ASIAN MYRMECOLOGY Volume 9, e009014, 2017

ISSN 1985-1944 | eISSN: 2462-2362 © Chi-Man Leong, Shiuh-Feng Shiao
and Benoit Guénard

DOI: 10.20362/am.009014

Ants in the city, a preliminary checklist of Formicidae(Hymenoptera) in Macau, one of the most heavily urbanizedregions of the world
Chi-Man Leong1, Shiuh-Feng Shiao1 and Benoit Guénard2*
National Taiwan University, Department of Entomology,
No.1, Sec.4, Roosevelt Road, Taipei, Taiwan
2
University of Hong Kong, School of Biological Sciences,
Kadoorie Biological Sciences Building, Pok Fu Lam Road,
Hong Kong SAR, China
1

Corresponding author: bguenard@hku.hk*

ABSTRACT. Macau is a small territory in South East China and one of themost densely populated regions in the world. Previous studies on insect groups
have shown that a relatively diverse, yet specific, fauna could still survive in thisregion. However, to this point, studies on the myrmecofauna of Macau are scarceand to date no species checklist exists. Here, we present the first checklist of
Macanese ant species by combining results from recent ant surveys using handcollections and Winkler extractors with published records. During the surveys,
82 species and morphospecies belonging to 37 genera and 8 subfamilies havebeen collected, with 37 species representing new records for Macau, including
an interesting new record of an undescribed Leptanilla species, the second
record of the Leptanillinae subfamily for South East China. To date, Macanese
ants comprise 105 species/morphospecies and 8 subspecies, after the removalof dubious records present in the literature (though some misidentifications
may remain). While still likely incomplete, these results represent the mostcomprehensive list of ants for Macau, and a baseline for future research on ant
diversity in heavily urbanized environments and for understanding the potential
consequences of urbanization on native and non-native diversity in Asia.
Keywords: Ants, Checklist, Exotic Species, Macao, Diversity Survey,
Urban Ecosystems.
INTRODUCTION
Globally, urban development has been constantly
expanding; in one analysis, by 2050, 70% of the
global human population is expected to live within cities leading to an increase of urbanized areas(Seto et al. 2013). In this context, understanding
what role urban habitats can play to maintain acertain level of biodiversity and which species
are present in these environments is key to future
ecology and conservation.

Located in the subtropical part of China,
Macau Special Administrative Region (SAR)(22.2˚N, 113.5˚E) lies on the south west side of
the Pearl River mouth, and is a small territory of
30.4 km2, encircled within the Chinese provinceof Guangdong. Similar to several cities in South
China, Macau has experienced rapid urban development over the past decades and as of 2017its population density is the highest in the world
with 20,166 inhabitants per km2: 2.5 and 3 times
higher than Singapore and Hong Kong respec-

2

Chi-Man Leong, Shiuh-Feng Shiao & Benoit Guénard

tively (Information Services Department Hong
Kong Special Administrative Region Government 2015; United Nations 2015; Statistics andCensus Service 2017). Historically, Macau consisted of a peninsula connected to Guangdong,and of two islands: Coloane Island (8.07 km2) andTaipa Island (7.6 km2). Over the past decades, important land reclamation over the ocean has led tothe merging of Coloane and Taipa Islands together. Despite this important urbanization and landchanges, a few “natural” areas and parks havebeen preserved. For example, Ka Ho and Hac
Sa Reservoirs on Coloane Island are freshwaterwetland ecological zones, and areas of these tworeservoirs are regulated for the conservation purpose of an endemic moss Fissidens macaoensis(Zhang & Hong 2011) and an endemic mosquito
Toxorhynchites macaensis (Ribeiro 1997).
Regarding the Macanese insect fauna,several groups like butterflies and moths (Lepidoptera) (Easton & Pun 1997a, b), water beetles(Coleoptera) (Jäch & Easton 1998), stink bugs
(Homoptera) (Easton & Pun 1999), mosquitoes,
sandflies, and blackflies (Diptera) (Ou 2001) oragain damselflies (Zygoptera) (Wilson & Xu
2007) have been previously surveyed in Macau,
but other major taxonomic groups that compose the Macanese biodiversity are still poorly
known, including ants (Formicidae). Yet, understanding ant diversity and species composition isimportant. For instance, within an urban matrix
composed of a diversity of habitats, even small
natural patches of natural habitats can still support high species diversity of ants (Menke et al.
2011; Guénard et al. 2015), while on the other
hand non-native or habitat disturbance specialist
species can represent a significant portion of themyrmecofauna within the habitats (Guénard etal. 2015). Moreover, as ants represent a dominantgroup among arthropods and show sensitivity toenvironmental changes in relation to disturbance,
they can represent a good bioindicator across different habitats and help in monitoring their respective ecological values (Andersen et al. 2002;
Del Toro et al. 2012).
To the best of our knowledge, no specific checklist or sampling has been realized tocharacterize ant diversity in Macau, and only
sporadic records have been reported in the literature. To date, the most comprehensive list of antsof Macau was provided in the 1920’s in a seriesof articles by the American myrmecologist William Morton Wheeler, and embedded with otherrecords for various parts of China (Wheeler 1921,
1928, 1930). A few other records from Macau
were reported or cited in various taxonomic revisions on Chinese or Asian ants (Xu 2003; Eguchi2008). Over the past two decades, several recordsof Macau ants were included within Chinese
species checklists, yet with no or little specificsampling efforts for this territory (e.g. Wu 1941;
Chapman & Capco 1951; Tang et al. 1995; Zhou
2001; Fellowes 2006; Hua 2006; Guénard &Dunn 2012; Ran & Zhou 2011, 2012, 2013); andexisting records of Macanese ants were not included in a checklist of ants in Guangdong (Zhaoet al. 2009). Despite these limited efforts and thesmall area that Macau represents, two species andsubspecies of ants have been described on the basis of specimens collected there, namely Carebara capreola laeviceps (Wheeler, 1928) andCrematogaster macaoensis Wu & Wang 1995,
with this latter recorded from other Chinese provinces as well (Guénard & Dunn 2012; antmaps.
org 2017). The subspecies C. capreola laeviceps
is endemic to Macau, however as no recent work
has examined the taxonomic validity of this species, the possibility that this species represents
a synonym should not be excluded. In addition,
most historical records were reported without
specific collecting locations in Macau and without habitat information, which limits the understanding of the distribution of these species. As
a result, knowledge on Macanese ants is at thispoint limited, and in order to establish a landmark
allowing improved future work on ant diversity
and conservation on this territory, a first species
checklist is currently needed.
Here, we present the results of a preliminary ant survey conducted in the main natural habitats of Macau. We then establish the firstcomprehensive species checklist of Macanese
ants based on newly collected materials andpublished records. This checklist may benefit
knowledge of the ants from Macau and SouthEast China, and provide detailed information for
future studies. In addition, the discussion on ant
community of each study site is also presented
to provide insights on the effects of urbanization
on ants.

Ants in the city, a preliminary checklist of Formicidae (Hymenoptera) in Macau

3

Fig. 1. Map of Macau SAR presenting the sampling locations (modified after Google Map 2017). Pie charts represent the ant composition in native (white), tramp (grey) and exotic species (black) for the sites with the mostcomplete sampling effort.

