bukit Batok East; Bukit Timah Nature
Reserve; Bukit Timah Road; Lower Peirce
Reservoir; MacRitchie Reservoir; Mandai;
Mandai Road; Pulau Ubin; Seletar Trail;
Singapore Botanic Gardens; Upper Thomson
Nature Park.
Habitat/Ecology. This species is associated mostly with primary and old/mature secondary forests in Singapore, but has sometimes also beenfound in more degraded or disturbed young
secondary forests, including mangrove backforest. Individual foragers have occasionallybeen collected from under bushes beachside,
presumably originating from adjacent coastal
beach forest. These ants were often found foraging in and around leaf litter.
Remarks. Type locality in Singapore.
Odontomachus simillimus Smith, 1858
Material examined. Queen, Bukit Timah, Hindhede Drive, 12 Nov 1989, H.K. Lua leg.,
ZRC_HYM_0000133; Commonwealth Road,
1.31115, 103.78171, 22-24 Nov 2016, G.W.
Yong leg., ZRC_HYM_0001329-1333; Hougang Avenue 4, 1.34561, 103.88884, 12-14 Dec
2016, G.W. Yong leg., ZRC_HYM_00013341338; queen, Jurong East, 27 Jul 2018, D. Safaruan leg., ZRC_ENT00007417; Labrador
beach, 29 Apr 1985, D.H. Murphy leg., ZRC_
HYM_0000670; Loyang mangroves, 29 Nov
2002, N. Sivasothi leg., ZRC_ENT00027922;
male, Block 78 Marine Drive, 5 Nov 2016,
W. Wang leg., ZRC_ENT00000895; male,
Pulau Semakau New Fragment, SMN3,
1°12’05.6”N, 103°45’46.5”E, 13-20 Sep
2012, J. Puniamoorthy et al. leg., malaise trap,
Reg. 29471, ZRC_BDP0015375; male, Pulau
Semakau Old Fragment, SMO2, 1°12’20.2”N,
103°45’35.8”E, 18-25 Oct 2012, same collectors as previous, malaise trap, Reg. 29538,
ZRC_BDP0016524; Pulau Ubin, 1.40308,
103.96924, 4 Dec 2016, G.W. Yong et al.
leg., ZRC_HYM_0001083; Seletar Link,
1.40518, 103.88555, 16-18 Oct 2016, G.W.
Yong leg., ZRC_HYM_0001339-1344; Sunset Way, 1.32609, 103.77187, 21-23 Sep 2016,
ZRC_HYM_0001323-1325; Temasek Polytechnic, adjacent waste woodland, 1.34239,
103.93598, 13-15 Dec 2016, G.W. Yong leg.,
ZRC_HYM_0001326-1328.

Material not physically examined. Nontypes, ANIC32-031799 (ANIC);
CASENT0009624/9624-D01/9624-D02,
ANTC7554 (CASC). Non-types, J.K. Wetterer leg. (JKWC) – Katong, Amber Road,
28 Jul 2014, vial #350; Marine Parade, 28 Jul
2014, vials #355, 371; City Hall, by church,
29 Jul 2014, vial #377; Harbourfront, 29 Jul
2014, vial #385; Dhoby Ghaut, Istana Park, 30
Jul 2014, vial #403; Tai Seng, by MRT station,
31 Jul 2014, vial #420; Katong, 1 Aug 2014,
vials #445, 449; Buona Vista, by MRT station,
5 Aug 2014, vial #486; Pasir Panjang, by MRT
station, 5 Aug 2014, vial #484.
Literature. Tan & Corlett (2012); Satria et al.
(2015).
Localities. Amber Road (Katong); Bukit Timah;
Buona Vista; City Hall; Commonwealth Road;
Dhoby Ghaut; Harbourfront; Hougang Avenue 4; Jurong East; Katong; Labrador beach;
Loyang mangroves; Marine Drive; Marine
Parade; Pasir Panjang; Pulau Semakau; Pulau
Ubin; Seletar Link; Singapore Botanic Gardens; Sunset Way; Tai Seng; (beside) Temasek
Polytechnic.
Habitat/Ecology. This species was found in awide range of habitats of varying quality andlevels of disturbance, including abandoned
plantation and waste woodland secondary forests, mangroves and beachside coastal forest.
Individuals including winged alates have also
been frequently found in or close to urban
residential areas and cultivated gardens orparks. Nests were found in soil at the bases ofliving trees, or in rotting wood or under rocks.
Odontoponera denticulata (Smith, 1858)
Material examined. Ang Mo Kio Garden West,
Jul-Nov 2013, Klara M.C. Chua leg., NT18B, ZRC_ENT00027904; Bedok Town Park,
same collection date and collector as previous, DT1-5A, ZRC_ENT00027903; Bukit
Batok East, 1.34304, 103.76235, 27-29 Sep
2016, G.W. Yong leg., ZRC_HYM_0001349;
Bukit Timah, Hindhede Drive, Dec 1989,
H.K. Lua leg., ZRC_ENT00000250; Choa
Chu Kang Avenue 2, 1 Mar 2017, D.J. Court
leg., ZRC_HYM_0001724; Commonwealth
Road, 1.31115, 103.78171, 22-24 Nov 2016,