MATERIALS AND METHODS
The main sampling sites were located on Guia
Hill on the Macau Peninsula, Tai Tam Hill and
Siu Tam Hill on Taipa Island and Ka Ho and Hac
Sa Reservoirs on Coloane Island (sampling locations shown on Figure 1). Field work was conducted from July to August 2015, July to August
2016, January 2017 and July to August 2017. It
should be noted that sampling effort among thedifferent sites was not standardized and more
effort was applied on sites located on Coloane
Island, to deliberately target more natural andmature habitats preserved on this island rather
than the urbanized and isolated habitats present
on Taipa Island and Macau Peninsula. In 2015, a
total of 23 field sessions were realized; in each ofthem we surveyed for about five hours using handcollection; in 2016 and 2017, a total of fourteen
field sessions were realized, again for about five
hours using hand collection but coupled with theuse of a Winkler extractor, this latter being an efficient method to collect cryptobiotic ants present in the leaf litter as well as ground dwelling
ants (Bestelmeyer et al. 2000). In addition, somespecimens collected in diverse locations of Macau were generously provided by Mr Martinho G.
Oxalá and Mr Chi-Yui Lei.
For hand collection, we searched forants visually with a specific focus on the ground,
under rocks, in rotten wood (breaking it open),and on tree trunks with the help of an insect aspirator. For Winkler extractor sampling, six litres ofsifted leaf litter (abundantly decomposed organic

4Chi-Man Leong, Shiuh-Feng Shiao & Benoit Guénard

materials) were collected within each site andchosen to maximize the number of micro-habitats
sampled and thus ant diversity (Ward 2000). Leaf
litter was dried in a Winkler extractor for sevendays. All collected specimens were preserved in
75% ethanol before being point mounted and later
processed for identification. All specimens of thisstudy were deposited in the Insect Biodiversity
and Biogeography Laboratory at the Universityof Hong Kong.
Taxonomic changes and updates
Regarding previous records, taxonomic updates
and changes were incorporated into our results
and followed taxonomic information available in
AntCat.org (2017). Voucher specimens from previous records in the literature were not depositedin Macau or did not have specific voucher specimens (Wu 1941; Tang et al. 1995; Zhou 2001;
Xu 2003; Hua 2006), which prevented us fromexamining these records. Thus, to identify potential dubious records, we used information on species’ known distributions available on antmaps.
org (Janicki et al. 2016; Guénard et al. 2017) andin the literature. Here we retained the subspecies
level to reflect the lack of recent taxonomic revisions in several genera, with some of the current
subspecies representing potential valid species in
the future. On the other hand, if these subspecies
are later on synonymized with other known species, this would allow to easily adopt future taxonomic changes.
RESULTSA total of 82 valid species and morphospecies,
belonging to 37 genera and from 8 subfamilies,
were collected during this survey, including 37
newly recorded species for Macau, which now

Table 1. Collection sites of this study with details on location and collection methods used (including GPS coordinate,
attitude, size of habitat, sampling times and methods such as H. = Hand collection, W. = Winkler extractor). The
last column (right) present the ant diversity retrieved at each site, as well as the number of exotic and tramp species.

Code

LocationGPS
coordinate

Attitude
(m)

Size
(ha)

Sampling
sessions

Methods

Total species(exotic, tramp)A

Ilha Verde Hill

22.2116 °N,
113.5374°E

55

4.3

1

H. + W.

3 (1,0)B

Guia Hill

22.1983 °N,
113.5511 °E

64

16.4

10

H. + W.

31 (2,5)C

Penha Hill

22.1833 °N,
113.5319 °E

37

5.3

1

H.

3 (1,1)D

Siu Tam Hill

22.1608 °N,
113.5466 °E

75

26

4

H. + W.

29 (4,0)E

Tai Tam Hill

22.1575 °N,
113.5650 °E

104

77.9

4

H.

16 (3,3)F

Ecological22.1406 °N,
Protection Area
113.5544 °E
I & II

0

55

1

H.

11 (3,3)G

Ka Ho
Reservoir

22.1244 °N,
113.5672 °E

93

177.6

6

H. + W.

32 (3,1)H

Hac SaReservoir

22.1344 °N,
113.5725 °E

65

264.7

8

H. + W.

50 (8,5)I

Tam Kong
Temple22.1138 °N,
113.5500 °E

11

4

2

H.

5 (2,1)

5

Ants in the city, a preliminary checklist of Formicidae (Hymenoptera) in Macau

Table 2. Species recorded during previous studies (extracted from Guénard & Dunn 2012; antmaps.org 2017)
and the current study. Single asterisk in front of species name shows new records from Macau. Also with notes(e.g. tramp and exotic species). The collecting methods (H. = Hand collection, W. = Winkler extractor) for eachspecies records are provided.
Species

Earliest record This study

Notes

Methods

AMBLYOPONINAE[1 genus, 1 species]
Stigmatomma rothneyi (Forel, 1900)

Wheeler 1928√

H. + W.

√H.

DOLICHODERINAE[5 genera, 9 species; 2 morphospecies]
Chronoxenus wroughtonii (Forel, 1895)
Chronoxenus wroughtonii formosensis (Forel, 1913)

Tang et al.1995
Hua 2006

Chronoxenus dalyi (Forel, 1895)

Wheeler 1928

Chronoxenus walshi (Forel, 1895)

Hua 2006

Dolichoderus taprobanae (Smith, 1858)

Wheeler 1928

Dolichoderus sp. mo01 [cf. sibiricus Emery, 1889]
Ochetellus glaber (Mayr, 1862)

Wheeler 1928
Wheeler 1921√

Tang et al.1995

H.
H. + W.

Tramp√

Tapinoma sp. mo01 [nr. melanocephalum (Fabricius, 1793)]
Technomyrmex brunneus Forel, 1895

Tramp

√

*Tapinoma indicum Forel, 1895
Tapinoma melanocephalum (Fabricius, 1793)√

H.
H. + W.

√

Tramp

H. + W.

√

Tramp

H.

DORYLINAE[1 genus, 1 species]*Ooceraea biroi (Forel, 1907)
FORMICINAE[8 genera, 23 species; 7 morphospecies]*Acropyga acutiventris Roger 1962
Acropyga sauteri Forel, 1912
Anoplolepis gracilipes (Smith, 1857)

Wheeler 1928

Camponotus albosparsus Bingham, 1903

Wheeler 1928

*Camponotus lighti Wheeler, 1927
Wheeler 1928

*Camponotus nicobarensis Mayr, 1865
Camponotus parius Emery, 1889

Wheeler 1921

*Camponotus vitiosus Smith, 1874
Camponotus variegatus (Smith, 1858)
Camponotus variegatus dulcis Dalla Torre, 1893
Camponotus variegatus proles Emery, 1925

H.

√

H. + W.

Wheeler 1928

Acropyga sp. mo02

Camponotus mitis (Smith, 1858)√√

Wu 1941

H. + W.

√

H.

√

H. + W.

√

H. + W.

√

H. + W.

√

H. + W.

Hua 2006
Wheeler 1928

ExoticTramp√

H. + W.

6

Chi-Man Leong, Shiuh-Feng Shiao & Benoit Guénard

Species

Earliest record This study

Notes

Methods

Camponotus sp. mo01 [cf. mitis (Smith, 1858)]√

H.

Colobopsis sp. mo01 [nr. nipponica Wheeler, 1928]√

H.

Colobopsis sp. mo02 [nr. vitrea Smith, 1860]√

H.

Lepisiota rothneyi watsonii (Forel, 1894)

Wheeler 1921

Nylanderia amia (Forel, 1913)

Wheeler 1928

Nylanderia bourbonica (Forel, 1886)
Nylanderia indica (Forel, 1894)
Nylanderia vividula (Nylander, 1846)√

Hua 2006

Tramp

H. + W.

Tramp

Wheeler 1928
Hua 2006√

H.

Nylanderia sp. mo01√

H. + W.

Nylanderia sp. mo02 [nr. vividula (Nylander, 1846)]√

H. + W.

Nylanderia sp. mo03 [cf. birmana (Forel, 1902)]√

H.

Nylanderia yerburyi (Forel, 1894)

Wheeler 1928

ExoticParaparatrechina sauteri (Forel, 1913)

Hua 2006

Paratrechina longicornis (Latreille, 1802)

Wheeler 1928

*Polyrhachis demangei Santschi, 1910
Polyrhachis dives Smith, 1857

Tang et al. 1995

*Polyrhachis illaudata Walker, 1859√

ExoticH. + W.