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

G.W. Yong leg., ZRC_HYM_00013501356; Kent Ridge, 1 Jun 1985, D.H. Murphyleg., ZRC_HYM_0000817; Kranji Road,
1.42638, 103.75413, 8-10 Nov 2016, G.W.
Yong leg., ZRC_HYM_0001362-1371; Linden Drive, 27 Oct 1974, D.H. Murphy leg.,
ZRC_HYM_0000818; MacRitchie Reservoir, 13 Apr 1993, collector unknown,
NS37, ZRC_HYM_0000198; Mandai Road,
1.41119, 103.80513, 29 Nov-1 Dec 2016,
G.W. Yong leg., ZRC_HYM_0001357;
males, Prince George’s Park Residences(NUS), Jun-Aug 2015, M.S. Foo & W. Wangleg., malaise trap, ZRC_BDP0045703,47328; Pulau Ubin, 1.41662, 103.99443, 4
Dec 2016, G.W. Yong & M.K.L. Wong leg.,
ZRC_HYM_0001095; Seletar Link, 1.40518,
103.88555, 18-25 Oct 2016, G.W. Yong leg.,
ZRC_HYM_0001358-1361; Sunset Way,1.32609, 103.77187, 29-31 Aug 2016, G.W.
Yong leg., ZRC_HYM_0001345; male, Telok
Blangah Street 31, 28 Apr 2019, N. Chinleg., ZRC_ENT00013541; Upper Peirce
Reservoir, 10 Jan 2014, Sk. Yamane leg.,
ZRC_HYM_0000410-411; queen and workers, Upper Thomson Nature Park, Oct 2016,
G.W. Yong leg., ZRC_HYM_0001372-1385.
Material not physically examined. Holotype
queen – CASENT0900664 (BMNH). Nontypes, J.K. Wetterer leg. (JKWC) – City Hall,by church, 29 Jul 2014, vial #377; Outram
Park, by MRT station, 29 Jul 2014, vial #379.
Literature. Type – Smith (1858) [as Ponera denticulata; see Remarks on erroneous type locality indicated in the original description].
Yamane (2009); Tan & Corlett (2012); Wanget al. (2018a).
Localities. Ang Mo Kio Garden West; Bedok
Town Park; Bukit Batok East; Bukit Timah;
City Hall; Choa Chu Kang; Commonwealth
Road; Kent Ridge; Kranji Road; Linden
Drive; MacRitchie Reservoir; Mandai Road;
Outram Park; Prince George’s Park Residences (NUS); Pulau Ubin; Seletar Link; Sunset
Way; Telok Blangah; Upper Peirce Reservoir;
Upper Thomson Nature Park.
Habitat/Ecology. This species seemed common incultivated vegetation patches and/or disturbed
forest fragments and fringes in urban or semiurban settings, including urban residential ar-eas and parks. Individuals were also collectedfrom both mature and young secondary forests, such as abandoned plantation and waste
woodland forests, usually while foraging at
ground level. The species has sometimes beencollected from open grassland or scrub.
Remarks. Type locality in Singapore – this species was first described based on the holotype
queen from Singapore. The type locality forO. denticulata was indicated as ‘South Africa:
Cape of Good Hope’ in its original description (see Smith 1858). However, this was laterclarified as an error due to mislabelling of thespecimen (Donisthorpe 1943).
Odontoponera transversa (Smith, 1857)
Material examined. Queen and workers, Bukit
Batok East, 1.34304, 103.76235, Aug-Oct
2016, G.W. Yong leg., pitfall trap, ZRC_
HYM_0001390-1395; Bukit Timah Nature Reserve, 8 Sep 2005, collector unknown, ZRC_
HYM_0000047-50; same locality as previous,
near BT07, 1.35499, 103.78167, 19 Jul 2017,
W. Wang leg., ZRC_ENT00047930; same locality as previous, plot III-2, 12 Oct 2015, D.J.
Court leg., pitfall trap, ZRC_ENT00048441;
Central Catchment Nature Reserve, Venus
Loop, 1.35549, 103.81578, 15 Jun 2015,
M.K.L. Wong leg., ZRC_ENT00000724;
Lower Peirce Reservoir, 25 Aug 1990, collector
unknown, ZRC_ENT00000117; same locality as previous, Aug 1990, collector unknown,
ZRC_ENT00000155; same locality as previous, 18 Aug 1990, collector unknown, ZRC_
ENT00000170; queens and workers, same
locality as previous, 7 Aug 1990, collector unknown, ZRC_ENT00000197; Mandai, 27 Jun
2019, J. Tan & N. Chin leg., NN-L01, ZRC_
ENT00013537; Mandai Road, Nov-Dec 2016,
G.W. Yong leg., ZRC_HYM_0001424-1459;
Nee Soon Swamp Forest, 1.38252, 103.80208,
25 Sep 2017, W. Wang leg.,Winkler extraction,
NS_W1, ZRC_ENT00027908; Rifle Range
Road, 3 Jun 1993, collector unknown, NS110,
ZRC_HYM_0000190; male, Seletar Reservoir,
16 May 1994, collector unknown, NS 134B,
ZRC_ENT00000267; Upper Thomson Nature
Park, 1.38311, 103.79839, Sep-Oct 2016, G.W.
Yong leg., ZRC_HYM_0001397-1423.

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Material not physically examined. Syntype workers, A.R. Wallace leg., SINC.55/9 (BMNH),
including CASENT0900666, ANTC19768(BMNH) [as type of Ponera transversa].
Non-types – CASENT0249126, PSW102561(PSWC); Harbourfront, Marang Trail, 4 Aug
2014, J.K. Wetterer leg., vial #473 (JKWC).
Literature. Type – Smith (1857) [as Ponera transversa]. Viehmeyer(1916); Overbeck (1924);
Yamane (2009); Tan & Corlett (2012).
Localities. Bukit Batok East; Bukit Timah Nature
Reserve; Bukit Timah Road; Central Catchment Nature Reserve; Harbourfront; Lower
Peirce Reservoir; Mandai; Mandai Road; NeeSoon Swamp Forest; Rifle Range Road; Seletar Reservoir; Upper Thomson Nature Park.
Habitat/Ecology. This species was found mostly
in primary and old/mature secondary forests in
Singapore, including swamp forest, frequently
in or on leaf litter. Sometimes, individuals
have been collected from well-developed
young secondary forest such as abandoned
plantation patches. In one instance, foragers
were collected from inside an Anoplolepis
gracilipes nest in a fallen rotting tree branch.
Remarks. Type locality in Singapore.
Parvaponera darwinii (Forel, 1893)
Material examined. Alate queens and males,
Raffles Lighthouse, 10 Aug 2019, J. Tan leg.,
ZRC_ENT00027936.
Material not physically examined. Alate queen,
CASENT0249179, PSW09596-2 (PSWC).
Literature. Viehmeyer (1916), Overbeck (1924)[both as Euponera (Trachymesopus) darwini
var. indica].
Localities. Bukit Timah Road; Kent Ridge.
Habitat/Ecology. Alate queens and males of thisspecies were collected from around lamps inurban parklands and a garden. Alates werealso collected on an offshore dive boat in surrounding waters of a lighthouse islet, where
they swarmed around lights on the vessel atnight. Actual habitat preferences of this species are unknown in the Singapore context at
the time of writing.