√

H.

√

H. + W.

√

H. + W.

√W.

LEPTANILLINAE[1 genus; 1 morphospecies]
Leptanilla sp. mo01
MYRMICINAE[12 genera, 40 species; 9 morphospecies]√*Cardiocondyla minutior Forel, 1899
Carebara capreola (Wheeler, 1927)

Tramp

H.

Xu 2003

Carebara capreola laeviceps (Wheeler, 1928)

Wheeler 1928

Carebara diversa (Jerdon, 1851)

Wheeler 1921

*Carebara diversa laotina (Santschi, 1920)√

H.

*Carebara zengchengensis (Zhou, Zhao & Jia, 2006)√

H. + W.

Carebara sp. 2 BG [nr. melasolena (Zhou & Zheng, 1997)]√

H. + W.

Carebara sp. mo02√

H. + W.

√

H.

*Crematogaster quadriruga Forel, 1911√

H. + W.

*Crematogaster rogenhoferi Mayr, 1879√

H. + W.

Meranoplus sp. mo01 [sp. nr. bicolor (GuérinMéneville, 1844)]√

H. + W.

Crematogaster biroi Mayr, 1897

Wheeler 1928

Crematogaster dohrni artifex Mayr, 1879

Wheeler 1928

*Crematogaster ferrarii Emery, 1888
Crematogaster macaoensis Wu & Wang, 1995

Wheeler 1928

7

Ants in the city, a preliminary checklist of Formicidae (Hymenoptera) in Macau

Species

Earliest record This study

Notes

Methods*Monomorium floricola (Jerdon, 1851)

√

Tramp

H.

*Monomorium pharaonis (Linnaeus, 1758)√

ExoticW.

Monomorium sp. mo01√

*Myrmecina sinensis Wheeler, 1921√

W.

*Pheidole tumida Eguchi, 2008√

H. + W.

Pheidole fervens Smith, 1858

Eguchi 2008√

Pheidole hongkongensis Wheeler, 1928

Eguchi 2008√

Pheidole indica Mayr, 1879

Wheeler 1928

Pheidole megacephala (Fabricius, 1793)

Eguchi 2008

H. + W.

Tramp

H. + W.
H. + W.

Tramp√

ExoticH. + W.

*Pheidole nodus Smith, 1874√

H. + W.

*Pheidole ochracea Eguchi, 2008√

H. + W.

√

H. + W.

Pheidole taipoana Wheeler, 1928

Eguchi 2008

Pheidole parva Mayr, 1865

Eguchi 2008

Pheidole sp. mo01√

H.

*Recurvidris recurvispinosa (Forel, 1890)√

H. + W.

Solenopsis geminata Buren, 1972

Wheeler 1928√

ExoticH.

Solenopsis invicta (Fabricius, 1804)

Hua 2006√

ExoticH. + W.

*Solenopsis jacoti Wheeler, 1923√

*Strumigenys emmae (Emery, 1890)√

Exotic√*Strumigenys exilirhina Bolton, 2000
Strumigenys membranifera Emery, 1869

H. + W.

Wheeler 1928

H. + W.
H. + W.

Exotic*Strumigenys minutula Terayama & Kubota, 1989√

H. + W.

*Strumigenys nepalensis Baroni Urbani & De
Andrade, 1994√

H. + W.

Strumigenys silvestrii Emery, 1906

Chapman andCapco 1951

ExoticSyllophopsis sp. mo01 [nr. sechellensis (Emery, 1894)]√*Tetramorium bicarinatum (Nylander, 1846)

√

Tramp

H. + W.

√

Tramp

H. + W.

√

Tramp

H. + W.

*Tetramorium kraepelini Forel, 1905
Tetramorium lanuginosum Mayr, 1870

Wheeler 1928

H. + W.

*Tetramorium nipponense Wheeler, 1928√

H.

*Tetramorium parvispinum (Emery 1893)√

H. + W.

*Tetramorium simillimum (Smith, 1851)√

Tetramorium sp. 1BG (obesum group Bolton, 1976)√

H. + W.

Tetramorium sp. mo02√

H. + W.

Tetramorium sp. mo03 [cf. wroughtonii (Forel, 1902)]√

H. + W.

ExoticH.

8

Chi-Man Leong, Shiuh-Feng Shiao & Benoit Guénard

Species

Earliest record This study

Notes

Methods

PONERINAE[11 genera, 17 species; 2 morphospecies]√*Anochetus risii Forel, 1900
Bothroponera rubiginosa (Emery, 1889)

Wheeler 1928

Brachyponera luteipes (Mayr, 1862)

Wheeler 1928

Brachyponera obscurans (Mayr, 1862)

Wheeler 1928

Diacamma rugosum (Le Guillou, 1842)

Hua 2006

H.
Tramp

√

H. + W.

√

H. + W.

√

H.

√

H. + W.

√

H. + W.

*Hypoponera exoecata (Wheeler, 1928)√

H. + W.

Hypoponera sp. mo01√

H. + W.

*Leptogenys chinensis (Mayr, 1870)

√

H.

√

H. + W.

√

H. + W.

√

H. + W.

√

H. + W.

Diacamma rugosum anceps Matsumura & Uchida, 1926 Wheeler 1928
Diacamma sp. mo01
Ectomomyrmex astutus (Smith, 1858)

Zhou 2001*Ectomomyrmex leeuwenhoeki (Forel, 1886)*Euponera pilosior Wheeler, 1928
Euponera sharpi Forel, 1901

Wheeler 1928

Harpegnathos venator (Smith, 1858)

Zhou 2001

Harpegnathos venator rugosus (Mayr, 1862)

Wheeler 1928

Leptogenys peuqueti (André, 1887)

Wheeler 1928

*Odontoponera denticulata (Smith, 1858)
Pseudoneoponera rufipes (Jerdon, 1851)

Wheeler 1930

PSEUDOMYRMECINAE[1 genus; 1 species]*Tetraponera allaborans (Walker, 1859)

includes a total of 105 species and 8 subspecies(Table 2). Among these, 11 species can be considered exotic or potentially exotic to Macau, and 15
species are known as tramp species.
The highest diversity in Macau was
found on two sites of Coloane Island, Hac Sa
and Ka Ho Reservoirs, in which 50 and 32 species were collected respectively, inclusive of asignificant number of exotic and tramp species(Table 1 & Figure 1). The Guia Hill on the Macau Peninsula and Siu Tam Hill on Taipa Islandpresented similar levels of species richness aswell as the presence of several exotic and tramp
species. While the other sites sampled had much
lower species diversity, exotic or tramp species

were commonly collected (Table 1 & Figure 1).
It should be noted that the sampling effort acrosssites was not standardized, and thus comparison
between sites should carefully consider this point.
Four species previously reported in Macau, Formica exsecta, Technomyrmex albipes,
Tetramorium guineense and Tetramorium tsushimae, are provisionally excluded from the Macanese ant fauna until further confirmation as theserecords represent distribution anomalies in Macau relative to the rest of their global distribution(antmaps.org 2017); they have not been collectedin this study nor could they be confirmed fromvoucher specimens. Notes on the four dubious
records: (1) Formica exsecta is distributed in re-Ants in the city, a preliminary checklist of Formicidae (Hymenoptera) in Macaugions of temperate climate. Therefore, F. exsecta
should not occur in Macau, and we consider therecord doubtful. (2) Although Technomyrmex albipes is a successful tramp species and spread easily, we have not collected this species. Moreover
this species has often been misidentified in thepast and is part of a group known to be difficult
to identify (Bolton 2007), and previous recordsof T. albipes in East Asia most likely represent
records of T. brunneus which was commonly collected in this study. (3) Tetramorium guineense isrestricted to the Afrotropical region (Hita Garcia
et al. 2010). The record from Macau, presented
by Wheeler (1928), does not contain enough information on the specimen diagnosis or description to confirm its presence and no new specimen
was collected in this study. Moreover, previous
records of T. guineense from Macau most likely
represent records of T. bicarinatum which wascommonly collected in this study, and has been
incorrectly reported as T. guineense in many locations around the world (Wetterer 2009). (4)Records of T. tsushimae are based on previous
records of T. caespitum jacoti and T. caespitum