Platythyrea parallela (Smith, 1859)
Material examined. Bukit Batok, 1 Sep 2016,
G.W. Yong leg., ZRC_ENT00047913; Bukit
Batok East, 28 Aug 2016, G.W. Yong leg.,
ZRC_ENT00048025; Bukit Timah Nature Reserve, 13 Dec 1967, D.H. Murphy leg., ZRC_
ENT00027924; queens, males and workers,
Bukit Timah forest, Jalan Jambul, 4 Oct 1973,
D.H. Murphy leg., A340-3, DHM-SG73-Plat1,
ZRC_ENT00027923; Kranji Road, 29 Jan
2016, G.W. Yong leg., ZRC_ENT00048026;
Lower Peirce, 7 May 2017, G.W. Yong leg.,
ZRC_ENT00048419; Mandai forest, 15 Jan
2020, W. Wang leg., ZRC_ENT00027938;
Mandai Road, 1.41258, 103.79839, 16 Nov
2016, G.W. Yong leg., ZRC_HYM_0001552;
male, Nee Soon Swamp Forest, 1°23’00.3”N,
103°48’46.5”E, 28 Mar-4 Apr 2012, J. Puniamoorthy et al. leg., malaise trap, NS1, Reg.
29157, ZRC_BDP0012762; alate queen,Prince George’s Park Residences (NUS),
1.29239, 103.77869, 15-22 Apr 2015, M.S.
Foo & W. Wang leg., malaise trap, NUS0013,
ZRC_BDP0045470; Pulau Ubin, 1.41662,
103.99443, 4 Dec 2016, G.W. Yong & M.K.L.
Wong leg., ZRC_HYM_0001080-1082; Seletar Link, 1.40581, 103.88629, 23 Oct 2016,
G.W. Yong et al. leg., ZRC_ENT00047914;
alate queen and workers, same locality as
previous, 23-25 Oct 2016, G.W. Yong leg.,
ZRC_HYM_0001546-1548; alate queens
and worker, Upper Thomson Nature Park,
1.38311, 103.79839, Oct 2016, G.W. Yongleg., ZRC_HYM_0001549-1551.
Material not physically examined. ANIC32065989, RWT68.11 (ANIC); ANIC32-06599165992, AreaK7, A64-6/A66-4 (ANIC);
ANIC32-065994, RWTAcc.68.35(ANIC);
queens [types of Platythyrea coxalis], FOCOL0847-849 (ZMHB); FOCOL0850851[types of P. pusilla var. egena] (ZMHB).
Non-types, Katong, Amber Road, 28 Jul 2014,
J.K. Wetterer leg., vial #337 (JKWC).
Literature. Viehmeyer (1916), Overbeck (1924)[as Platythyrea coxalis and P. pusillia var.
egena].

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Localities. Amber Road (Katong); Bukit Batok;
Bukit Batok East; Bukit Timah Nature Reserve; Bukit Timah Road; Kranji Road; Mandai; Mandai Road; Nee Soon Swamp Forest;
Prince George’s Park Residences (NUS);
Pulau Ubin; Seletar Link; Singapore Botanic
Gardens; Upper Thomson Nature Park.
Habitat/Ecology. This species is associated mainly with both mature and young secondary forests, including native-dominated, abandoned
plantation, and waste woodland forests, often
fragments in urban or semi-urban settings.
Sometimes the ants have been found on low
vegetation and foliage in urban parklands. The
species is considered arboreal and individuals
are often found on tree trunks or branches,or foliage in general. Nests have been foundunder bark of living trees or large fallen logs.
Remarks. Platythyrea parallela is known to exhibit broad (body) size variation among both
sympatric and allopatric populations. Other
variable traits include shape and size of propodeal angles, shape of the petiolar node (specifically that of the posterodorsal margin), andbody sculpture. At the point of writing, all size
and morphological variants observed from local material have thus been treated as conspecific, pending future more refined analyses todistinguish between different species of thispotential species complex.
Platythyrea sagei Forel, 1900
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916); Overbeck (1924).
Localities. Bukit Timah Road
Habitat/Ecology. Workers of this species were
found in a half-decayed tree trunk on the
ground in a garden.
Remarks. At the time of writing, this species isonly known from Singapore based on dated
literature records.

Ponera menglana Xu, 2001
Material examined. Bukit Timah forest, 6
Jun 1968, D.H. Murphy leg., D7-4, ZRC_
HYM_0000644; same locality and collection
data as previous, D7-4, ZRC_HYM_0000649;
Bukit Batok abandoned park, 28 Jan 2015,
W. Wang leg., ZRC_ENT00027964; queen
and workers, Mandai Lake Road, Northern Node, 1°24’36.5”N, 103°47’04.0”, 53
m, 18 Jul 2019, J.S. Tan & N.L. Chin leg.,
Winkler extraction, JTNC_MIS_013, ZRC_
ENT00028291.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Batok; Bukit Timah Nature Reserve; Mandai Lake Road.
Habitat/Ecology. This species is associated mostly with different grades of secondary forest(i.e., young, mature, old) at varying levels of
disturbance, including waste woodland forestof an abandoned park. The ants were typicallycollected from rotten wood, leaf litter, andupper soil substrate.
Ponera swezeyi (Wheeler, 1933)
Material examined. Prince George’s Park Residences (NUS), 1.29239, 103.7787, 6 Dec
2017, W. Wang leg., ZRC_ENT00027912;
alate queen, Kent Ridge Park, YM site 2, 7 Dec
2017, W. Wang leg., ZRC_ENT00027984;
Bukit Timah Nature Reserve, 1.35599,
103.77397, 8 Aug 2017, W. Wang leg., WWSG17-058, ZRC_ENT00028318; same locality as previous, 8 Sep 2005, collector unknown, ZRC_ENT00048385.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Kent
Ridge Park; Prince George’s Park Residences(NUS).
Habitat/Ecology. This species is associated mostly with old/mature secondary forests at variedlevels of disturbance. Individuals were collected from leaf litter and soil substrate via
Winkler extraction, and also soil at bases oftrees. On one occasion, some workers were
found with a Brachyponera colony in rottingwood.

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Remarks. Despite being fairly widespreadthroughout Australasia and Oceania, P.
swezeyi is suspected to actually be native
to the Oriental region because of its sharedcharacters with three other minute Oriental
species: P. baka Xu, 2001, P. xantha Xu, 2001,
and P. shennong Terayama, 2009 (Leong et al.
2019). These similarities include: relatively
small body size (HW 0.28-0.35), thick petiolar
node in lateral view, yellowish-brown body,and 5-segmented antennal club (sometimes
indistinct).
Pseudoneoponera havilandi (Forel, 1901)
Material examined. None available.
Material not physically examined. Syntype –CASENT0907253, ANTC27462 (MHNG).
Literature. Type – Forel (1901) [as Pachycondyla(Bothroponera) havilandi].
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore context.
Remarks. Type locality in Singapore. At the timeof writing, the species is known only fromSingapore, by its type series. While there has
been no recent material for objective comparisons, based on type images of P. havilandi
on AntWeb and its original description, thevalidity of the species seems well-justified.
The more striking characters that distinguishP. havilandi from other congeners known to
the region would be: 1) small eyes, each with
only 9-10 ommatidia along its maximum diameter, 2) petiolar node in dorsal view 1.31.5x wider than long, truncate with convex
sides and slightly concave posterior margin.
Pseudoneoponera insularis (Emery, 1889)
Material examined. Alate queen, Nee Soon
Swamp Forest, NS1, 1°23’00.3”N,
103°48’46.5”E, 11-18 Apr 2012, J. Puniamoorthy et al. leg., malaise trap, Reg. 29181,
ZRC_BDP0012698.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[as Pachycondyla (Bothroponera) insularis].
Localities. Bukit Timah Road.

Habitat/Ecology. Individuals were found in atermite ‘hill’ between the roots of a tree ina garden. An alate queen was collected fromswamp forest, caught in a malaise trap sample.
Pseudoneoponera rufipes (Jerdon, 1851)
Material examined. Bukit Timah Nature Reserve,
Jun-Aug 1967, D.H. Murphy leg., pitfall trap,
ZRC_ENT00027992; same locality as previous, plot III-2, 28 Apr 2016, D.J. Court leg.,
pitfall trap, ZRC_ENT00048451; same locality as previous, collection date unknown,
D.H. Murphy leg., ZRC_HYM_0000873;
same locality as previous, Aug 1967, D.H.
Murphy leg., ZRC_HYM_0000869; Lower
Peirce Reservoir, 7 Aug 1990, collector unknown, ZRC_HYM_0000242; Sunset Way,1.32609, 103.77187, 21-23 Sep 2016, G.W.
Yong leg., pitfall trap, ZRC_HYM_0001184;
same locality, collection date and collector as
previous, ZRC_HYM_0001633.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Lower
Peirce Reservoir; Sunset Way.
Habitat/Ecology. This species is associated mostly
with primary or old/mature secondary forests,
sometimes also in young secondary forests
such as abandoned plantation forests, in Singapore. These ants are usually ground-foraging
and were often collected using pitfall traps.
Pseudoneoponera tridentata (Smith, 1858)
Material examined. Bukit Timah Nature Reserve,
Jun-Aug 1967, D.H. Murphy leg., pitfall trap,
ZRC_ENT00027993; same locality and collector as previous, 14 May 1967, pitfall trap, A670/C2-2, ZRC_ENT00048392; Lower Peirce
Reservoir, 7 Aug 1990, collector unknown,
ZRC_HYM_0000237-238; Mandai, 1.4083,
103.77803, 12 Nov 2015, M.K.L. Wong leg.,
ZRC_ENT00000728; Mandai Road, 1.41258,
103.79839, 12 Jul 2016, G.W. Yong leg.,
M1D7GH2506, ZRC_HYM_0001634; alatemale, Nee Soon Swamp Forest, NS1, malaise
trap, Reg. 29181, ZRC_BDP0012698; University Campus, 1 Apr 1972, D.H. Murphyleg., ZRC_HYM_0000909; Upper Thom-