9

jacoti annectens by Wheeler (1928), which are
now considered primary synonyms of T. tsushimae. Species of the genus Tetramorium are particularly difficult to identify in Asia and these
species are regularly misidentified. However, T.
tsushimae is primarily a temperate-distributed
species and its presence in the subtropical climateof Macau can be considered doubtful.
Figure 2 presents the accumulationof Macanese ant species records through time,
which currently includes a total of 105 species
and 8 subspecies based on literature records andnewly collected material (Figure 2 and Appendix 1). The sharp increase following this survey
is noticeable and demonstrates the incomplete
knowledge on Macanese ants for both nativeand exotic species. The majority of the species
newly recorded in the Macanese myrmecofauna
are considered native, but it should be noted that
newly recorded exotic species have also beenreported recently. Two notorious invasive species, Solenopsis invicta, native to South America (first recorded by Hua 2006), and Pheidole
megacephala, native to Africa (first recorded by

Fig. 2. Accumulation of Macanese ant species knowledge based on literature records and this study. Pie chartspresent the ant composition in native (white), tramp (grey) and exotic species (black) for each time period (details
presented in appendix 1).

10

Chi-Man Leong, Shiuh-Feng Shiao & Benoit Guénard

Eguchi 2008), represent recent introductions andseem now widespread in Macau based on theirmultiple occurrences in the recent survey. Here,
Strumigenys emmae, native to the Australasianregion, is for the first time recorded in Macau.
We also report the first record of Ooceraea biroi.
While this species could be native to Macau, its
exact native range is uncertain and this species
is known to have successfully colonized various
regions around the world: Taiwanese and Japanese islands, Pacific islands, Caribbean islands,
Malagasy islands and Christmas Island (Australia). Both species possess cryptobiotic habits andlive within the leaf litter and soil. The records ofMonomorium pharaonis and Tetramorium simillimum, both native from the Afrotropical region,
represent new records for Macau. Other notorious exotic species first recorded nearly 90 years
ago (e.g. Anoplolepis gracilipes, Paratrechina
longicornis and Solenopsis geminata) were stillcommonly encountered in Macau.
The comprehensive species checklist of
Macanese ants is shown in Table 2, in addition tocollecting methods used for each species (wheninformation is available).
Checklist of species collected in this study* indicates newly recorded species from Macau.
AMBLYOPONINAE [1 genus; 1 species]
Stigmatomma rothneyi (Forel, 1900)
Material examined. Guia Hill, 9.VIII.2015, 1
worker, CM Leong leg.; Guia Hill, 27.VIII.2016,
1 worker and 1 male, CM Leong leg.
DOLICHODERINAE [4 genera; 4 species, 2morphospecies]
Dolichoderus sp. mo01 [cf. sibiricus Emery, 1889]
Material examined. Hac Sa Reservoir,
29.VII.2015, 1 worker, CM Leong leg.; Ecological Protection Area I & II, 12.I.2017, 1 worker,
CM Leong leg.

Ochetellus glaber (Mayr, 1862)
Material examined. Guia Hill, 28.VII.2015, 1
worker, CM Leong leg.; Ecological Protection
Area I & II, 12.I.2017, 3 workers, CM Leong leg.
*Tapinoma indicum Forel, 1895
Material examined. Siu Tam Hill, 5.VIII.2015,
1 worker, CM Leong leg. Hac Sa Reservoir,
20.VIII.2016, 1 worker, CM Leong leg. Ilha
Verde Hill, 25.I.2017, 1 dealate queen, CM Leong leg.
Tapinoma melanocephalum (Fabricius, 1793)
Material examined. Guia Hill, 5.VIII.2015,
1 worker, CM Leong leg.; Ecological Protection Area I & II, 12.I.2017, 2 workers and 1 dealate queen, CM Leong leg.; Hac Sa Reservoir,
2.VIII.2017, 1 worker, B Guénard leg.
Tapinoma sp. mo01 [nr. melanocephalum (Fabricius, 1793)]
Material examined. Ka Ho Reservoir,
14.VII.2016, 1 worker, CM Leong leg.; Guia
Hill, 20.VIII.2016, 1 worker, CM Leong leg.;Hac Sa Reservoir, 20.VIII.2016, 1 worker, CM
Leong leg.; Siu Tam Hill, 26.VIII.2016, 1 worker,
CM Leong leg.; Guia Hill, 27.VIII.2016, 1 worker, CM Leong leg.
Technomyrmex brunneus Forel, 1895
Material examined. Guia Hill, 13.VII.2015, 1
worker, CM Leong leg.; Guia Hill, 15.VII.2015, 1
worker, CM Leong leg.; Guia Hill, 27.VIII.2016,
1 worker, CM Leong leg.
DORYLINAE [1 genus, 1 species]*Ooceraea biroi (Forel, 1907)
Material examined. Tai Tam Hill, 25.VIII.2016,
1 worker, MG Oxalá leg.
FORMICINAE [7 genera; 14 species, 7 morphospecies]*Acropyga acutiventris Roger 1962
Material examined. Guia Hill, 24.VII.2015,
1 dealate queen, CM Leong leg.; Siu Tam Hill,
26.VIII.2016, 1 dealate queen, CM Leong leg.;Ants in the city, a preliminary checklist of Formicidae (Hymenoptera) in Macau

Hac Sa Reservoir, 29.VII.2017, 1 dealate queen
and 7 workers, CM Leong leg. Hac Sa Reservoir,
2.VIII.2017, 4 dealate queens and 7 workers, B
Guénard leg.
Acropyga sp. mo02
Material examined. Hac Sa Reservoir,
29.VII.2017, 3 workers, CM Leong leg.
Anoplolepis gracilipes (Smith, 1857)
Material examined. Hac Sa Reservoir,
7.VIII.2015, 1 worker, CM Leong leg.; Ka Ho
Reservoir, 14.VII.2016, 1 worker, CM Leong leg.;Hac Sa Reservoir, 20.VIII.2016, CM Leong leg.
*Camponotus lighti Wheeler, 1927
Material examined. Tai Tam Hill, 29.VII.2015,
1 major, CM Leong leg.; Tam Kong Temple,
30.VII.2015, 1 minor, CM Leong leg.; Tam Kong
Temple, 31.VII.2015, 1 minor, CM Leong leg.;
Penha Hill, 6.VIII.2015, 1 minor, CM Leong leg.;
Guia Hill, 9.VIII.2015, 1 minor, CM Leong leg.
Camponotus mitis (Smith, 1858)
Material examined. Siu Tam Hill, 18.VII.2015,
1 minor and 1 major, CM Leong leg.; Hac Sa
Reservoir, 31.VII.2015, 1 minor, CM Leong leg.;
Siu Tam Hill, 31.VII.2015, 1 minor, CM Leongleg.; Hac Sa Reservoir, 7.VIII.2015, 2 majors,
CM Leong leg.; Hac Sa Reservoir, 20.VIII.2016,
1 dealate queen, CM Leong leg.; Siu Tam Hill,
26.VIII.2016, 2 minors and 1 major, CM Leongleg.; Hac Sa Reservoir, 9.I.2017, 1 minor and 2
majors, CM Leong leg.
*Camponotus nicobarensis Mayr, 1865
Material examined. Hac Sa Reservoir,
7.VIII.2015, 1 minor and 1 major, CM Leongleg.; Tai Tam Hill, 30.IX.2015, 1 minor, CY Lei
leg.; Ka Ho Reservoir, 14.VII.2016, 1 major, CM
Leong leg.; Guia Hill, 20.VIII.2016, 1 major, CM
Leong leg.; Hac Sa Reservoir, 9.I.2017, 1 minor,
CM Leong leg.
Camponotus parius Emery, 1889
Material examined. Ka Ho Reservoir,
14.VII.2016, 1 minor, CM Leong leg.; Hac Sa
Reservoir, 2.VIII.2017, 2 media, B Guénard leg.