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

son Nature Park, 1.38311, 103.79839, 1113 Sep 2016, G.W. Yong leg., UT2GP226,
ZRC_HYM_0001635; Upper Thomson Road,Oct 2016, G.W. Yong leg., GY-SG16-Pseu1,
ZRC_ENT00028000.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[both as Pachycondyla (Bothroponera) tridentata].
Localities. Bukit Timah Nature Reserve; Lower
Peirce Reservoir; Mandai; Mandai Road; NeeSoon Swamp Forest; University Campus (notspecified if Bukit Timah or Clementi/Kent
Ridge); Upper Thomson Nature Park; Upper
Thomson Road.
Habitat/Ecology. This species is associated mostly with primary or old/mature secondary forests, sometimes also in well-developed young
secondary forest fragments in Singapore. A
single alate was collected from swamp forest in a malaise trap sample. These are verylarge ground-foraging ants that form relatively
small colonies.
PROCERATIINAE (3 genera, 7 species)
Discothyrea bryanti (Wheeler, 1917)
Material examined. Seletar Trail, 1.39151,
103.80056, 16 Apr 2016, M.K.L. Wong leg.,
Winkler extraction, ZRC_ENT00000675.
Material not physically examined. ANIC32030767, D4-60PP.RA-38 (ANIC).
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Seletar.
Habitat/Ecology. This species is associated withprimary or old/mature secondary forests in
Singapore, typically collected from leaf litterand topsoil debris.
Discothyrea sauteri Forel, 1912
Material examined. Bukit Timah Nature Reserve, 20 Jun 1967, D.H. Murphy leg., D46, ZRC_HYM_0000253; same locality as
previous, BT09, 1.35127, 103.78161, 5 Oct
2016, W. Wang leg., Winkler extraction,
ZRC_HYM_0000563.
Material not physically examined. Unknown.
Literature. None. New record.

Localities. Bukit Timah Nature Reserve
Habitat/Ecology. This species is associated withprimary or old/mature secondary forests in
Singapore. Individuals were typically foundin leaf litter and soil substrate from maturesecondary forest.
Remarks. At the time of writing, this species isknown from only one locality in Singapore.
Probolomyrmex vieti Eguchi, Yoshimura & Yamane, 2006
Material examined. Alate queen, Pulau Semakau,
SMO3, 1.20664, 103.76044, 22-29 Nov 2012,
J. Puniamoorthy et al. leg., malaise trap, Reg.
29604, ZRC_BDP0015248.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Pulau Semakau
Habitat/Ecology. A single alate queen of this species was found in a malaise trap sample fromold growth mangroves on an offshore landfill.
Remarks. At the time of writing, this species isknown from only one locality in Singapore,
based on a single alate. The presence of a reproductive individual may suggest that thisspecies is not a mere transient visitor to thearea.
Probolomyrmex watanabei Tanaka, 1974
Material examined. Bukit Timah Nature Reserve, 4 Jul 1969, D.H. Murphy leg., D12-6,
ZRC_HYM_0000017; Central Catchment
Nature Reserve, 24 Aug 2011, E.J.Y. Soh leg.,
CASENT0317191, ZRC_ENT00048018;
queen, Mandai Lake Road, Northern Node,1°24’36.5”N, 103°47’04.0”, 53 m, 18 July
2019, J.S. Tan & N. Chin leg., Winkler extraction, ZRC_ENT00028309; queen and workers, Seletar Trail, 1.39151, 103.80056, 16 Apr
2016, M.K.L. Wong leg., Winkler extraction,
ZRC_ENT00000676.
Material not physically examined.
ANIC32-030944, D9-1/A124-7 (ANIC);
CASENT0172416, D8-1 (ANIC).
Literature. None. New record.
Localities. Bukit Timah Nature Reserve; Central
Catchment Nature Reserve; Mandai Lake
Road; Seletar.

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Habitat/Ecology. This species was found in primary and/or old/mature secondary forests in
Singapore. Individuals were mostly collectedfrom leaf litter.
Remarks. Some specimens examined were observed to have petiole in profile elongate withnearly flat dorsal margin, closely resemblantof Probolomyrmex itoi or P. longiscapus.
However, the same specimens in dorsal vieweach had posterodorsal margin of petiole medially produced into a horn, a feature uniqueto and diagnostic of P. watanabei.
Proceratium deelemani Perrault, 1981
Material examined. Queens and workers, Bukit
Timah Nature Reserve, 1.35127, 103.78161,
8 May 2017, W. Wang leg., WW-SG17-014,
ZRC_HYM_0000610; Seletar Trail, 1.39151,
103.80056, 16 Apr 2016, M.K.L. Wong leg.,
Winkler extraction, ZRC_ENT00000677.
Material not physically examined. ANIC32025871, RWTacc.68.34 (ANIC); FMNHINS0002821932-2821935, FMHD_81-282(FMNH).
Literature. Baroni Urbani & de Andrade (2003).
Localities. Bukit Timah Nature Reserve; Seletar.
Habitat/Ecology. This species was found inprimary or old/mature secondary forests in
Singapore. Nests have been found in soil substrate and debris in fallen wood (at the base ofa large liana). Specimens have also been collected in litter from forest floors and crevicesof tree buttresses.
Proceratium malesianum de Andrade, 2003
Material examined. Mandai Lake Road, NorthernNode, 1°24’36.5”N, 103°47’04.0”, 53 m, 18
July 2019, J.S. Tan & N. Chin leg., Winkler
extraction, ZRC_ENT00028314.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Mandai Lake Road
Habitat/Ecology. This species was found in a mature secondary forest fragment in Singapore.
Individuals were collected from a mixture ofleaf litter and topsoil substrate.
Remarks. At the time of writing, this species isknown from only one locality in Singapore.

Proceratium papuanum Emery, 1897
Material examined. Queen, Pulau Ubin, 1.41662,
103.99443, 4 Dec 2016, G.W. Yong & M.K.L.
Wong leg., ZRC_HYM_0001092; queen,
Sungei Buloh Wetland Reserve, 28 Mar 2018,
W. Wang leg., ZRC_ENT00000961.
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Pulau Ubin; Sungei Buloh Wetland
Reserve.
Habitat/Ecology. This species was identified
based on queens collected from disturbed old/
mature secondary forest, and also mangroves.
Actual nesting sites are unknown.
Remarks. The queens from Singapore identified
as P. papuanum confer mostly with the original description of the species, but differ fromthe holotype queen in terms of body coloration– the Singapore queens appear darker in coloration. At the time of writing, we undertakethe assumption that this might be a case of
intraspecific variation among allopatric populations.
PSEUDOMYRMECINAE (1 genus, 11 species)
Tetraponera allaborans (Walker, 1859)
Material examined. Kent Ridge, 3 Jan 1983, D.H.
Murphy leg., ZRC_HYM_0000765; same
locality and collector as previous, 13 May
1985, ZRC_HYM_0000766; Mandai mangroves, 2 Oct 1978, D.H. Murphy leg., ZRC_
HYM_0000764; Pulau Semakau New Fragment, SMN3, 1°12’05.6”N, 103°45’46.5”E,
13-20 Sep 2012, J. Puniamoorthy et al. leg.,malaise trap, Reg. 29471, ZRC_BDP0016113;
queen and workers, University Hall (NUS),
1.29711, 103.77658, May-Aug 2015, M.S.
Foo & W. Wang leg., malaise trap, ZRC_BDP(multiple); Upper Thomson Road, 13 Sep
2016, G.W. Yong leg., ZRC_ENT00048105;
Sungei Buloh, 8 Jan 2013, Sk. Yamane leg.
(SKYC).
Material not physically examined. Alate
queen, CASENT0752596, ANTC42318(CASC); CASENT0795901, tc1162110327(PSWC); queens, males, and workers,
CASENT0795901-795912 (PSWC).

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Literature. Viehmeyer (1916), Overbeck (1924)[both as Sima (Tetraponera allaborans, andS. allaborans var. sumatrensis); Ward (2001);
Wang et al. (2018a).
Localities. Bukit Timah; Kent Ridge; Lim Chu
Kang; Mandai mangroves; Pulau Semakau;
Sungei Buloh; University Hall (NUS); Upper
Thomson Road.
Habitat/Ecology. This arboreal species was foundin a broad range of habitats at varying levelsof disturbance in Singapore, such as mangroves, young secondary forest fragments andscrubland in urban or semi-urban settings. In
mangroves, individuals were found on low
vegetation, in dead twigs of coastal plants
such as Hibiscus tiliaceus; nests have beenfound in Sonneratia branches. The ants werealso collected from secondary forest fringes,
roadsides and urban parklands, in fallen deadtwigs of various tree or shrub species including: Clerodendron disparifolium, Mallotussp., Citharexylum spinosum, Delonix regia,
and Vitex pubescens. Sometimes individuals
were found on foliage in secondary forest,
including on Bromhedia orchids.
Remarks. Specimens examined were observed
to be morphologically rather variable acrosssamples, but this is to be expected of the species (see Ward 2001) and all morphologicalvariants were treated as conspecific at the timeof writing.
Tetraponera attenuata Smith, 1877
Material examined. Queen, Neo Tiew Lane, 16
Jul 2009, H.K. Lua leg., LHK 513, ZRC_
HYM_0000236; Rifle Range Road, 27 Jul
2016, G.W. Yong leg., ZRC_ENT00048104;
Upper Thomson Nature Park, 1.38009,
103.8258, 4 Oct 2016, G.W. Yong et al. leg.,
uT7GH1860, ZRC_ENT00048099.
Material not physically examined.
CASENT0752611, ANTC42333 (CASC).
Literature. Forel (1907) [as Sima attenuata var.
thagatensis]; Viehmeyer (1916), Overbeck(1924) [both as Sima (Tetraponera) attenuata]; Ward (2001).
Localities. Bukit Timah Road; Neo Tiew Lane;
Rifle Range Road; Seletar Reservoir; Upper
Thomson Nature Park.

Habitat/Ecology. This arboreal species is associated mainly old/mature secondary forests in
Singapore, including abandoned plantation orjust disturbed forest fragments in semi-urban
settings. Some specimens were collected fromabove ground foliage using sweepnets.
Tetraponera crassiuscula (Emery, 1900)
Material examined. None available.
Material not physically examined. Unknown.
Literature. Viehmeyer (1916), Overbeck (1924)[both as Sima (Tetraponera) allaboranscrassiuscula].
Localities. Bukit Timah Road
Habitat/Ecology. Individuals were found in agarden.
Remarks. At the time of writing, this species isknown from Singapore only based on literature records.
Tetraponera difficilis (Emery, 1900)
Material examined. Mandai Road, 1.41488,
103.79917, 6-8 Dec 2016, G.W. Yong et al.
leg., ZRC_ENT00048097; Upper Thomson
Nature Park, 1.38012, 103.82616, 4 Oct 2016,
G.W. Yong et al. leg., ZRC_ENT00048098;
Pulau Ubin, 7 Jan 2014, Sk. Yamane leg.
(SKYC).
Material not physically examined.
CASENT0907466, ANTC28118 (MHNG)[syntype of Sima stipitum]; queen andworkers, FOCOL1176-1178 (ZMHB)[types of Sima fulva]; CASENT0220759,
PSW09602 (UCDC); males and workers,
CASENT0796174-796183 (PSWC).
Literature. Forel (1912) [as Sima stipitum]; Viehmeyer (1916), Overbeck (1924) [both as Sima
(Tetraponera) fulva]; Ward (2001).
Localities. Bukit Timah Road; Changi Beach;
Mandai Road; National University of Singapore; Peirce Reservoir; Pulau Ubin; Seletar
Reservoir; Singapore Botanic Gardens; Upper
Thomson Nature Park.
Habitat/Ecology. This species is associated mainly with primary and old/mature secondary forests, sometimes young secondary habitats, ofvarying levels of disturbance in Singapore.
These include native-dominated and aban-