11

Camponotus variegatus dulcis Dalla Torre, 1893
Material examined. Guia Hill, 20.VIII.2016, 1
minor, CM Leong leg.
*Camponotus vitiosus Smith, 1874
Material examined. Siu Tam Hill, 13.VII.2016,
3 minors, MG Oxalá leg. Guia Hill, 20.VIII.2016,
1 minor, CM Leong leg.
Camponotus sp. mo01 [cf. mitis (Smith, 1858)]
Material examined. Siu Tam Hill, 12.X.2015, 1
minor, CY Lei leg.
Colobopsis sp. mo01 [nr. nipponica (Wheeler,
1928)]
Material examined. Tai Tam Hill, 5.VIII.2015, 1
minor, CM Leong leg.
Colobopsis sp. mo02 [nr. vitrea Smith, 1860]
Material examined. Siu Tam Hill, 5.VIII.2015, 1
minor, CM Leong leg.
Nylanderia amia (Forel, 1913)
Material examined. Ka Ho Reservoir,
14.VII.2016, 1 worker, CM Leong leg.; Guia Hill,
20.VIII.2016, 1 worker, CM Leong leg; Hac Sa
Reservoir, 2.VIII.2017, 1 worker, B Guénard leg.
Nylanderia yerburyi (Forel, 1894)
Material examined. Guia Hill, 15.VII.2015, 1
worker, CM Leong leg.
Nylanderia sp. mo01
Material examined. Guia Hill, 15.VII.2015,
1 worker, CM Leong leg.; Hac Sa Reservoir,
3.X.2015, 1 worker, CY Lei leg.; Ka Ho Reservoir, 13.X.2016, 1 worker, CY Lei leg.
Nylanderia sp. mo02 [nr. vividula (Nylander,1846)]
Material examined. Hac Sa Reservoir,
20.VIII.2016, 1 worker, CM Leong leg.; Siu Tam
Hill, 26.VIII.2016, 1 worker, CM Leong leg.
Nylanderia sp. mo03 [cf. birmana (Forel, 1902)]
Material examined. Guia Hill, 15.VII.2015, 1
worker, CM Leong leg.

12

Chi-Man Leong, Shiuh-Feng Shiao & Benoit Guénard

Paratrechina longicornis (Latreille, 1802)
Material examined. Hac Sa Reservoir,
29.VII.2015, 1 worker, CM Leong leg.; Tam
Kong Temple, 30.VII.2015, 1 worker, CM Leong leg.; Penha Hill, 6.VIII.2015, 2 workers,
CM Leong leg.; Ka Ho Reservoir, 14.VII.2016,
2 workers, CM Leong leg.; Hac Sa Reservoir,
20.VIII.2016, 1 worker, CM Leong leg.; Siu Tam
Hill, 26.VIII.2016, 1 worker, CM Leong leg.;Hac Sa Reservoir, 9.I.2017, 1 worker, CM Leongleg.; Ecological Protection Area I & II, 12.I.2017,
1 worker, CM Leong leg.
*Polyrhachis demangei Santschi, 1910
Material examined. Ka Ho Reservoir,
23.VIII.2015, 1 worker, CM Leong leg.
Polyrhachis dives Smith, 1857
Material examined. Hac Sa Reservoir, 22.I.2017,
1 worker, CM Leong leg.
*Polyrhachis illaudata Walker, 1859
Material examined. Ka Ho Reservoir,
11.VII.2015, 1 worker, CM Leong leg.; Ka Ho
Reservoir, 14.VII.2016, 1 worker, CM Leongleg.; Hac Sa Reservoir, 20.VIII.2016, 1 worker,
CM Leong leg; Tai Tam Hill, 02.VIII.2017, 1
worker, B Guénard leg.
LEPTANILLINAE [1 genus; 1 morphospecies]
Leptanilla sp. mo01
Material examined. Ilha Verde Hill, 25.I.2017, 5
workers, CM Leong leg.
MYRMICINAE [12 genera; 30 species, 9 morphospecies]*Cardiocondyla minutior Forel, 1899
Material examined. Tai Tam Hill, 2.VIII.2017, 2
workers, B Guénard leg.
*Carebara diversa laotina (Santschi, 1920)
Material examined. Tai Tam Hill, 02.VIII.2017,
50 workers, B Guénard leg.
*Carebara zengchengensis (Zhou, Zhao & Jia,
2006)

Material examined. Ka Ho Reservoir,
11.VII.2015, 1 minor, CM Leong leg.; Siu Tam
Hill, 18.VII.2015, 1 minor and 1 major, CM
Leong leg.; Siu Tam Hill, 31.VII.2015, 1 minor
and 1 major, CM Leong leg.; Ka Ho Reservoir,
31.VII.2015, 1 minor, 1 major and 1 supermajor,
CM Leong leg.; Tai Tam Hill, 25.IX.2015, 1 minor and 2 majors, CY Lei leg.; Ka Ho Reservoir,
14.VIII.2016, 1 minor and 1 major, CM Leongleg.; Hac Sa Reservoir, 20.VIII.2016, 2 minors,
2 majors and 1 supermajor, CM Leong leg.; Ka
Ho Reservoir, 23.VIII.2016, 1 minor, CM Leongleg.; Siu Tam Hill, 26.VIII.2016, 1 minor, 1 major
and 1 supermajor, CM Leong leg.
Carebara sp. 2 BG [nr. melasolena (Zhou &Zheng, 1997)]
Material examined. Guia Hill, 18.VIII.2016, 3
minors, CM Leong leg.; Guia Hill, 27.VIII.2016,
1 major, CM Leong leg.
Carebara sp. mo02
Material examined. Guia Hill, 15.VII.2015, 1
minor, CM Leong leg.; Siu Tam Hill, 18.VII.2015,
1 minor, CM Leong leg.
*Crematogaster ferrarii Emery, 1888
Material examined. Hac Sa Reservoir,
29.VII.2015, 1 worker, CM Leong leg.; Ka Ho
Reservoir, 23.VIII.2015, 1 worker, CM Leong leg.
*Crematogaster quadriruga Forel, 1911
Material examined. Ka Ho Reservoir,
14.VII.2016, 1 worker, CM Leong leg.
*Crematogaster rogenhoferi Mayr, 1879
Material examined. Tai Tam Hill, 29.VII.2015,
1 worker, CM Leong leg.; Ka Ho Reservoir,
31.VII.2015, 1 worker, CM Leong leg.; Siu Tam
Hill, 5.VIII.2015, 1 worker, CM Leong leg.; Tai
Tam Hill, 5.VIII.2015, 1 worker, CM Leong leg.;
Tai Tam Hill, 20.VIII.2015, 1 worker, CM Leongleg.; Tai Tam Hill, 25.IX.2015, 2 workers, Lei CYleg.; Hac Sa Reservoir, 20.VIII.2016, 1 worker,
CM Leong leg.; Guia Hill, 20.VIII.2016, 1 worker, CM Leong leg.; Siu Tam Hill, 26.VIII.2016, 1
worker, CM Leong leg.; Guia Hill, 27.VIII.2016,
1 worker, CM Leong leg.; Ecological Protection
Area I & II, 12.I.2017, 2 workers, CM Leong leg.