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doned plantation secondary forests. The ants
have been found around forest edges, in loose
dead twigs of plants such as Mallotus sp. The
species also occurs sporadically in urban parklands or gardens, and coastal beach forest.
Colonies in Singapore have been observed
to be polydomous, with aggressive, activelystinging workers.
Tetraponera extenuata Ward, 2001
Material examined. Paratype - Bukit Timah, 100m,
20 Nov 1988, P.S. Ward leg., ZRC_6_20705.
Non-types - Mandai Bike Trail, 1.4083,
103.77803, 19 Jun 2016, M.K.L. Wong leg.,
ZRC_ENT00000678; Nee Soon Swamp Forest, NS2, 1°23’04.2”N, 103°48’40.7”E, 2-9
May 2012, J. Puniamoorthy et al. leg., malaise
trap, Reg. 29221, ZRC_BDP0014635; Pulau
Ubin, PU1, 1°24’36.3”N, 103°59’25.5”E, 1219 Oct 2013, M.S. Foo et al. leg., malaise trap,
Reg. 30212, ZRC_BDP0016184.
Material not physically examined. Holotype – CASENT0902824, PSW09570(BMNH). Paratypes – CASENT0761615,
PSW09570 (PSWC); queen and workers,
CASENT0796004-796007,
PSW09570 (PSWC). Non-types –CASENT0796008-796011 (PSWC);
CASENT0217591, ANTC17842 (CASC).
Literature. Type — Ward (2001).
Localities. Bukit Timah [type]; Kent Ridge; Mandai; Nee Soon Swamp Forest; Peirce Reservoir; Pulau Ubin; Seletar Reservoir.
Habitat/Ecology. This arboreal species is associated mostly with primary or old/maturesecondary forests in Singapore, including
native-dominated secondary rainforest andfreshwater swamp forest. Both colonies andindividuals, including the type series, havebeen found in dead twigs of Timonius sp. andRhodamnia trinervia. Individuals have occasionally been collected in malaise traps set
up in mangroves and swamp forest.
Remarks. Type locality in Singapore.

Tetraponera inversinodis Ward, 2001
Material examined. Locality unknown [but deduced as Nee Soon based on ‘swamp forest’on original label], 14 Apr 1993, Sk. Yamaneleg. (SKYC).
Material not physically examined. Unknown.
Literature. None. New record.
Localities. Unknown.
Habitat/Ecology. A single specimen of this species was collected from a swamp forest.
Remarks. Closely related to T. difficilis, T. inversinodis can be differentiated by its larger
eyes, shorter and broader profemur, and shorter metatibia (Ward 2001).
Tetraponera modesta (Smith, 1860)
Material examined. Bukit Timah Nature Reserve,
BT06, 1.35284, 103.77995, 23 Nov 2016, W.
Wang leg., ZRC_HYM_0000261; same locality as previous, 25 Mar 1968, D.H. Murphyleg., ZRC_HYM_0000768-769; Seletar Trail,
1.39532, 103.80228, 5 Mar 2016, M.K.L.
Wong leg., ZRC_ENT00000679.
Material not physically examined.
CASENT0907461,
ANTC28113(MHNG) [syntype of Sima fulva];
CASENT0796039-796040, PSW0857501/
PSW10255-01 (PSWC).
Literature. Viehmeyer (1916), Overbeck (1924)[both as Sima (Tetraponera) fulva]; Baroni
Urbani (1977) [as Tetraponera fulva]; Ward
(2001).
Localities. Bukit Timah Nature Reserve; Bukit
Timah Road; Seletar.
Habitat/Ecology. This arboreal species is associated with primary or old/mature secondary
forests in Singapore. Colonies were found inhollows and cavities of fallen branches andtwigs. Individuals were sometimes collectedfrom foliage by beating.

Remarkable diversity in a little red dot: a comprehensive checklist of known ant species in Singapore(Hymenoptera: Formicidae) with notes on ecology and taxonomy

Tetraponera nitida (Smith, 1860)

Tetraponera pilosa (Smith, 1858)

Material examined. HortPark, 1.27954,
103.79759, 25 Feb 2016, Boyi Zhou leg.,
ZRC_ENT00000680; Kent Ridge,12 Jun 1970,
D.H. Murphy leg., ZRC_HYM_0000810;
Kranji Road, 27-29 Jan 2016, G.W. Yongleg., ZRC_ENT00048101-48102; Pulau Semakau Old Fragment, SMO3, 1°12’23.9”N,
103°45’37.6”E, 8-15 Nov 2012, J. Puniamoorthy et al. leg., malaise trap, Reg. 29578,
ZRC_BDP0016208; Pulau Ubin, 7 Jan 2014,
Sk. Yamane leg., ZRC_HYM_0000473.
Material not physically examined.
CASENT0220763,
PSW10251(UCDC); queens, males and workers,
CASENT0796308-796314 (PSWC); queen
and workers, FOCOL1172-1175 (ZMHB)[types of Sima siggi var. setifera]; males,
FOCOL1182-1184 (ZMHB) [types of Simadifficilis r. longiceps].
Literature. Viehmeyer (1916), Overbeck (1924)[as Sima (Tetraponera) difficilis longiceps andS. siggi var. setifera]; Ward (2001).
Localities. Bukit Timah; Bukit Timah Road; HortPark; Kent Ridge; Kranji Road; Lim Chu
Kang; Mandai mangrove; Pulau Semakau;
Pulau Ubin; Seletar Reservoir.
Habitat/Ecology. This species is associated mostly with disturbed secondary habitats especially rainforest edges/fringes and mangroves
or mangrove forests in Singapore. On a few
occasions, individuals were collected fromsecondary forest fragments in semi-urban settings. At rainforest edges, individuals havebeen found in dead twigs of Mallotus sp. andon Bromheadia (orchids). In mangroves ormangrove forest, the species was found on
low vegetation, and in dead twigs of Avicennia
alba. The species has also been found nesting in hollow branches of Chiku trees, andthe hollowed-out pith-channel of a bough ina mangosteen tree.