Ants in the city, a preliminary checklist of Formicidae (Hymenoptera) in Macau

Meranoplus sp. mo01 [nr. bicolor (GuérinMéneville, 1844)]
Material examined. Siu Tam Hill, 26.VIII.2016,
1 worker, CM Leong leg.
*Monomorium floricola (Jerdon, 1851)
Material examined. Penha Hill, 6.VIII.2015, 1
worker, CM Leong leg.
*Monomorium pharaonis (Linnaeus, 1758)
Material examined. Hac Sa Reservoir,
29.VII.2017, 1 worker, CM Leong leg.
Monomorium sp. mo01
Material examined. Siu Tam Hill, 26.VIII.2016,
1 worker, CM Leong leg.
*Myrmecina sinensis Wheeler, 1921
Material examined. Hac Sa Reservoir,
29.VII.2017, 1 worker, CM Leong leg.
Pheidole fervens Smith, 1858
Material examined. Hac Sa Reservoir, 9.I.2017,
1 minor, CM Leong leg.
Pheidole hongkongensis Wheeler, 1928
Material examined. Guia Hill, 18.VIII.2016,
1 minor, CM Leong leg.; Siu Tam Hill,
26.VIII.2016, 1 major and 1 minor, CM Leongleg.; Guia Hill, 27.VIII.2016, 1 minor, CM Leongleg.; Hac Sa Reservoir, 29.VII.2017, 1 major and2 minors, CM Leong leg.
Pheidole megacephala (Fabricius, 1793)
Material examined. Guia Hill, 28.VII.2015,
2 minors, CM Leong leg.; Hac Sa Reservoir,
29.VII.2015, 2 minors, CM Leong leg.; Tam
Kong Temple, 30.VII.2015, 1 major, CM Leongleg.; Siu Tam Hill, 26.VIII.2016, 1 major and 1
minor, CM Leong leg.; Ecological Protection
Area I & II, 12.I.2017, 1 major and 1 minor,
CM Leong leg.; Ilha Verde Hill, 25.I.2017, 1 minor and 1 major, CM Leong leg; Tai Tam Hill,
2.VIII.2017, 13 workers, B Guénard leg.
*Pheidole nodus Smith, 1874
Material examined. Guia Hill, 15.VII.2015,
2 minors, CM Leong leg.; Tam Kong Temple,
30.VII.2015, 1 minor, CM Leong leg.; Ka Ho Reservoir, 1.VI.2016, 1 worker, 1 dealate queen and 1

13

male, CY Lei leg.; Hac Sa Reservoir, 20.VIII.2016,
1 minor, CM Leong leg; Hac Sa Reservoir,
2.VIII.2017, 12 workers, B Guénard leg.
Pheidole parva Mayr, 1865
Material examined. Hac Sa Reservoir,
21.VIII.2015, 2 minors, CM Leong leg.; Hac
Sa Reservoir, 2.VIII.2017, 2 minors, B Guénard
leg.; Guia Hill, 20.VIII.2016, 2 majors and 1 minor, CM Leong leg.
*Pheidole ochracea Eguchi, 2008
Material examined. Ka Ho Reservoir,
14.VII.2016, 1 minor, CM Leong leg.; Hac Sa
Reservoir, 2.VIII.2017, 2 minors, B Guénard leg.
*Pheidole tumida Eguchi, 2008
Material examined. Ka Ho Reservoir,
14.VII.2016, 1 minor, CM Leong leg.; Hac Sa
Reservoir, 20.VIII.2016, 1 minor, CM Leongleg.; Hac Sa Reservoir, 29.VII.2017, 2 minors,
CM Leong leg.; Siu Tam Hill, 26.VIII.2016, 1
minor, CM Leong leg.
Pheidole sp. mo01
Material examined. Ecological Protection Area
I & II, 12.I.17, 1 minor, CM Leong leg.
*Recurvidris recurvispinosa (Forel, 1890)
Material examined. Siu Tam Hill, 1 worker, 26.
VIII.2016, CM Leong leg.
Solenopsis geminata (Fabricius, 1804)
Material examined. Hac Sa Reservoir,
7.VIII.2015, 1 alate queen and 2 males, CM
Leong leg.; Hac Sa Reservoir, 21.VIII.2015, 1
worker, CM Leong leg.
Solenopsis invicta Buren, 1972
Material examined. Siu Tam Hill, 5.VIII.2015,
1 worker, CM Leong leg.; Tai Tam Hill,
5.VIII.2015, 1 worker, CM Leong leg.; Hac Sa
Reservoir, 8.VIII.2015, 1 worker, CM Leongleg.; Ka Ho Reservoir, 14.VII.2016, 1 worker,
CM Leong leg.; Ecological Protection Area I &II, 12.I.2017, 1 worker, CM Leong leg.

14

Chi-Man Leong, Shiuh-Feng Shiao & Benoit Guénard

*Solenopsis jacoti Wheeler, 1923
Material examined. Hac Sa Reservoir,
29.VII.2017, 2 workers, CM Leong leg.; Siu Tam
Hill, 26.VIII.2016, 2 workers, CM Leong leg.
*Strumigenys emmae (Emery, 1890)
Material examined. Guia Hill, 18.VIII.2016,
1 worker, CM Leong leg.; Siu Tam Hill,
26.VIII.2016, 1 dealate queen, CM Leong leg; Tai
Tam Hill, 02.VIII.2017, 1 worker, B Guénard leg.
*Strumigenys exilirhina Bolton, 2000
Material examined. Hac Sa Reservoir,
29.VII.2017, 1 worker, CM Leong leg.; Ka Ho
Reservoir, 14.VII.2016, 2 workers, CM Leong leg.
*Strumigenys minutula Terayama & Kubota,
1989
Material examined. Hac Sa Reservoir,
20.VIII.2016, 1 dealate queen, CM Leong leg.;Hac Sa Reservoir, 29.VII.2017, 1 dealate queen,
CM Leong leg.; Ka Ho Reservoir, 20.VIII.2016,
1 worker, MG Oxalá leg.
*Strumigenys nepalensis Baroni Urbani & De
Andrade, 1994
Material examined. Hac Sa Reservoir, 1 worker,
20.VIII.2016, CM Leong leg.
Syllophopsis sp. mo01 [nr. sechellensis (Emery,1894)]
Material examined. Ka Ho Reservoir,
14.VII.2016, 2 workers, CM Leong leg.; Hac Sa
Reservoir, 20.VIII.2016, 1 worker, CM Leongleg.; Ka Ho Reservoir, 23.VIII.2016, 1 worker,
CM Leong leg.; Siu Tam Hill, 26.VIII.2016, 1
worker, CM Leong leg.
*Tetramorium bicarinatum (Nylander, 1846)
Material examined. Guia Hill, 24.VII.2015,
1 worker, CM Leong leg.; Tam Kong Temple,
30.VII.2015, 1 worker, CM Leong leg.; Guia Hill
,5.VIII.2015, 1 worker, CM Leong leg.; Hac Sa
Reservoir, 3.X.2015, 1 worker, CY Lei leg.; Guia
Hill, 18.VIII.2016, 1 worker, CM Leong leg.;Hac Sa Reservoir, 20.VIII.2016, 1 worker, CM
Leong leg.; Ka Ho Reservoir, 20.VIII.2016, 1
worker, MG Oxalá leg.; Guia Hill, 27.VIII.2016,
2 workers, CM Leong leg.; Ecological Protection
Area I & II, 12.I.2017, 1 worker, CM Leong leg.

*Tetramorium kraepelini Forel, 1905
Material examined. Ka Ho Reservoir,
14.VII.2016, 1 worker, CM Leong leg.; Hac Sa
Reservoir, 20.VIII.2016, 3 workers, CM Leongleg.; Ka Ho Reservoir, 23.VIII.2016, 1 worker,
CM Leong leg.
Tetramorium lanuginosum Mayr, 1870
Material examined. Guia Hill, 20.VIII.2016, 1
worker, CM Leong leg.; Guia Hill, 27.VIII.2016,
2 workers, CM Leong leg; Hac Sa Reservoir,
2.VIII.2017, 4 workers, B Guénard leg.; Tai Tam
Hill, 02.VIII.2017, 1 worker, B Guénard leg.
*Tetramorium nipponense Wheeler, 1928
Material examined. Ka Ho Reservoir,
20.VIII.2016, 1 worker, MG Oxalá leg.
Tetramorium parvispinum (Emery, 1893)
Material examined. Ka Ho Reservoir,
14.VIII.2016, 1 worker, CM Leong leg.; Hac Sa
Reservoir, 20.VIII.2016, 2 workers, CM Leong leg.
Note that the specimens are similar to
the type of Tetramorium parvispinum formosae
Forel 1912, now considered a junior synonymof Tetramorium parvispinum, with a distinct andlarge trapezoidal cell present just on the anterodorsal position of eyes.
*Tetramorium simillimum (Smith, 1851)
Material examined. Hac Sa Reservoir,
2.VIII.2017, 4 workers, B Guénard leg.
Tetramorium sp. 1BG (obesum group Bolton,
1976)
Material examined. Guia Hill, 18.VIII.2016,
1 worker, CM Leong leg.; Siu Tam Hill,
26.VIII.2016, 1 worker, CM Leong leg; Hac Sa
Reservoir, 2.VIII.2017, 4 workers, B Guénard leg.
Tetramorium sp. mo02
Material examined. Siu Tam Hill, 26.VIII.2016,
1 dealate queen, CM Leong leg.
Tetramorium sp. mo03 [cf. wroughtonii (Forel,
1902)]
Material examined. Siu Tam Hill, 26.VIII.2016,
1 worker, CM Leong leg.

Ants in the city, a preliminary checklist of Formicidae (Hymenoptera) in Macau

PONERINAE [10 genera; 10 species, 2 morphospecies]*Anochetus risii Forel, 1900
Material examined. Ka Ho Reservoir,
20.VIII.2016, 1 worker, MG Oxalá leg.
Brachyponera obscurans (Mayr, 1862)
Material examined. Guia Hill, 18.VII.2016, 1
worker, CM Leong leg.; Guia Hill, 20.VIII.2016,
1 worker, CM Leong leg.; Hac Sa Reservoir,
20.VIII.2016, 1 worker, CM Leong leg.; Ka Ho
Reservoir, 23.VIII.2016, 1 worker, CM Leongleg.; Siu Tam Hill, 26.VIII.2016, 1 worker, CM
Leong leg.; Guia Hill, 27.VIII.2016, 1 worker,
CM Leong leg.; Ecological Protection Area I &II, 12.I.2017, 2 workers, CM Leong leg.
Diacamma sp. mo01
Material examined. Siu Tam Hill, 31.VII.2015,
1 worker, CM Leong leg.; Ka Ho Reservoir,
14.VII.2016, 2 workers, CM Leong leg.; Guia
Hill, 18.VII.2016, 1 worker, CM Leong leg.; Hac
Sa Reservoir, 9.I.2017, 1 worker, CM Leong leg.
*Ectomomyrmex leeuwenhoeki (Forel, 1886)
Material examined. Tai Tam Hill, 26.VIII.2016,
3 workers, MG Oxalá leg.; Hac Sa Reservoir,
29.VII.2017, 1 worker, CM Leong leg.
*Euponera pilosior Wheeler, 1928
Material examined. Ka Ho Reservoir,
14.VII.2016, 1 worker, CM Leong leg; Hac Sa
Reservoir, 29.VII.2017, 1 alate gyne, CM Leongleg.; Hac Sa Reservoir, 2.VIII.2017, 1 worker, B
Guénard leg.
Harpegnathos venator (Smith, 1858)
Material examined. Hac Sa Reservoir,
7.VIII.2015, 1 worker, CM Leong leg.; Guia Hill,
18.VIII.2016, 1 worker, CM Leong leg.; Hac Sa
Reservoir, 9.I.2017, 1 worker, CM Leong leg.
*Hypoponera exoecata (Wheeler, 1928)
Material examined. Ka Ho Reservoir,
14.VII.2016, 2 workers, CM Leong leg.; Hac Sa
Reservoir, 20.VIII.2016, 1 worker, CM Leongleg.; Siu Tam Hill, 26.VIII.2016, 1 worker, CM
Leong leg.

15

Hypoponera sp. mo01
Material examined. Ka Ho Reservoir,
14.VII.2016, 1 worker, CM Leong leg.
*Leptogenys chinensis (Mayr, 1870)
Material examined. Tai Tam Hill, 25.IX.2015, 1
worker, CY Lei leg.
Leptogenys peuqueti (André, 1887)
Material examined. Guia Hill, 15.VII.2015, 1
worker, CM Leong leg.; Guia Hill, 18.VIII.2016,
1 worker, CM Leong leg.; Guia Hill, 20.VIII.2016,
1 worker, CM Leong leg.; Guia Hill, 27.VIII.2016,
1 worker and 1 male, CM Leong leg.; Hac Sa
Reservoir, 29.VII.2017, 1 worker, CM Leong leg.
*Odontoponera denticulata (Smith, 1858)
Material examined. Guia Hill, 15.VII.2015,
1 worker, CM Leong leg.; Ka Ho Reservoir,
23.VIII.2015, 1 worker, CM Leong leg.; Tai Tam
Hill, 29.VII.2015, 1 worker, CM Leong leg.; Siu
Tam Hill, 25.IX.2015, 1 worker, CY Lei leg.; Siu
Tam Hill, 12.X.2015, 1 worker, CY Lei leg.; Ka
Ho Reservoir, 14.VII.2016, 1 worker, CM Leongleg.; Guia Hill, 18.VIII.2016, 1 worker, CM Leong leg.; Hac Sa Reservoir, 9.I.2017, 1 worker,
CM Leong leg.
Note that although some specimens of
O. denticulata were mentioned by Leong et al.
(2017) for comparing the sculpture variation of
O. denticulata in Asia, the formal record for Macau is firstly provided in this study.
Pseudoneoponera rufipes (Jerdon, 1851)
Material examined. Hac Sa Reservoir,
7.VIII.2015, 1 worker, CM Leong leg.; Ka Ho
Reservoir, 14.VII.2016, 1 worker, CM Leong leg.
PSEUDOMYRMECINAE [1 genus; 1 species]*Tetraponera allaborans (Walker, 1859)
Material examined. Guia Hill, 27.VIII.2016, 1
worker, CM Leong leg.