Material examined. Bukit Timah Nature Reserve, plot II-I, 15 Aug 2011, J. Koh leg.,
ZRC_ENT00000889; Icube (NUS), 1.29347,
103.77633, 6-13 May 2015, M.S. Foo & W.
Wang leg., malaise trap, ZRC_BDP0044952;
queen, Kent Ridge, 10 Nov 1991, collector unknown, ZRC_HYM_0001750; alate
queen, Lim Chu Kang Mangrove, 19 May
1987, Serena Teo leg., ZRC_ENT00048805;
Mandai, 7 Apr 2016, M.K.L. Wong leg.,
ZRC_ENT00000681-682; queen, Pulau Ubin,
PU1, 1°24’36.3”N, 103°59’25.5”E, 1-8 Sep
2012, J. Puniamoorthy et al. leg., malaise
trap, Reg. 29449, ZRC_BDP0016492; same
locality and collectors as previous, 25 Aug-1
Sep 2012, malaise trap, Reg. 29436, ZRC_
BDP0015989; Rifle Range Road, 27 Jul 2016,
G.W. Yong leg., ZRC_ENT00048103; Toh
Tuck wasteland, 5 Mar 1978, D.H. Murphyleg., ZRC_HYM_0000809.
Material not physically examined.
CASENT0220765, PSW09567 (UCDC);
males and workers, CASENT0796714-796718(PSWC); queens, FOCOL1163-1165 (ZMHB)[types of Pseudomyrma pilosa].
Literature. Viehmeyer (1916), Overbeck (1924)[as Sima pilosa]; Ward (2001); Wang et al.
(2018a).
Localities. Bukit Timah Nature Reserve; Bukit
Timah Road; Kent Ridge; Lim Chu Kang
Mangrove; Mandai; Peirce Reservoir; Pulau
Ubin; Rifle Range Road; Toh Tuck.
Habitat/Ecology. This species is associated mostly with mature secondary forest and the edges/
fringes of such rainforest. It has sometimes
also been found in scrub, wasteland, or disturbed secondary forest fragments in urban orsemi-urban settings. Alates have been collected from mangroves either using malaise traps,
or in mud lobster (Thalassina sp.) mounds.
Individuals in secondary forest or forest edges
were often found in dead twigs, such as that of
Hevea brasiliensis (rubber), and on low vegetation. Colonies in semi-urban settings suchas gardens have also been found in dead twigs
of mangosteen trees, and in withered bamboo.

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Tetraponera rufonigra (Jerdon, 1851)

Tetraponera vivax Ward, 2001

Material examined. Choa Chu Kang Avenue 2, 1 Mar 2017, D.J. Court leg., ZRC_
HYM_0001725; Jurong Lake Gardens, 29 Nov
2018, H.H. Tan leg., ZRC_ENT00007632;
Jurong mangroves, 7 Nov 1976, D.H. Murphy leg., ZRC_HYM_0000912; KranjiRoad, 1.42638, 103.75413, Sep-Nov 2016,
G.W. Yong leg., ZRC_HYM_00016391640; Seletar, 20 Dec 1982, D.H. Murphyleg., ZRC_HYM_00000802-805; University
Campus, Jan 1977, D.H. Murphy leg., ZRC_
HYM_0000806.
Material not physically examined. Queens, males,
and workers, CASENT0796770-796774,
tc1162117908/PSW09580 (PSWC).
Literature. Viehmeyer (1916), Overbeck (1924)[both as Sima rufonigra]; Ward (2001).
Localities. Bukit Timah Nature Reserve; Choa
Chu Kang; Jurong Lake Gardens; Jurong
mangroves; Kranji Road; Mandai; Seletar;
University Campus (either Bukit Timah or
Clementi/Kent Ridge).
Habitat/Ecology. This species was found in a
broad range of habitats in Singapore, including mangroves and degraded or heavily modified environments such as urban parklands,
gardens and semi-urban waste woodland secondary forests. Individuals were frequently
collected from flowering shrubs on granite
and tree trunks. The species typically nests
in cavities of both dead and living wood. In
mangroves, a colony was found occupying
live beetle-bored branches of Sonneratia ovata, the nest cavities expanded into the boleof the tree. On one occasion, the ants were
found predating on Pomacea (apple snail) egg
masses on wooden stakes supporting planted
trees near the edge of a man-made water body,in urban parkland. Workers of the species are
known to be behaviourally aggressive andretaliatory when provoked.

Material examined. None available.
Material not physically examined. Holotype
alate queen, collection date and locality unspecified, C.F. Baker leg. (MCZ).
Literature. Type – Ward (2001).
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore context.
Remarks. Type locality in Singapore. At the timeof writing, this species is known from Singapore only by the holotype queen. The species
was first described based on three queens collected from Singapore, Sabah and Java respectively.
Tetraponera volucris Ward, 2001
Material examined. None available.
Material not physically examined. Holotype
alate queen, collection date and locality unspecified, 9209, C.F. Baker leg. (MCZ).
Literature. Type – Ward (2001).
Localities. Unknown.
Habitat/Ecology. Unknown in the Singapore context.
Remarks. Type locality in Singapore. At the timeof writing, the only record of this species in
Singapore is that of the holotype queen. The
queen of T. volucris appears similar to that of
T. vivax, but can be distinguished by its tridentate clypeal margin and widely separated
frontal carinae, also other traits such as larger
eyes, absence of malar pit, and more elongate
postpetiole.
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ACKNOWLEDGEMENTS
We thank all friends, family and colleagues who
contributed, in one way or another, towards thecompletion of this milestone publication. We areespecially grateful to the following experts whohelped with identification, and/or provided precious material or specimen data (in no particularorder): the late Prof. ‘Paddy’ D.H. Murphy, Katsuyuki Eguchi, Shingo Hosoishi, Koichi Arimoto, Fuminori Ito, Masashi Yoshimura, Weeyawat
Jaitrong, Herbert Zettel, John (Jack) Longino,
James (Jim) Wetterer, Lam Weng Ngai. We would
also like to thank Teo Siyang for the use of thevector base map of Singapore. Finally, we thankthe reviewers for their invaluable advice and comments on increasing the impact of this manuscript.

ASIAN MYRMECOLOGY
A Journal of the International Network for the Study of Asian Ants
Communicating Editor: Adam Cronin