16

Chi-Man Leong, Shiuh-Feng Shiao & Benoit GuénardDISCUSSION
Here, we present the first ant survey specificto Macau, with results providing new data for
82 species/morphospecies from 37 genera in 8
subfamilies. More than 2/5 (37 species) of thespecies collected represent new species records
from Macau, which brings the known Macanese
myrmecofauna to 105 species and 8 subspecies (Table 2); with this checklist summarizing
literature records collected for nearly a century
and excluding potential dubious records. Whileduring this period the urbanization of Macau has
increased, and it cannot be excluded that someof the species previously collected in Macau are
now locally extinct, the results presented heresuggest the presence of a diverse ant fauna in
Macau. This is particularly interesting as Macau
is a small region (30.4 km2) with the highest human population density globally, and with mostof its natural habitats small and highly fragmented within the urban matrix. For comparison, andwith the limitations due to climatic differences,the myrmecofauna of Macau is probably as diverse as the myrmecofauna found in Oregon(110 species; 254,806 km2) or South Korea (112
species; 100,210 km2), both regions being several thousands of times larger than Macau (datafrom antmaps.org 2017, Guénard et al. 2017).
While the results presented here should be perceived as preliminary, nonetheless, they provide
a basal framework for future research on Macanese ant fauna and can be informative to measurepotential extinction and introduction rates of antsin the region.
Typical of heavily urbanized environments (McKinney 2008; Guénard et al. 2015),
the proportion of species considered either as
exotic or tramp species was high. Here, nearly aquarter of the species collected (19/82; 23.2%)
belong to one of these two categories, with eightspecies considered exotic (Table 2), Anoplolepis gracilipes, Monomorium pharaonis, Paratrechina longicornis, Pheidole megacephala,
Solenopsis geminata, Solenopsis invicta, Strumigenys emmae, Tetramorium simillimum andeleven other species considered tramp species:
Cardiocondyla minutior, Monomorium floricola,
Nylanderia amia, Ochetellus glaber, Ooceraea
biroi, Pheidole fervens, Tapinoma melanoceph-

alum, Technomyrmex brunneus, Tetramorium
bicarinatum, Tetramorium kraepelini, Tetramorium lanuginosum. Exotic species prevalence
is particularly high in Macau as shown by theirpresence in all sites sampled in this study. Moreover, within the best sampled sites tramp or exotic species represented a significant portion of
the myrmecofauna (Fig. 1), consistent with McKinney’s (2008) speculation of a high proportionof non-native and habitat disturbance specialist
species retrieved within heavily urbanized environments. Finally, several of the exotic species
collected are considered among the world’s worst
invasive alien species (Anoplolepis gracilipes,
Pheidole megacephala and Solenopsis invicta:
Lowe et al. 2000) which could result in problems
for local biodiversity conservation (Holway et al.
2002). Our results also indicate that new exotic
species can be detected in Macau. However, at
this point, it is unclear if these species represent
recent species introductions or much older establishment that has been missed until now due tothe limited extent and nature of sampling. Nonetheless, measures to control establishment of new
exotic species through trading activities should
be considered as potential new invaders present
in the Asia/Pacific region (e.g. Linepithema humile, Wasmannia auropunctata) could establish
populations in Macau (antmaps.org 2017).
The array of sampling methods used inthis study is likely insufficient to capture the full
spectrum of the Macanese myrmecofauna, e.g. noarboreal or subterranean traps being used, whichmight in the future allow the collection of new
species. Yet, the use of Winkler extractors within
this study, a collecting method used infrequently
in South East China, has enhanced the collectionof cryptobiotic ant species, including within urban habitats. For instance, in 2015, 37 species/
morphospecies from 23 genera were collectedusing only hand collection over 26 sampling sessions; while in 2016 and 2017, 68 species/morphospecies from 37 genera were collected from a
combination of both hand collections and Winkler
extractors over 14 sampling sessions. Particularly, the use of Winkler extractors has allowed
the collection of four species of Carebara (formerly Oligomyrmex), one Myrmecina species,
four Strumigenys species, one Syllophopsis species, two Hypoponera species and a new record

Ants in the city, a preliminary checklist of Formicidae (Hymenoptera) in Macauof the rarely collected subfamily Leptanillinae,
the second record for South East China with thefirst recently reported from Hong Kong (Wong &Guénard 2016c). This survey represents a starting
point for future myrmecological research in Macau. However future studies are needed in particular to improve taxonomic knowledge to delimit
and identify current morphospecies, with some
representing new species (e.g. Leptanilla species;
Leong et al. submitted); and to complete the survey of Macanese ants through the use of a wider
range of sampling methods. For instance, the useof complementary and efficient methods such asarboreal, subterranean and pitfall traps are likely
to provide new records for Macau and potentially to Guangdong province. Recently, the use of
subterranean pitfall traps and Winkler extractors
in Singapore, another heavily urbanized city ofSouth East Asia, has allowed the recent collectionof three new ant species (Wong & Guénard 2016a,b, c) and the discovery of the species of Leptanillasuggests that similar outcomes might be possible.
Based on current knowledge, the diversity of Macanese ants is comparable to that ofother diverse groups such as 145 moths and 74
butterflies (Lepidoptera; Easton & Pun 1997a,b), and significantly higher than the 37 stinkbugs(Homoptera; Easton & Pun 1999), 35 mosquitoes(Diptera; Ou 2001), 25 water beetles (Coleoptera;
Jäch & Easton 1998) or 7 damselflies (Zygoptera;
Wilson & Xu 2007) recorded thus far. Although
it should be noted that further inventory in someof these groups (e.g. moths, Odonata), as for
ants, is probably necessary, the high taxonomicand ecological diversity of ants observed within
an urban matrix combined with the presence ofnumerous native and non-native disturbance specialists should encourage their use as a bioindicator in Macau and other urbanized regions ofAsia; in particular if coupled with studies withinmore pristine habitats in the region to determine
composition changes following urbanization. Finally, future studies should focus on the distribution and potential ecological impacts of the tramp
and exotic species observed in the ant community surveyed, as most of the ecological impactsof these species on the regional native fauna andflora are still unknown and should be investigatedin the future to benefit our knowledge on urbanized habitats within Asia.

17

In summary, although Macau is smalland highly urbanized, the need for biodiversity
inventory and the protection of the few natural
terrestrial habitats left is important as these might
still host a diverse fauna, including some species of peculiar importance (e.g. Leptanilla species). To enhance our knowledge of the Macanese
myrmecofauna and guide the dilemmas of urban
development, future studies should include comprehensive collecting methods, further study onthese morphospecies, and ecological impacts oftramp and exotic species.
ACKNOWLEDGEMENTS
We would like to express our gratitude to Dr
Seiki Yamane (Kagoshima University Museum,Japan), Dr Chung-Chi Lin (National Changhua
University of Education, Taiwan), Dr Feng-long
Jia (Sun Yat-Sen University, China), Mr Hao
Ran (Guangxi Normal University, China), andMr Yun Hsiao (NTU) for their suggestions andhelp. Thank you Mr Roger Ho Lee, Mrs Ying
Y. Luo, Mr Brian Worthington (HKU), Mr TszYeung Wong and Miss Man-Kwan Wong (NTU)
for their assistance in Hong Kong. We would liketo also thank Dr Zheng-Hui Xu (Southwest Forestry College, China), Dr John Fellowes and Dr
Adam Cronin (Tokyo Metropolitan University,
Japan) for their valuable suggestions on an earlier version of the manuscript. Lastly, the first author CML extends his thanks to Mr Chi-KeongChan, Mr Chi-Yui Lei, Mr Ken Kwan, Mr Eric
Kwan, Mr Tim Chan, Mr Martinho G. Oxalá, myhigh school classmates and teachers (Kwong Tai
Middle School, Macau), friends and family forproviding support and assistance during surveys
in Macau.
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ASIAN MYRMECOLOGY
A Journal of the International Network for the Study of Asian Ants
Communicating Editor: Bakhtiar Effendi Yahya

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Chi-Man Leong, Shiuh-Feng Shiao & Benoit Guénard

APPENDIX
Appendix 1. Summary of the accumulation of species known from Macau over time with details on
the number of reported native, tramp and exotic species.
Species
First record

Subspecies

Total

0

1

4

5

4

5

32

19

5

4

5

33

Wu 1941

19

5

4

6

34

Chapman and Capco 1951

19

5

5

6

35

Tang et al. 1995

21

6

5

6

38

Zhou 2001

23

6

5

6

40

Xu 2003

24

6

5

6

41

Hua 2006

27

8

7

7

49

Eguchi 2008

30

9

8

7

54

Present study 2017

79

15

11

8

113

NativespeciesTramp
species

Exoticspecies

Wheeler 1921

2

1

Wheeler 1928

18

Wheeler 1930